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HARVARD    UNIVERSITY 

Library  of  the 

Museum  of 

Comparative  Zoology 


The  Great  Basin  Naturalist 


VOLUME  35,  1975 


Editor:  STi:i'Hr.N  L.  W( 


Published  at  Brigham  Young  University,  by 
Brigham  Young  University 


TABLE  OF  CONTENl  S 

Volume  36 

Number  1  -  March  31.  1975 

Evolution   of  the   sceloporine   lizards    (Iguaniclae).    Kenneth   R.    Larsen 

and    Wilmer    W.    Tanner    1 

New   synonymy   and   new   species   of   American   bark    beetles    (Coleop- 

tera:   Scolytidae) .  Stephen  L.  Wood  21 

Genetics,   environment,   and  subspecies   differences:    the  case  of  Polites 

sabuleti   (Lepidoptera:    Hesperiidae).  Arthur  M.   Shapiro  33 

Life  history  and  ecology  of  Megarcys  signata   (Plecoptera:   Perlodidae), 

Mill  Creek,  Wasatch  Mountains,  Utah.  Mary  R.  Gather  and  Arden 

R.    Gaufin    39 

Records  of  stoneflies    (Plecoptera)    from  Nevada.  Marv  R.   Gather,  Bill 

P.  Stark,  Arden  R.  Gaufin ' 49 

Growth  of  Plecoptera    (stonefly)   nymphs  at  constant,  abnormally  high 

temperatures.  Joseph  M.  Branham,  Arden  R.   Gaufin,  and  Robbin 

L.    Traver    51 

Water  balance  and  fluid  consumption  in  the  southern  grasshopper  m.ouse, 

Onychomys  torridus.  Vernon  C.  Bleich  and  Orlando  A.  Schwartz    ...       62 
A  systematic  study  of  Coenia  and  Paracoenia    (Diptera:    Ephvdridae). 

Wayne    N.    Mathis 1 65 

Environmental  factors  in  relation  to  the  salt  content  of  Salicornio  pa- 

cifica  var.  utahensis.  D.  J.  Hansen  and  D.  J.  Weber 86 

New  records  of  stoneflies  (Plecoptera)  from  New  Mexico.  Bill  P.  Stark, 

Theodore  A.  Wolff,  and  Arden  R.  Gaufin 97 

The  authorship  and  date  of  publication  of  Siren  intermedia  (Amphibia: 

Caudata).    Hobart   M.    Smith,    Rozella    B.    Smith,    and    H.    Lewds 

Sawin ..-. 100 

New   mites   from   the   Yampa   Valley    (Acarina:    Cryptostigmata:    Ori- 

batulidae,     Passalizetidae).     Harold     G.     Higgins     and     Tvler     A. 

Woolley ■ 103 

The  identity  of  Boucourt's  lizard  Eunieces  capito  1879.  Hobart  M.  Smith, 

Rozella  B.  Smith,  and  Jean  Guibe 109 

Studies    in   nearctic    desert   sand    dune   Orthoptera.    Part   XV.    Eremog- 

raphy  of  Spaniacris  with  biological  notes.  Ernest  R.  Tinkham  113 

Roosting  behavior  of  male  Euderma   maculatum   from   Utah.   Richard 

M.  Poche  and  George  A.  Ruffner  121 

The  nest  and  larva  of  Diploplectron  hrunneipes   (Cresson)    Hymenop- 

tera:    Sphecidae).  Howard  E.  Evans  123 

Number  2  -  .Tune  30,  1975 

A  revision   of   the   Phacelia   Crenulatae   group    (Hydrophyllaceae)    for 

North  America.   N.   Duane  Atwood  127 

Rodent  populations,  biomass,  and  community  relationships  in  Arte- 
misia tridentata.  Rush  Vallev,  Utah.  D.  W.  Nichols,  H.  D.  Smith, 
and    M.    F.    Baker    ." 191 

Computerized  reduction  of  meteorologic  measurements  from  irrigated 
and  nonirrigated  plots  in  central  Utah.  Ferron  L.  Andersen  and 
Paul   R.   Roper   - 203 


3-^   ' 


Clarence    Cottani,    1899-1974;    a    distinguished    alumnus    of    Brigham 

Young  University.   Vasco  M.   Tanner   231 

Evolutionary  divergence  in  closely  related  populations  of  Mimulus 
guttatus  (Scrophulariaceae).  Karen  W.  Hughes  and  Robert  W. 
Vickery,  Jr 240 

Number  3  -  September  30,  1975 

Urosaurus  and  its  phylogenetic  relationship  to  Uta  as  determined  by 
osteolog}'  and  myology  (Reptilia:  Iguanidae).  Charles  Fanghella, 
David  F.  Avery,  and  Wilmer  W.  Tanner  245 

Distribution  and  adundance  of  the  black-billed  magpie    {Pica  pica)    in 

North  America.  Carl  E.  Bock  and  Larry  W.  Lepthien  269 

Nectar    composition    of   hawkmoth-visited    species    of   Oenothera    (Ona- 

graceae).  Robert  E.  Stockhouse,  II 273 

A  revision  of  the  nearctic  species  of  Clinohelea  Kieffer  (Diptera:  Cera- 

topogonidae).  William  L.  Grogan,  Jr.,  and  Willis  W.  Wirth 275 

Basidiomycetes   that   decay   junipers   in   Arizona.   R.   L.    Gilbertson   and 

J.    P.    Lindsey 288 

Body    size,   organ   size,   and   sex   ratios    in   adult   and   yearling   Belding 

ground  squirrels.  Martin  L.  Morton  and  Robert  J.  Parmer  305 

Photoperiodic  responses  of  phenologically  aberrant  populations  of  pie- 
rid  butterflies    (Lepidoptera).  Arthur  M.   Shapiro  310 

Additional  records  of  reptiles  from  Jalisco,  Mexico.  Philip  A.  Medica, 

Rudolf  G.  Arndt,  and  James  R.  Dixon 317 

Invasion  of  big  sagebrush  {Artemisia  tridentata)  by  white  fir  {Abies 
concolor)  on  the  southeastern  slopes  of  the  Warner  Mountains, 
California.  Thomas  R.  Vale 319 

Morpholog}'  of  ephemeral  and  persistent  leaves  of  three  subspecies  of 
big  sagebrush  grown  in  a  uniform  environment.  W.  T.  McDonough, 
R.  0.  Harniss,  and  R.  B.  Campbell 325 

Number  4  -  December  31,  1975 

Endangered,  threatened,  extinct,  endemic,  and  rare  or  restricted  Utah 
vascular  plants.  Stanley  L.  Welsh,  N.  Duane  Atwood,  and  James 
L.  Reveal  ". 327 

Utah  plant  novelties  in  Cymopterus  and  Penstemon.  Stanley  L.  Welsh....     377 

The  Zygoptera    (Odonata)    of  Utah  with  notes  on  their  biology.  A.   B. 

Provonsha :. 379 

New   synonymy   and   new   species   of   American   bark    beetles    (Coleop- 

tera:   Scolytidae),  Part  II.  Stephen  L.  Wood  391 

Correlates  of  burrow  location  in  Beechey  ground  squirrels.  Donald  H. 

Owings    and    Mark    Borchert 402 

Arachnids  as  ecological  indicators.  Dorald  M.  Allred  405 

Notes  on  the  genus  Bomhylius  Linnaeus  in  Utah,  with  key  and  descrip- 
tions of  new  species  (Diptera:  Bombyliidae).  D.  Elmer  Johnson 
and    Lucile   Maughan   Johnson 407 

Breeding  range  expansion  of  the  starling  in  Utah.  Dwight  G.  Smith  ....     419 

Some  parasites  of  paddlefish   {Polydon  spathula)   from  the  Yellowstone 

River,  Montana.  Lawrence  L.  Lockard  and  R.  Randall  Parsons  ....     425 

Reproductive  cycle  of  the  Belding  ground  squirrel  [Sperrnnphilus  hel- 
dingi) :  seasonal  and  age  differences.  Martin  L.  Morton  and 
John  S.   Gallup 427 


A  new  combination  in  Penstemon  (Scrophulariaceae).  Stephen  L.  Clark..  434 
Some  relationships  between  water  fertility  and  egg  production  in  brown 

trout  (Salmo  trutta)  from  Montana  streams.  Lawrence  L.  Lockard..  435 
Some   relationships   between   internal   parasites   and   brown   trout   from 

Montana  streams.  Lawrence  L.  Lockard,  R.  Randall  Parsons,  and 

Barry  M.  Schaplow 442 

Sexual    dimorphism   in   malpighian   tubules   of   Pteronarcys   californica 

Newport   (Plecoptera) .  Ralph  R.  Hathaway 449 

New    records    of    the    bat    Plecotus    phyllotis    from    Utah.     Richard 

M.  Poche  451 


€  GREAT  BASIN  NATURALISl 


ime35Na1         March  31, 1975 


Brigham  Young  Universit: 


MUQ.  COMP.  ZOOU 
L.JBRARY 

JUN   1  1  1975 

HARVARD 
UNIVERSITY 


i  ^\ 


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,  Zoology;  Stanley  L. 

Welsh,  Botany;  Clayton  M.  White,  Zoology. 
Ex  Officio  Editorial  Board  Members.  A.  Lester  Allen,  dean.  College  of  Biological  and 

Agricultural  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  continuously  published  from  one  to  four  times  a  year  since  then  by  Brigham 
Young  University,  Provo,  Utah.  In  general,  only  original,  previously  unpublished 
manuscripts  pertaining  to  the  biological  natural  history  of  the  Great  Basin  and  western 
North  America  will  be  accepted.   Manuscripts  are  subject  to  the  approval  of  the  editor. 

Subscriptions.  The  annual  subscription  is  $9  (outside  the  United  States  $10).  The 
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for  sale.  All  matters  pertaining  to  the  purchase  of  subscriptions  and  back  numbers 
should  be  directed  to  Brigham  Young  University  Press,  Marketing  Department,  204 
UPB,  Provo,  Utah  84602. 

Scholarly  Exchanges.  Libraries  or  other  organizations  interested  in  obtaining  this 
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  the  Great  Basin  Naturalist 
should  be  addressed  to  the  editor  as  instructed  on  the  back  cover. 


The  Great  Basin  Naturalist 

Published  at  Provo,  Utah,  by 
Brigham  Young  University 


Volume  35 


March  31,  1975 


No.  1 


EVOLUTION  OF  THE  SCELOPORINE  LIZARDS  (IGUANIDAE) 

Kenneth  R.   Larsen^   and  Wilmer  W.  Tanner^ 


Abstract. —  Phylogenetic  relationships  among  Sceloporine  genera  are  briefly  discussed.  Species  re- 
lationships witliin  the  genus  Sceloporus  are  analyzed,  and  evolutionary  lines  of  descent  are  proposed. 

The  genus  Sceloporus  is  composed  of  three  monophyletic  groups:  Group  I,  the  most  primitive,  prob- 
ably developed  from  Salor-\\ke  ancestral  stock  in  Miocene  times.  This  group  speciated  from  stock 
similar  to  Sceloporus  gadoviae  in  southern  Mexico  to  S.  merriami  in  the  North  and  contains  7  species 
in  3  species  groups.  We  propose  that  these  species  be  included  in  the  genus  Lysoptychus  Cope.  Group 
II  arose  from  Group  I  and  evolved  from  centrally  located  Sceloporus  pictus  in  all  directions  throughout 
Mexico.  This  intennediate  group  contains  approximately  19  species  in  5  species  groups.  Group  III 
also  arose  from  the  primitive  stock  of  Group  I  and  radiated  from  several  desert  refugia  created  by 
Pleistocene  glaciation.  Evolution  of  this  group  in  Mexico  was  generally  from  north  to  south  with 
Sceloporus  malachiticus  extending  as  far  south  as  Panama.  This  group  contains  approximately  33  spe- 
cies in  5  species  groups. 


In  a  previous  paper  (Larsen  and  Tan- 
ner, 1974)  we  presented  our  analysis  of 
the  species  in  the  lizard  genus  Sceloporus. 
Numerical  statistical  methods  were  used 
to  analyze  the  species  in  the  genus  Scel- 
oporus using  cranial  osteology,  external 
meristic  and  numeric  characters,  karyol- 
ogy,  display  behavior,  and  geographic  dis- 
tribution. A  new  classification  for  the 
genus  was  proposed  with  three  major 
branches  or  groups.  Group  I  contained  7 
species  in  3  species  groups.  Group  II  con- 
tained approximately  19  species  in  5  spe- 
cies groups.  Group  III  contained  approxi- 
mately 33  species  in  5  species  groups.  This 
classification  was  supported  by  the  cluster 
analysis  of  several  different  sets  of  data. 
Cranial  osteology,  zoogeograph}',  behavior, 
and  karyology  were  shown  to  be  taxon- 
omically  significant  as  numeric  charac- 
ters. Stepwise  discriminate  analysis 
showed  that  this  classification  of  the  spe- 
cies of  Sceloporus  into  3  major  groups  and 
13  species  groups  was  significant  at  the 
.999  confidence  level. 

The  purpose  of  this  paper  is  to  present 
our  views  on  the  evolution  of  the  species 
in  the  genus  Sceloporus.  We  also  propose 
a   ph3dogeny  of   closely  related    (Scelop- 


orine) genera.  We  are  grateful  for  the 
assistance  of  H.  M.  Smith,  C.  C.  Carpen- 
ter, W.  P.  Hall,  and  the  following  per- 
sons at  Brigham  Young  University:  A.  L. 
Allen,  F.  L.  Anderson,  J.  R.  Murphy,  M. 
S.  Peterson,  J.  K.  Rigby,  N.  M.  Smith,  D. 
A.  White,  and  S.  L.  Wood. 

Intergeneric  Phylogeny 

In  1828  Weigmann  described  several 
genera,  including  Sceloporus  (S.  torqua- 
tus) .  He  distinguished  Sceloporus  from  the 
South  American  Tropidurus  mainly  on 
the  basis  of  femoral  pores  (S'c^j/o^ thigh, 
porus=\)OYe) .  In  1852  Baird  and  Girard 
described  the  genus  Uta  (U.  stansburiana) 
which  is  distinguished  from  the  smaller 
species  of  Sceloporus  by  its  gular  fold  and 
granular  dorsal  scales.  In  1854  Hallowell 
erected  the  genus  Urosaurus  (U.  gracio- 
sus),  which  is  similar  to  Uta  but  has  sev- 
eral rows  of  enlarged,  carinate,  imbricate 
vertebrals  or  paravertebrals.  Two  years 
later  Dimieril  (1856)  described  the  genus 
Phymatolepis  (Urosaurus  bicarinatus)  on 
the  basis  of  enlarged  paravertebrals.  In 
1859  Baird  placed  Hallowell's  genus  Uro- 
saurus  in    synonymy   with    Uta,   and    in 


V07  North  500  West,  Provo,  Utah  84601 . 

-Department    of    Zoology,    Brigham    Young    Universitj-.    Provo.    Utah   84602. 


GREAT  BASIN  NATURALIST 


Vol.  35,  No.  1 


1864  Cope  did  the  same  with  Dmneril's 
Phymatolcpis.  Boulenger  (1885)  raised 
Cope's  Uta  thalassina  to  generic  status 
(Petrosauriis) ,  but  Cope  (1900)  rejected 
this  proposal  and  made  Petrosaurus  a  third 
synon;y^n  of  Uta.  In  1888  Cope  erected 
the  genus  LrsoptycJius  (L.  Iateralus^= 
Sceloporus  couchi)  on  the  basis  of  a  single 
specimen  that  appeared  to  have  a  well- 
developed  gular  fold.  Subsequent  investi- 
gation (Stejneger,  1904)  showed  the  "gu- 
lar fold"  to  be  an  artifact  of  preparation 
on  a  single  specimen  which  "was  pre- 
served in  such  a  manner  as  to  make  a  fold 
across  the  neck,  which  formed  the  basis 
for  the  erection  of  the  genus"  (Smith, 
1939,  p.  242).  Dickerson  (1919)  de- 
scribed the  genus  Sator  (S.  grandaevus) 
which  has  persisted  despite  Sator's  close 
similarity  to  Uta,  Urosaurus  and  Scel- 
oporus.  In  1942  Mittleman  resurrected 
the  genera  Urosaurus  and  Petrosaurus.  He 
also  erected  the  genus  Streptosaurus  based 
on  Uta  mearnsi,  which  is  most  similar  to 
Petrosawus.  He  proposed  that  Uta,  Uro- 
saurus, and  Sator  all  arose  independently 
from  Sceloporus.  He  placed  PJirynosoma 
with  the  above  genera  in  a  distinct  group. 
Smith  (1946)  moved  Sauromalus  and 
Dipsosaurus  to  more  primitive  positions 
but  otherwise  retained  Mittleman's  ar- 
rangement. Savage  (1958)  placed  Strept- 
osaurus in  synonymy  with  Petrosaurus. 
He  separated  Uta  from  Urosaurus  mainly 
on  the  basis  of  sternal  and  costal  mor- 
phology. He  placed  Uta  and  Petrosaurus 
with  the  sand  lizards  (Holhrookia,  Unia, 
and  Callisaurus) ,  leaving  Sceloporus,  Sa- 


SCEIOPORUS 

GROUP  III 

SCELOPORUS 

\                GROUP  II 

COPHOSAURUS 
HOIBROOKIA 

\/ 

\     /►'^ElOPORUS 
/            GROUP  1 

Y 

SATOR 
UMA      \ 

~\ 

/^^CALIISAURUS 

1 

^/ 

/Urosaurus 

PHRYNOSOMA^"-\^                                   "'*    \ 

/ 

^x 

petrosaurus 

ANCESTRAL          STOCK 

Fig.   1.     Phylogeny  of  sceloporine  genera  and 
the  three  major  groups  in  Sceloporus. 


tor,  and  Urosaurus  together.  Etheridge 
(1964)  rejected  Savage's  wide  separation 
of  Uta  and  Urosaurus,  and  placed  Uta, 
Urosaurus,  Sator,  and  Sceloporus  on  one 
side  and  Uma,  Holbrookia,  and  Callisau- 
rus on  the  other.  Primitive  to  both  groups 
was  Petrosaurus.  A  sand  lizard  resur- 
rected by  Clarke  (1965)  was  Troschel's 
(1852)  genus  Cophosaurus  (C.  texanus, 
previously  Holbrookia  texana) . 

Presch  (1969)  rejected  Etheridge's  re- 
moval of  PJirynosoma  from  the  scelopor- 
ines  and  placed  Phrynosoma  with  the 
sand  lizards  as  a  primitive  member  of  that 
group.  On  the  basis  of  scleral  ossicles, 
Presch  (1970)  indicated  that  Petrosaurus 
is  a  primitive  member  of  the  Sceloporus 
branch.  Ballinger  and  Tinkle  (1972)  pro- 
posed an  early  separation  of  the  Uta  and 
Petrosaurus  stock  from  the  ancestor  of 
Urosaurus,  Sator,  and  Sceloporus. 

Several  characters  suggest  further  modi- 
fication of  the  above  arrangement.  Our 
jiroposed  phylogeny  of  sceloporine  genera 
is  illustrated  in  Figure  1.  Urosaurus 
shows  a  tendency  for  enlarged  scales  near 
the  midline  of  the  dorsum.  This  trend  is 
further  developed  in  Sator,  which  has  en- 
larged dorsals  and  granular  laterals.  The 
migration  of  enlarged  scales  around  the 
sides  of  the  body  and  the  increase  in  scale 
size  and  degree  of  imbrication,  mucrona- 
tion,  and  carination  is  a  general  trend 
along  the  chain  of  genera  from  Petrosau- 
rus to  Sceloporus.  The  new  phylogeny  is 
also  supported  by  the  gradual  decrease  in 
development  of  the  gular  fold,  which  is 
completely  lost  in  all  species  of  Sceloporus 
in  Group  III.  Most  of  the  species  in 
Group  I  have  what  Smith  (1939)  called 
a  rudimentary  gular  fold.  Some  of  the 
species  in  Group  II  show  a  less  pro- 
nounced tendency  to  develop  a  gular 
fold,  and  Group  III  lacks  it  completely. 
The  gradual  loss  of  the  gular  fold  in  the 
Sceloporus  complex  is  more  probable  than 
a  loss  (from  Petrosaurus  to  Sceloporus) 
and  subsequent  redevelopment  (from 
Sceloporus  to  Uta,  Urosaurus,  or  Sator). 
This  reversal  of  the  phylogeny  resolves  a 
question  raised  by  Smith  (1946:178):  "It 
is  a  curious  fact  that  all  genera  that  have 
sprung  from  Sceloporus  have  developed 
a  gular  fold — including  Sator,  a  Baja  Cali- 
fornia genus.  The  tendency  to  develop 
this  fold  a})pears  to  be  restricted  to  the 
])rimitive  groups  of  Sceloporus  .  .  .  and 
these    are    the    groups    from    which    Uta, 


March  1975 


LARSEN,  TANNER:    SCELOPORINE  LIZARDS 


Urosaurus,  and  Sator  independently  ap- 
pear to  have  been  derived." 

Although  Smith  pointed  to  this  prob- 
lem, he  nevertheless  accepted  Mittleman's 
arrangement  of  the  sceloporine  genera. 
More  recently,  Smith  (per  comm.):  has 
agreed  that  Sceloporus  ma}'  be  derived 
with  respect  to  Uta^  Urosaurus.  and  Sator. 
This  position  has  also  been  suggested  by 
Hall  (pers.  comm.):  "Inspection  of  the 
structure  of  the  femoral  pores  and  their 
surrounding  scales,  and  the  development 
of  mucronation  and  carination  of  the  body 
scales,  to  mention  but  two  sets  of  charac- 
ters in  various  primitive  Sceloporus  and 
in  other  sceloporine  genera,  will  suggest 
that  Sceloporus  is  derived  even  in  respect 
to  Uta  and  Urosaurus. '' 

We  suggest  the  following  conclusions 
with  regard  to  the  new  phylogeny  and 
published  data  on  hip  ratios  of  displaying 
males  (Purdue  and  Carpenter,  1972a, 
1972b).  The  hip  ratio  (vertical  hip  move- 
ment to  vertical  eye  movement)  increased 
from  Petrosaurus  (0.68)  to  Uta  (average 
0.74)  to  Urosaurus  (average  1.06).  After 
the  transition  from  Sator  (no  published 
data  on  hip  ratios)  to  Sceloporus,  the 
trend  reversed  and  hip  ratios  decreased 
from  an  average  of  1.21  in  Group  I  to 
0.66  in  Group  II  to  0.34  in  Group  III 
(averages  computed  from  Purdue  and 
Carpenter,  1972b). 

Etheridge  (1964)  illustrated  clavicles 
and  scapulocoracoids  of  8  sceloporine 
genera  (excluding  Phrynosoma) .  If  his 
drawings  are  superimposed  on  the  new 
phylogeny  (Fig.  2),  two  trends  are  ap- 
parent: (1)  a  gradual  development  of 
the  scapular  fenestra  (top  groove)  from 
Petrosaurus  to  Sceloporus  Group  III,  and 
(2)  an  increase  in  size  of  the  clavicular 
hook.  If  Urosaurus  and  Uta  were  derived 
from  Sceloporus,  the  scapular  fenestra 
would  have  developed  and  then  disap- 
peared from  Petrosaurus  to  Sceloporus  to 
Uta.  This  improbable  reversal  is  similar 
to  the  problem  with  the  gular  fold.  We 
are  persuaded  that  the  new  phylogeny  is 
more  probable. 

Intrageneric  Phylogeny 

The  first  ph^dogenetic  schemes  for  the 
genus  Sceloporus  were  proposed  by  Smith 
(1934,  1937a,  1937b,  1938,  1939).  Other 
workers  have  recently  modified  the  phy- 
logeny on  the  basis  of  karyology    (Cole, 


1970,  1971a,  1971b;  Hall,  1971,  1973), 
and  behavior  (Bussjaeger,  1971). 

Larsen  and  Tanner  (1974)  redefined 
relationships  among  the  species  in  the 
genus  Sceloporus.  We  used  Ward's  clus- 
ter analysis  (Wishart,  1968)  to  cluster  55 
species  on  the  basis  of  external  characters, 
cranial  osteology,  karyology,  behavior, 
and  zoogeography  (Fig.  3).  We  then  used 
step- wise  discriminate  analysis  (Dixon 
1967)  and  found  that  the  arrangement 
of  groups  and  subgroups  is  significant  at 
the   .999   level   of  confidence    (Table    1). 

Although  Ward's  cluster  analysis  pro- 
vides a  phenetic  dendogram,  it  does  not 
give  any  indication  as  to  which  branch  of 
a  cluster  is  derived  and  which  is  primi- 
tive. In  1939  Smith  said,  "The  most 
primitive  form  of  this  group  is  undoubted- 
ly lunaei  which  is  closely  related  to  for- 
mosus  malachiticus''  (p.  60).  In  other 
words,  lunaei  is  the  most  primitive  form 


PETROSAURUS 


Fig.  2.  Clavicles  and  scapulocoracoids  of  sev- 
eral sceloporines.  All  illustrations  except  Scelo- 
porus I,  Sceloporus  II,  and  Sceloporus  III  are 
from  Etheridge  (1964). 


4 

GREAT  BASIN  NATURALIST 

Vol.  35,  No.  1 

Tabt.k  1. — Groups 

and  subgroups  in  the  genus  Sceloporus 

Group  I  (7  spp 

) 

Group  II  (20  spp.) 

Group  III  (33  spp.) 

Subgroup  A  ( 1 

spp.) 

Subgroup  A  (7  spp.) 

Subgroup  A  (9  spp.) 

gadoviae 

grammicus 

spinosus 

Subgroup  B  (2 

spp.) 

pictus 

orcutti 

couchi 

megalepidurus 

clarki 

memami 

cryptus 

melanorhinus 

Subgroup  C  (4 

spp.) 

shannonorum  * 

magister 

maculosus 

heterolepis 

olivaceus 

parvus 

asper 

cautus 

jalapae 

Subgroup  B  (2  spp.) 

horridus 

ochotei-enae 

pyrocephalus 

edwardtaylori 

nelsoni 

Subgroup  B  (7  spp.) 

Subgroup  C  (3  spp.) 

formosus 

scalaris 

lunaei 

goldmani* 

nialachiticus 

aeneus 

acanthinus 

Subgroup  D  (4  spp.) 

Subgroup  C  (5  spp.) 

siniferus 

undulatus 

carinatus 

virgatus 

utifonnis 

woodi 

squamosus 

occidentalis 

Subgroup  E  (4  spp.) 

graciosus 

variabilis 

Subgroup  D  (4  spp.) 

cozumelae 

jarrovi 

teapensis 

lineolateralis 

chrysostictus 

ornatus 
dugesi 
Subgroup  E  (8  spp.) 
torquatus 
cyanogenys 
bulleri 
insignis* 
macdougalli 
mucronatus 
serrifer 
poinsetti 

*  Species  not  examined  in  this  study. 


ill  the  spinosus  s])ecies  group  because  it  is 
most  similar  to  a  member  of  the  next 
closest  group  (formosus).  This  statement 
by  Smith  is  consistent  with  the  following 
method  of  converting  a  phenetic  dendro- 
gram into  a  phylogeny  (Fig.  4):  If  "A" 
is  primitive  to  "B"  it  is  less  derived  from 
(more  similar  to)  the  stem  species  "G." 
The  more  primitive  member  of  the  other 
cluster  ("C"  or  "D")  will  also  be  more 
similar  to  "G."  The  more  primitive  mem- 
bers of  the  two  clusters  will  therefore  be 
phylogenetically  "closer"  and  phenotypi- 
cally  more  similar  than  any  other  com- 
bination from  the  two  clusters.  This  rule 
can  be  applied  objectively  with  a  similar- 
ity matrix. 

When  all  possible  pairs  between  adja- 
cent clusters  are  compared,  the  two  most 
similar  species  are  considered  jjrimitive 
within  their  res|)ective  clusters.  This 
technique  will  convert  a  dendrogram  into 
a   phylogeny. 

Ward's  cluster  analysis  and  the  above 
phylogeny  techni(|ii(>  were  repeated  sever- 


al times  using  external  and  osteological 
characters,  distribution,  karyology,  be- 
havior, and  combinations  of  the  above. 
(See  Larsen  and  Tanner,  1974,  for  a  pre- 
sentation of  results.)  The  differences 
among  results  were  resolved  subjectively 
to  produce  a  composite  phylogeny  (Fig. 
5).  This  ])rocedure  is  based  on  several 
assumptions  which  are  admittedly  vul- 
nerable. To  restrict  the  scope  of  our  study 
it  was  assumed  that  the  alpha  taxonomy 
is  complete  and  correct.  That  is,  it  was 
assvimed  that  all  species  of  Sceloporus  are 
now  named  and  correctly  defined  in  the 
literature.  Of  course,  this  assumption  may 
be  incorrect.  But  the  purpose  of  our  study 
is  to  produce  a  general  overview  and  not 
a  detailed  taxonomic  review.  The  de- 
tails near  the  ends  of  branches  are  there- 
fore tonlativo  and  stibject  to  future  re- 
view. 

In  spite  of  the  large  number  of  charac- 
ters considered  (over  80),  these  results 
are  also  subject  to  errors  due  to  parallel- 
ism, convergence,  varying  rates  of  diver- 


March  1975 


LARSEN,  TANNER:    SCELOPORINE  LIZARDS 


Formosus  __^ 
spinosus  .^_^ 
Horndus  _— 

Olivaceus 

Cqutus   

Adieri  

Molachiticus  — 
Luna«i 

Lundalli  ^^— 
Acantninus  ~__ 
Edwprdtoylori. 

Orcutti    —^— 

Magistcr  

Undulatus   

Occidentalis  ^ 
Virgatus '— ^— 
Graciosus  ^^— i 
Torquatus 
sernfer 

Mucronatus  

Cyonogenyi 

Bulleri 
Poinsetti 
jarrovi    -^—— 
Linaolateralis- 
Ornotus 
Dugesi   — — ^— 

Atper   

Heterolepis_ 
Grammicus 


Megolopidurus. 
Pictus  ^^-^^^ 


Ochoterenae  — 

Jalopae 

scolons 

Aeneus   

pyrocephalus- 
Nelsoni     — - 
Melonorhinus. 
siniferus 
Connotui 

Utiformis 

Variabilis 

Cozumela* 

Teapensif  '^^— 
Chrysostictus- 
squamosus^^ 
Parvus  ^^.— 
Maculosus^— 
Couchi  ■ 
MerriamI— 
Cadoviae  


^^ 


^ 


^ 


^ 


i 


;=^ 


0.5 


16 


Fig.  3.     Dendrogram   generated   by   Ward's   cluster    analysis    of    external, 
diaracters  (82  characters). 


skull,    and    distribution 


gence,  pleiotrophy,  and  other  cases  in 
which  the  phenotype  is  not  a  direct  mani- 
festation of  the  genotype.  All  phylogene- 
tic  conclusions  are  subject  to  these  liinita- 
tions,  and  the  systematist  can  do  little 
more  than  acknowledge  the  circumstantial 
nature  of  his  evidence. 

We  propose  that  SceJoporus  is  derived 
from  Uta  through  Urosaurus  and  Sator 
(see  above).  Smith  (1938)  suggested  that 
tlie  connection  between  these  genera  is 
from  Urosaurus  ornatus  to  Sceloporus 
couchi.  Smith  included  couchi  in  the 
variabilis  species  group. 

Figure  6  shows  the  arrangement  of 
species  in  Smith's  variabilis,  maculosus, 
and  mcrrianii  groups  according  to  Smith 
(1939,  Fig.  42)  and  the  new  phylogeny. 
Four  of  these  species  {couchi,  parvus, 
maculosus,  and  merriami)  are  transferred 
to  Group  I.  Smith  may  have  allowed  for 
this  by  placing  these  four  species  on  one 
side  of  his  tree  next  to  Uta.    If  Uta  {Uta, 


Urosaurus,  and  Sator)  is  considered  primi- 
ti^'e  to  Sceloporus,  then  Smith's  evidence 
supports  our  conclusion  that  Group  I  is 
primitive  to  the  other  two  groups  in 
Sceloporus.  The  remaining  species  in 
Smith's  variabilis  group  {variabilis,  coz- 
umelae,  and  teapensis)  are  placed  in 
Group   II. 

Smith  (1939:239)  allowed  for  the  re- 
moval of  parvus  and  couchi  from  the  var- 
iabilis grouj)  with  this  statement: 

That  parvus  and  couchi  are  only  dis- 
tantly related  to  the  remainder  of  the  group 
is  shown  by  the  widely  different  charac- 
ter of  the  ventral  coloration  in  the  males, 
smooth  head  scales,  larger  number  of  fem- 
oral pores,  and  general  habitus.  ...  It 
is  my  belief  that  this  section  approaches 
more  closely  the  ancestral  stock  of  Uta  than 
the   other   species   of   the    variabilis   group. 

Smith  (p.  239)  also  associated  merriami 
with  Uta:  "It  w^ould  appear  that  merri- 
ami is  closely  related  to  Uta.  and  that  Uta 


GREAT  BASIN  NATURALIST 


Vol.  35,  No.  1 


B 


E 

1 

1 

F 

1 

1 
G 

IF     A-C  =  l 

A-D=2 

B-C  =  2 

B-D 

=  3 

THEN-- 

Fig.  4.  Phylogeny  theory-  K  the  phenetic 
distance  between  "A"  and  "C"  is  less  than  that 
between  any  other  pair,  then  "A"  and  "C"  are 
primitive  members  in  clusters  "E"   and  "F." 

arose  from  the  forms  now  extinct  which 
closed  the  present  gap  between  couchi 
and  merriami.'"  Note  that  our  new  ar- 
rangement places  merriami  and  couchi 
together. 

Another  divergence  from  Smith's  phylo- 
genetic  tree  is  the  addition  of  chrysostic- 
tus  to  the  variabilis  group.  Smith  (p.  239) 
supports  this  inclusion  (and  the  close 
proximity  of  the  siniferus  group) :  "An- 
other group  close!)'  related  to  the  variab- 
ilis section  is  the  siniferus  series,  which 
closely  approaches  the  variabilis  group 
through  cuprous.  .  .  .  The  chrysostictus 
group  is  also  closely  related." 

Thus  it  can  be  seen  that  Smith  allowed 
for  the  possibility  of  removing  parvus  and 
couchi  and  adding  chrysostictus,  which 
changes  his  variabilis  group  into  the  new 
variabilis  group. 


Smith  stated  that  the  siniferus  group 
"closely  approaches  the  variabilis  group" 
and  yet  his  illustration  (1939,  Fig.  3)  has 
these  groups  separated  by  several  other 
groups.  In  the  new  phylogeny  they  are 
adjacent. 

Figure  7  compares  Smith's  arrange- 
ment of  his  chrysostictus.  utiformis  and 
siniferus  groups  with  the  new  arrange- 
ment of  the  same  species.  Besides  the 
placing  of  chrysostictus  in  the  variabilis 
group  (which  has  already  been  ex- 
plained), the  only  major  difference  in 
Figure  4  is  the  removal  of  ochoterenae 
to  place  it  in  Group  I.  (The  inclusion 
of  utiformis  in  the  siniferus  group  is  mi- 
nor) .  Smith  listed  1 1  diagnostic  characters 
of  the  siniferus  group.  In  three  cases  he 
said  "except  ochoterenae''  and  in  another 
"except  ochoterenae  and  cupreusT  He 
(p.  301 )  said,  "Postanals  tending  to  be 
poorly  developed  (except  ochoterenae  and 
cupreus);  two  postrostrals  (except  ocho- 
terenae, without  postrostrals) ;  .  .  .  ventral 
scales  pointed  or,  at  least  not  notched  (ex- 
cept ochoterenae  in  which  they  are 
notched)  .  .  .  males  without  distinctive 
ventral  coloration   (except  ochoterenae).'" 

If  size  is  discounted,  then  ochoterenae 
is  different  in  4  of  the  10  diagnostic  char- 
acters for  the  siniferus  group.  S.  ocho- 
terenae also  has  more  femoral  pores  than 
any  other  species  in  Smith's  siniferus 
group.  Smith's  conclusions,  therefore, 
would  not  be  seriously  challenged  if 
ochoterenae  were  removed  from  the  sinif- 
erus group  and  placed  in  Group  I  next  to 
jalapae.  In  fact,  when  describing  ocho- 
terenae., Smith  (p.  309)  said,  "three  or 
four  scales  on  anterior  border  of  ear,  not 
so  large  as  in  jalapae.''  So  apparently 
he  was  comparing  these  two  species. 

Smith  included  jalapae  in  his  scalaris 
group,  which  is  otherwise  identical  to  the 
new  scalaris  group  (Fig.  5).  Removing 
jalapae  from  the  scalaris  group  to  place  it 
in  the  primitive  Group  I  is  supported  by 
the  following  statement  b^-  Smith  (p. 
331): 

The  only  species  doubtfully  inchuled  in 
this  group  is  jalapae,  which  differs  from 
the  remaining  fonns  in  having  lateral  scales 
in  distinctly  oblique  rows,  and  in  lacking 
postrostrals  [as  does  ochoterenae].  .  .  . 

5.  jalapae  is  clearly,  the  most  primitive 
member  of  the  group.  S.  scalaris,  aeneus 
and  goldmani  are  clearly  more  closely  re- 
lated to  each  other  than  any  one  of  these 
is  to  jalapae. 


March  1975 


LARSEN,  TANNER:    SCELOPORINE  LIZARDS 

HORRIDUS 


MELANORHINUS 


MALACHITICUS 

ACANTHINUS 


OCCIDENTALIS 


SCALARIS  I    UTirORMIS 

GOLDMANI 


VARIABILIS 

TEAPENSIS 


Fig.  5.     Proposed  phylogeny  for  the  genus  Sceloporus.     (*    =    species   not   examined.) 


GREAT  BASIN  NATURALIST 


Vol.  35,  No.  1 


PARVUS  MACULOSUS         MERRIAMI 


TEAPENSIS      COZUMELAE 


Fig.  6.     Phylogeny   of   Smith's    (1939)    variabilis,    maculosus.    and    merriami    groups   according   to 
Smith    (A)    and  the  new  phylogeny   (B). 


CHRYSOSTICTUS 


VARIABILIS  GROUP 


SCALARIS  GROUP 


OGHOTERENAE 


CHRYSOSTIGTUS 


VARIABILIS  GROUP 


SCALARIS  GROUP 


OGHOTERENAE 


Fig.  7.     Phylogeny   of   Smith's    (1939)    chrysostictus,   utiformis,   and  siniferus  groups   according   to 
Smith    (A)    and  the  new  phylogeny    (B). 


March  1975 


LARSEN.  TANNER:    SCELOPORINE  LIZARDS 


(iroup  I  includes:  parvus,  couchi,  ma- 
( ulosus,  mcrriami,  ochoterenat\  jalapae, 
aiul  gadoviac.  the  most  primitive.  Smith 
( p.  362)  inchided  gadoviac  with  nelsoni 
and  pyrocephalus  in  the  pyrocephalus 
group.  But  once  again  he  outhned  rea- 
sons why  gadoviac  could  be  removed  and 
[)laced  in  Group  I.  "5.  gadoviae  differs 
widely  from  other  members  of  the  group 
in  having  very  small  dorsal  scales,  a  large 
number  of  femoral  pores,  a  postfemoral 
dermal  pocket,  very  small  scales  on  pos- 
terior surface  of  the  thighs,  and  many 
other  minor  characters."  S.  gadoviae  is 
also  the  only  member  of  this  group  to 
have  a  vestigial  gular  fold  as  mentioned 
by  Smith  (p.  374):  "scales  immediately 
preceding  gidar  fold  region  somewhat  re- 
duced in  size."  All  of  these  characters 
are  diagnostic  of  Group  I,  and  this  primi- 
tive placement  is  therefore  natural.  In 
fact.  Smith  (p.  363)  said,  "I  assume 
gadoviae  to  be  nearest  the  primitive  type, 
as  it  retains  certain  characters  of  the 
variabilis  group,  from  which  I  believe  it 
was  derived." 

The  main  character  on  which  Smith  (p. 
363)  based  his  inclusion  of  gadoviae  with 
the  pyrocephalus  group  is  the  strong  com- 
pression of  the  tail:  "That  the  group  is  a 
natural  one  is  more  or  less  assured  by  its 
compact  range  and  by  the  common  char- 
acter of  the  compressed  tail,  which  is 
otherwise  unknown  in  the  genus."  In 
view  of  the  many  characters  supporting 
the  placement  of  gadoviac  in  Group  I,  we 
propose  that  a  compressed  tail  developed 
twice:  once  in  the  pyrocephalus  group, 
and  once  in  gadoviae.  Smith  (p.  363) 
gave  further  support  to  this  placement  of 
gadoviae:  "The  assumption  that  gadoviac 
is  a  remnant  of  a  primitive  stock  is  sup- 
ported by  its  secretive  habits  and  its  re- 
striction to  a  somewhat  arid  region." 

The  most  serious  difference  between 
the  new  phylogen^-  and  that  of  Smith  is 
the  placement  of  the  gramniicus  and  me- 
galepidurus  groups.  In  both  phylogenies 
the  species  are  arranged  in  a  similar  man- 
ner within  these  groups.  But  Smith 
placed  these  groups  next  to  the  jormosus 
group  with  the  large-scaled,  large-sized 
species,  and  we  ha\e  moved  them  to  a 
primitive  position  in  Group  II.  How- 
ever, we  propose  that  the  grammicus 
group  (we  have  combined  Smith's  gram- 
micus and  hetcrolcpis  groups)  is  the  most 
primitive    in    Group    II.     In    fact,    Smith 


(1938:552)  said  "the  microlepidurus  [our 
grammicus^  group  is  assumed  to  be  the 
most  primitive  of  these  [the  large-scaled, 
large-sized  sjiecies],  largely  because  of 
its  very  small  scales."  This  greater  separ- 
ation between  the  grammicus  and  jormos- 
us groups  is  further  justified  by  the  fact 
that  the  diploid  number  of  chromosomes 
is  22  (derived)  in  the  jormosus  group  and 
32  (primitive)  in  the  grammicus  group. 
We  propose,  therefore,  that  some  of,  the 
similarities  between  grammicus  and  jor- 
mosus (coloration,  dorsal-scale  count,  ovo- 
viviparity,  and  preference  for  an  arboreal 
habitat)  are  a  result  of  convergence  as 
is  true  of  gadoviae  and  the  pyrocephalus 
group. 

The  only  remaining  difference  from 
Smith's  jormosus  group  is  his  inclusion  of 
asper,  which  we  have  moved  to  the  gram- 
micus group.  This  move  is  justified  by 
the  fact  that  asper  has  32  chromosomes, 
as  do  the  other  members  of  the  grammicus 
group.  If  the  grammicus  grou]:)  is  re- 
moved from  Smith's  large-scaled,  large- 
sized  branch,  the  remaining  species  are 
the  same  as  those  included  in  Group  III. 
This  grouping  (the  omission  of  grammi- 
cus)  was  allowed  by  Smith   (1938:552): 

The  relatively  small  size  of  the  species  of 
the  undulatus  group  must  be  assumed  as 
a  parallel  development  rather  than  a  direct 
inlieritence  of  the  small  size  of  the  ancestor 
in  the  variabilis  group,  for  the  close  rela- 
tionship of  the  spinosus  and  undulatus 
groups  cannot  logically  be  disputed,  nor  is 
the  close  relationship  of  the  spinosus,  lor- 
qualus   and  formosus  groups   doubtful." 

Smith  and  Taylor  (1950)  included  the 
following  species  within  the  undulatus 
group:  undulatus,  cautus,  occidentalism 
and  woodi.  Since  then,  virgatus  has  been 
raised  from  subspecific  to  specific  status 
(Cole,  1963).  Smith  (1939)  placed  fjrac/- 
osus  adjacent  to  the  undulatus  group,  so 
the  only  discrepanc}'  between  the  two 
classifications  is  the  placement  of  cautus, 
which  we  have  moved  to  the  spinosus 
group  next  to  olivaceus.  This  mo^'ement 
is  justified  by  the  fact  that  there  is  a  zone 
of  intergradation  between  cautus  and  oli- 
vaceus  (Hall,  pers.  comm.). 

Bussjaeger  (1971:151)   remarked: 

The  relation  of  cautus  and  olivaceus 
and  the  undulatus  group  of  Sceloporus  has 
been  questioned.  Hall's  data  indicated  that 
these  two  species  were  the  same  and  limited 
data  on  their  displays  indicate  that  they  are 
similar.    If  one  accepts  that  they  are  syn- 


10 


GREAT  BASIN  NATURALIST 


Vol.  35,  No.  1 


onyms,  then  olivaceus  (cautus)  would  be 
the  connecting  link  between  the  spinosus 
and  undulatus  groups. 

However,  rather  than  use  these  forms  as 
a  link  between  species  groups,  we  have 
placed  them  together  in  the  spijiosus 
group. 

Smith  (1938:554)  indicated  that  the 
torquatus  group  consited  of  2  subgroups: 
"It  appears  that  soon  after  the  separation 
of  the  torquatus  stock  from  the  other 
groups  of  Sceloporus,  there  was  a  separa- 
tion into  two  divisions,  one  of  which  ex- 
hibited a  tendency  to  develop  small  scales, 
the  other  large  scales."  We  have  recog- 
nized the  small-scaled  division  as  the 
jarrovii  group. 

Figure  8  shows  the  phylogeny  of  the 
jarrovii  group  according  to  Smith  (1938, 
Fig.  4)  and  the  new  arrangement.  Al- 
though he  placed  lineolateralis  further 
away  from  jarrovii  in  his  diagram.  Smith 
(p.  556)  did  say,  "S".  jarrovii  appears  to 
be  most  closely  related  to  lineolateralis. 
From  this  species,  or  its  ancestors,  the  re- 
maining species  of  the  small-scaled  divi- 
sion have  obviously  been  derived." 

Figure  9  shows  the  phylogeny  of  the 
torquatus  group  according  to  Smith  (1938, 


LINEOLATERALIS 


Fig.  8.     Phylogeny  of  jarrovi  group  according 
to  Smith  (1938)   (A)  and  the  new  phylogeny  (B). 

Figs.  3-4)  and  the  new  arrangement. 
There  seems  to  be  little  similarity  here, 
except  that  torquatus  is  derived  from 
serrifer,  and  poinsetti  is  derived  from 
cyanogenys  in  both  trees.  Smith  (1938: 
555)  raised  a  question  about  the  ancestral 
position  of  serrifer: 

S.  serrifer  appears  to  be  the  oldest  of  the 
large-scaled  species.  The  postulation  that 
this  species,  which  is  one  of  the  larger  ones 


POINSETTI 


CYANOGENYS 


Fig.  9     Phylogeny  of  torquatus  group  according  to  Smith  (1938)   (A)  and  the  new  phylogeny  (B). 


March  1975 


LARSEN,  TANNER:    SCELOPORINE  LIZARDS 


11 


of  the  genus,  and  one  having  large  scales, 
is  nearest  to  the  ancestral  type  of  the  large- 
scaled  division  of  the  torquatus  group  may 
appear  to  be  contradictory  to  the  postula- 
tion  that  Sceloporus  is  derived  from  small 
species  with  small  scales.  However,  my  as- 
sumption seems  to  be  justified  by  the  fact 
that  serrifer  occupies  a  southern  position  on 
the  periphery  of  the  geographical  area  now 
occupied  by  the  torquatus  group. 

The  reason  for  this  paradox  is  that  Smith 
assumed  speciation  in  Group  III  was  from 
south  to  north.  The  data  in  1938  strongly 
supported  this  conclusion.  Obviously, 
Smith  did  not  believe  that  a  peripheral 
location  is  necessarily  primitive,  because 
on  the  next  page  (556)  he  said,  "S".  mu- 
cronatus  appears  to  be  the  nearest  to  the 
ancestral  type  of  these  three  species  {cy- 
anogenys,  poinsetti  and  omiltemanus)  de- 
spite the  fact  that  it  has  larger  scales  than 
they.  I  so  conclude  because  of  its  central- 
ized geographical  position  with  relation 
to  the  area  occupied  by  the  other  three 
forms." 

So  the  basic  problems  can  be  solved, 
and  the  trend  is  indeed  from  small  to 
large  size  and  small  to  large  scales  if  this 
group  was  developed  from  north  to  south 
rather  than  south  to  north.  Smith  indi- 
cated a  northward  development  from  ser- 
rifer to  torquatus  to  mucronatus  to  cyano- 


genys,  and  our  phylogeny  indicates  a 
southward  development  from  cyanogenys 
to  mucronatus  to  serrifer  to  torquatus.  An 
ancestral  placement  of  cyanogenys  is  fur- 
ther supported  by  Smith  (1939:209): 
"Species  of  this  group  are  as  a  rule  con- 
fined to  rocky  habitats.  So  far  as  I  am 
aware,  only  cyanogenys  tends  to  live  on 
or  near  the  ground."  Thus,  the  new 
]:)hylogeny  indicates  a  trend  in  this  group 
from  small-sized,  small-scaled  ground 
dwellers  to  large-sized,  large-scaled  rock 
dwellers.  With  this  reversal  in  direction, 
the  remaining  differences  between  the  two 
phylogenies  in  Figtire  9  are  negligible 
and  the  trends  within  this  group  fit  the 
overall  phylogeny  of  the  genus. 

In  the  genus  Sceloporus,  the  spinosus 
group  has  been  the  object  of  more  system- 
atic study  than  any  other.  No  less  than 
four  different  phylogenetic  trees  have  been 
proposed  by  Smith,  Bussjaeger,  Cole,  and 
Hall.  The  confusion  is  further  compound- 
ed by  the  fact  that  the  spinosus  group  is 
the  largest  in  number  of  species  and  sub- 
species. The  four  phylogenetic  trees  and 
our  conclusions  are  presented  in  Figure 
10.  Smith  (1939)  included  acanthinus, 
lunaei.  and  lundelli  with  this  group.  In 
1950,   he   and   Tavlor  moved   acanthinus 


Fig.   10.     Phylogeny  of  spinosus  group  according  to  Smith  (1939),  Cole  (1970),  Bussjaeger  (1971), 
Hall  (pers.  comm.  1973),  and  the  new  phylogeny  (L  and  T). 


12 


GREAT  BASIN  NATURALIST 


Vol.  35,  No.  1 


and  lunaei  into  the  formosus  group.  How- 
ever, in  1939  Smith  (p.  60)  said,  "The 
most  primitive  form  of  the  group  is  un- 
doubtedly lunaei.  which  is  closely  related 
to  formosus  malachiticus.  S.  acanthinus 
is  a  near  relative  of  lunaei.  as  is  also  lun- 
delli.'"  It  should  therefore  be  acceptable 
to  remove  lundelli  from  the  spinosus  group 
and  place  it  in  the  formosus  group  next 
to  lunaei  as  we  have  done. 

Behavioral  data  also  support  this  ar- 
rangement. Bussjaeger  (1971:136)  ob- 
served: 

The  display-action-patterns  of  lundelli 
gaigei  of  the  spinosus  group  and  asper,  acan- 
thinus acanthinus  and  a.  lunaei  of  the 
formosus  group  were  quite  similar  with 
peaked  single  units  and  multiple  units. 
Sceloporus  asper  and  lundelli  seemed  to 
share   more   elements. 

In  his  conclusions,  Bussjaeger  (p.  151)  an- 
ticipated the  new  position  of  S.  lundelli: 

The  status  of  lundelli  is  questionable.  .  .  . 
Its  display-action-pattern  was  between  acan- 
thinus and  orcuiti;  but  the  pattern  was 
based  on  only  one  female.  More  data  are 
needed  to  establish  this  species  relationship. 
At  present  it  should  be  left  in  the  spinosus 
group,  although  it  appears  to  be  closer  to 
the  formosus  group. 

Cole's  (1970)  phylogenetic  tree  would 
xiot  allow  the  removal  of  lundelli  from 
this  group  unless  melanorhinus  and  clarki 
were  placed  elsewhere.  Cole  (p.  39,  Fig. 
17)  showed  how  four  centric  fusions  could 
change  the  melanorhinus-clarki  karyotype 
into  the  typical  pattern  for  this  group. 
According  to  Cole's  assumption  that  only 
fusions  (i.e.,  no  fissions)  are  possible, 
melanorhinus  and  clarki  are  primitive  not 
only  for  this  group,  but  also  for  the 
genus  Sceloporus.  and  for  the  entire  fam- 
ily Iguanidae!  As  demonstrated  by  Web- 
ster, Hall,  and  Williams  (1972),  chromo- 
somal evolution  can  occur  by  fission  as 
well  as  fusion.  We  believe  this  is  the 
only  acceptable  explanation  for  the  karyo- 
type in  melanorhinus  and  clarki.  If  fission 
is  accepted  as  well  as  fusion,  Cole's  data 
provide  support  for  our  arrangement  of 
orcutti,  clarki.  and  melanorhinus.  (They 
also  confirm  the  primitive  position  of 
lundelli  and  permit  its  placement  in  the 
formosus  group.) 

If  clarki  and  melanorhinus  are  derived 
from  orcutti  and  if  lundelli  is  removed 
from  the  group,  then  the  only  difference 
between  Cole's  tree  and  ours  is  a  minor 


shift  in  the  position  of  edwardtaylori. 
The  single  remaining  difference  between 
Smith's  tree  and  ours  is  the  placement  of 
edwardtaylori.  The  close  relationship  of 
edwardtaylori  to  spinosus  and  horridus 
has  been  proposed  by  Cole  and  also  by 
Hall.  The  justification  is  that  the  species 
clustering  on  one  side  {olivaceus.  cautus, 
edwardtaylori.  spinosus.  and  horridus)  all 
have  22  chromosomes,  whereas  orcutti  has 
34,  magister  has  26,  and  clarki  and  me- 
lanorhinus each  have  40. 

Zoogeography 

The  phylogeny  of  the  genus  Sceloporus 
can  be  considered  with  its  present  geo- 
graphical distribution  to  produce  a  theo- 
retical history  of  events  in  the  speciation 
in  this  genus.  We  conclude  that  the  an- 
cestral sceloporine  was  a  tropical  or  sub- 
tropical lizard  (as  Smith  reasoned)  \vith 
a  distribution  somewhat  matching  the  sub- 
tropical conditions  of  western  America 
before  the  Madro-Tertiary  revolution 
(Ballinger  and  Tinkle(  1972:^63).  This  dis- 
tribution was  not  restricted  to  southern 
Mexico,  where  Smith  pro])osed  the  begin- 
ning of  Sceloporus  evolution,  but  covered  a 
vast  area  in  the  western  United  States  ex- 
tending as  far  north  as  Canada. 

Milstead  (1960:76)  said,  "Formation 
of  the  western  deserts  is  presumed  to  have 
begun  in  Miocene  times  and  continued 
through  Pliocene  and  into  early  Pleisto- 
cene times."  Accordingly,  the  derivation 
of  the  Scelporine  genera  could  have  oc- 
curred in  late  Miocene  and  early  Pliocene 
times  during  the  development  of  the  west- 
ern deserts  (Ballinger  and  Tinkle,  1972). 

The  formation  of  deserts  trapped  a 
mesic-adapted  relict  (Petrosaurus)  in  Baja 
California.  The  remaining  sceloporine 
stock  began  adapting  to  the  oncoming 
desert  conditions  with  such  characters  as 
a  lengthened,  sinuous  nasal  passage  and 
the  behavior  called  "shimmy  burial" 
(Stebbins,  1944).  The  separation  of  the 
generic  lines  of  Uta,  Urosaurus,  Sator, 
and  Sceloporus  was  accomplished  during 
the  initial  stages  of  adaptation  to  desert 
conditions. 

As  tropical  conditions  moved  south- 
ward during  middle  and  late  Pliocene 
(Axelrod,  1948),  the  ancestral  stock  of 
Group  I  moved  south  almost  as  far  as 
the  Isthmus  of  Tehuantepec.  Some  popu- 
lations did  not  migrate,  but  remained  and 


March  1975 


LARSEN,  TANNER:    SCELOPORINE  LIZARDS 


13 


adapted  to  more  xeric  conditions  (Group 
III).  The  mountains  of  central  and  south- 
ern Mexico  J)ro^  ided  a  barrier  that  sepa- 
rated the  western  Group  I  and  eastern 
Group  II  populations.  A  relict  genus 
(Sator)  was  isolated  in  Baja  California  at 
this  time  (Fig.  11).  The  subsequent  de- 
velopment of  Grou])s  I  and  II  was  a  mat- 
ter of  adaptive  radiation  and  centrifugal 
speciation    (Brown,   1957). 

Figure  12  shows  the  routes  of  speciation 
in  Group  I.  The  eastern  branch  extended 
from  gadoviac  (in  southern  Michoacan, 
Guerrero,  Morelos,  southern  Puebla,  and 
northwestern  Oaxaca)  northward  across 
the  Oaxaca  Upland,  the  Neovolcanic  Pla- 
teau and  into  the  Sierra  Madre  Oriental 
to  parvus  (in  Nuevo  Leon,  southeastern 
Coahuila,  San  Luis  Potosi,  and  Hidalgo). 
Speciation  continued  northward  along  the 
Sierra  Madre  Oriental  to  couchi  (Nuevo 
Leon,  eastern  Coahuila,  and  southern  Tex- 
as) and  merriami  (northern  Coahuila  and 
adjacent  Texas).  (Locality  information 
in  this  discussion  is  from  Smith  and  Tay- 
lor, 1950.  Topographical  terminology  is 
from  Raisz,   1964.) 

The  second  branch  of  Group  I  extended 
from  parvus  to  jalapac  (Veracruz,  Pueb- 
la, and  Oaxaca).  This  radiation  then 
moved  across  the  Mixtec  Upland  (along 
the  northern  border  of  Oaxaca)  and  north- 
ward along  the  western  flank  of  the  Sier- 
ra Madre  del  Sur  (through  Guerrero, 
Michoacan,    Colima,     and     Jalisco)     and 


Fig.   12.     Speciation  in  Group  I. 

further  northward  along  the  western  flank 
of  the  Sierra  Madre  Occidental  (through 
Nayarit  and  Sinaloa  and  into  Durango) . 
The  Durango  populations  became  niaculo- 
sus,  and  most  of  the  pathway  is  now  oc- 
cupied by  ochotcrenae. 

Figure  13  shows  the  initial  radiation 
from  the  ancestral  stock  of  Group  II.  This 
ancestral  stock  is  now  represented  by 
pictus  (in  central  Puebla  and  central 
western  Veracruz).  The  first  radiation 
involved  four  species  in  four  directions: 
aencus  to  the  north,  pyrocephalus  to  the 
west,  sinifcrus  to  the  south,  and  cozumelac 
to  the  east. 

Subsequent  radiation  from  these  cen- 
ters is  shown  in  Figure  14.  Sceloporus 
aeneus  (Puebla,  Veracruz,  Oaxaca,  Hidal- 
go, Morelos,  Mexico,  (juanajuato,  Micho- 
acan, and  Jalisco)  produced  scalar  is  (ni 
Durango,  Guanajuato,  Hidalgo,  Jalisco, 
Mexico,  Michoacan,  Puebla,  and  Zacate- 
cas).  S.  pyrocephalus  (Guerrero,  Michoa- 
can, and  Colima)  produced  nrlsoni  (in 
Chihuahua.  Jalisco,  Sinaloa,  and  NaA'arit). 


Fig.    li.     Isolation    of    early    Sceloporus    stocks 
response  to  desert  formation  in  middle  Pliocene. 


Fig.   13.     Early    radiation    ui    Group    II. 


14 


GREAT  BASIN  NATURALIST 


Vol.  35,  No.  1 


Fig.   14.     Second 


These  two  species  occupy  most  of  the 
western  flank  of  the  Sierra  Madre  Occi- 
dental. According  to  Hall,  the  separation 
of  nelsoni  and  pyrocephalus  occurs  along 
a  river  in  Nayarit  (the  Rio  Grande  de 
Santiago).  Concerning  this  river,  Hall 
(pers.  comm.,  1973;  see  also  Hall,  1973: 
115-125)  said: 

Evidence  from  the  fresh  water  fish  fauna 
in  the  Rio  Grande  de  Santiago  (Salvador 
Contreras  B..  pers.  comm.)  suggests  that  at 
one  time  this  major  river  drained  the 
greater  part  of  the  Mexican  Plateau.  Even 
now  it  is  the  outlet  for  Lake  Chapala  and 
the  entire  Rio  Lenna  e.xtending  east  as 
far  as  the  western  border  of  the  Distrito 
Federal.  Although  rivers  usually  are  not 
very  effective  natural  barriers,  the  steep 
gradient  of  this  river  as  it  falls  off  the 
Plateau  and  the  comparative  narrowness  of 
the  costal  plain  probably  would  have  made 
it  an  extremely  effective  barrier  during  the 
Pleistocene  pluvial  times,  which  would 
have  provided  ample  opportunity  for  the 
splitting  of  the  ^troio-nelsoni  into  two  stocks. 

The  southern  speciation  produced  sin- 
iferus  (in  Oaxaca,  Chiapas,  and  Guerre- 
ro), carinatus  (in  Chiapas),  squamosus 
(along  the  Pacific  slopes  from  Chiapas  to 
Costa  Rica),  and  utiformis  (to  the  north 
along  the  Pacific  slopes  of  Michoacan, 
Colima,  Jalisco,  Nayarit,  and  Sinaloa). 
The  eastern  branch  to  cozumclae  (in  the 
northern  peninsular  states  of  Yucatan  and 
Quintana  Roo)  produced  chrysostictus  (in 
the  entire  Yucatan  Peninsula),  teapensis 


(in  southern  Veracruz,  Tabasco,  Cam- 
peche,  Quintana  Roo,  northern  Guate- 
mala, and  British  Honduras),  and  variabi- 
lis (which  has  developed  subspecies  along 
the  Gulf  Coast  plain  from  south-central 
Texas,  through  Nuevo  Leon,  Tamaulipas, 
San  Luis  Potosi,  Queretaro,  Hidalgo,  Tlax- 
cala,  Puebla,  and  Veracruz,  across  the 
Isthmus  of  Tehuantepec,  through  Oaxaca 
and  Chiapas,  and  into  western  Guatema- 
la). 

The  central  stock  of  Group  II  also  pro- 
duced a  second  wave  of  speciation.  A 
southern  speciation  from  pictus  produced 
cry  plus  in  the  Oaxaca  highlands.  A  west- 
ern speciation  resulted  in  asper  (in  the  Si- 
erra Madre  del  Sur  in  Guerrero  and  Mi- 
choacan and  extending  as  far  north  as  the 
Sierra  Madre  Occidental  in  Nayarit). 
This  branch  also  produced  heterolepis  in 
the  coastal  mountains  of  Jalisco. 

An  eastern  branch  from  pictus  pro- 
duced megalepidurus  in  Northern  Puebla 
on  the  eastern  slopes  of  the  Neovolcanic 
Plateau.  The  most  recent  derivation  from 
the  pictus  stock  is  grammicus.  This  spe- 
cies has  invaded  most  of  the  Plateau  re- 
gions in  Mexico.  The  distribution  of 
grammicus  is  widespread,  and  Hall  (1971) 
has  suggested  that  there  ma}'  be  as  many 
as  6  cryptic  species  in  the  grammicus  com- 
plex. Further  discussion  of  this  species 
must  therefore  be  deferred  .until  the  alpha 
taxonomy  is  more  complete. 


March  1975 


LARSEN,  TANNER:    SCELOPORINE  LIZARDS 


15 


Speciation  in  Group  III  was  more  com- 
plex and  probably  more  recent  than  in 
the  others.  Other  workers  have  suggested 
that  considerable  speciation  resulted  from 
repeated  glaciation  in  Pleistocene  times 
(Savage,  1960;  Ballinger  and  Tinkle, 
1972).  Each  glacial  period  forced  desert 
species  into  southern  refugia  from  which 
they  later  speciated  through  adaptive 
radiation  and  centrifugal  speciation. 

Group  III  remained  originally  in  the 
north  and  adapted  to  the  xeric  conditions 
of  the  southwest  during  middle  and  late 
Pliocene,  as  did  Uta  and  Urosaurus.  Sub- 
sequent Pleistocene  glaciation  forced  the 
desert-adapted  populations  into  southern 
refugia  with  massive  northern  extinctions. 
The  five  refugia  south  of  30°  latitude  in- 
clude Baja  California,  the  Sonoran  Desert, 
the  Mexican  Plateau,  the  Gulf  Coastal 
Plain,  and  Florida.  Barriers  include  the 
Gulf  of  California,  the  Sierra  Madre  Oc- 
cidental, the  Sierra  Madre  Oriental,  and 
the  Gulf  of  Mexico.  Ballinger  and  Tinkle 
(1972)  discussed  the  first  three  refugia 
in  considerable  detail  with  reference  to 
the  e^■olution  of  Uta. 

After  each  glacial  period,  the  isolated 
populations  expanded  in  all  directions 
from  their  refugia.  (A  worldwide  increase 
in  rainfall  would  restrict  the  midlatitude 
deserts  from  both  sides.  A  subsequent  de- 
crease in  rainfall  would  cause  a  movement 
of  xeric  conditions  both  northward  and 
'southward  from  a  small  latitudinal  band.) 
Each  southerly  movement  was  preserved 
as  the  species  adapted  to  subtropical  con- 
ditions, but  the  northerly  radiations  would 
bo  eliminated  during  the  next  glacial  peri- 
od (southern  rains  could  be  tolerated  bet- 
ter than  northern  snows) . 

Each  invasion  to  the  south  required  a 
secondary  adaptation  to  the  ancestral  en- 
vironment. This  explains  why  formosus 
has  not  yet  lost  a  behavioral  trait  called 
"shimmy  burial."  Hall  (pers.  comm.;  see 
also  Hall   1973:99-102)    said: 

One  gathers  from  Cole's  (1970)  discus- 
sion that  he  uncritically  accepts  Smith's 
(1939)  idea  that  the  arboreal,  tropical  for- 
mosus group  is  primitive  in  the  genus. 
Smith  (pers.  comm.)  believed,  not  unreas- 
onably on  the  limited  infoiTnation  then 
available,  that  the  closest  primitive  relatives 
of  sceloporus  were  the  South  American  trop- 
idurines  (from  which  Weigmann  separated 
Sceloporus),  and  that  its  close  xeric  adap- 
ted relatives  (i.e.  "Uta"  =--  Petrosaurus, 
Urosaurus,  and  Uta)  were  derived  from 
within    tlie    radiation    of    Sceloporus.     The 


work  of  Savage  (1958),  Etheridge  (1964), 
and  Presch  (1969)  tends  to  refute  this 
idea.   .   .  . 

Furthermore,  it  is  interesting  to  note  that 
the  behavioral  trait  of  'shimmy  burial'  .  .  . 
is  also  found  in  most  other  Sceloporines.  .  .  . 

From  this  analysis,  it  would  seem  that 
all  sceloporines  above  Petrosaurus  at  least 
primitively  know  how  to  use  loose  sand 
for  escape  and  sleeping  cover.  It  seems  un- 
likely that  this  behavior  would  evolve  in  a 
supposedlj'  primitive  fomi  like  formosus, 
which  lives  in  inountain  rain  forests  where 
the  lizards  would  rarely  or  never  encounter 
a  suitable  substrate  for  shimmy  burial.  Its 
presence  in  this  species  probably  indicates 
only  that  formosus  has  only  very  recently 
entered  the  rain  forest  habitat.  On  the  other 
hand,  shinnnj'  burial  would  be  selectively 
valuable  to  a  species  inhabiting  dry  plains 
or  deserts  where  loose  sand  might  fre- 
quently be  the  only  cover  available  for 
escape  or   sleeping. 

This  quotation  explains  why  Smith  (1939) 
and  Cole  (1970)  proposed  phylogenies 
from  south  to  north.  We  propose  a  re- 
versal of  these  phylogenies,  which  means 
that  most  trends  in  Group  III  are  from 
the  north  and  that  the  Group  III  forms 
moved  southward  and  adapted  to  a  climate 
similar  to  the  one  in  which  the  ancestors 
lived. 

The  smaller  size  and  greater  isolation  of 
Baja  California  have  limiited  the  genetic 
potential  of  its  populations.  This  has  al- 
lowed continental  species  to  move  north 
from  the  Sonoran  Desert  and  enter  the 
peninsula  to  trap  southern  relicts  (see 
Savage,   1960). 

Another  possible  explanation  for  relict 
species  in  Baja  California  is  the  separa- 
tion and  westward  drift  of  the  peninsula 
in  Miocene-Pliocene  times.  Concerning 
this  movement,  Moore  and  Buffington  (p. 
1241)  said,  "Therefore,  from  about  4  to 
10  million  years  ago,  during  late  Miocene 
and  Pliocene  times,  a  proto-Gulf  of  Cali- 
fornia existed.  .  .  .  The  present  cycle  of 
spreading  began  about  4  million  years 
ago." 

lanner  (1966:191)  stated  that  this 
same  event  could  apply  to  the  night 
snakes: 

Thus  the  distribution  of  Eridiphus  stock 
may  have  reached  southern  Baja  California 
by  a  shorter  route  before  the  present  Gulf 
of  California  was  formed.  Assuming  this 
to  be  correct,  Eridiphus  is  a  relic  of  a  once 
more  widespread  group  of  snakes  in  West- 
ern Me.xico. 

Hall  (1973)  has  suggested  that  such  a 
mechanism    is   responsible    for   speciation 


16 


GREAT  BASIN  NATURALIST 


Vol.  35,  No.  1 


in  Baja  California  and  that  the  Cape  re- 
gion was  isolated  from  the  rest  of  the 
peninsula  as  well  as  the  mainland  during 
an  intermediate  stage. 

The  first  glacial  advance  divided  Scelo- 
porus  into  four  refugia:  an  orcutti  stock 
in  Baja  California,  a  formosus  stock  in 
the  Sonoran  Desert,  a  virgatus  stock  on 
the  Mexican  Plateau  and  a  cyanogenys 
stock  on  the  Gulf  Coastal  Plain.  Subse- 
quent postglacial  speciation  is  illustrated 
in  Figure  15. 

The  virgatus  stock  expanded  northward 
and  as  far  eastward  as  Florida.  It  also  ex- 
panded westward  into  the  Sierra  Madre 
Occidental.  Most  of  the  expansion  from 
this  stock  was  reduced  to  refugia  during 
a  second  glacial  advance.  The  second  gla- 
cial advance  was  less  severe  than  the  first 
(Ballinger  and  Tmkle,  1972:63)  and  a 
population  survived  in  Florida  (ivoodi) . 
The  main  virgatus  stock  was  again  con- 
fined to  the  5lexican  Plateau,  but  some 
of  the  mountain  ])opulations  moved  west 
into  the  Sonoran  refuge.  This  isolation 
produced  graciosus. 

The  subsequent  northward  migration  of 
graciosus  and  the  northern  speciation  of 
undulatus  and  occidentalis  from  virgatus 
is  shown  in  Figure  16. 

The  orcutti  stock,  which  was  confined 


to  the  Baja  California  refuge  during  the 
first  glaciation,  emerged  with  sufficient 
adaptive  specialization  to  displace  the 
formosus  stock  as  far  south  as  Guerrero. 
The  displacement  of  a  mainland  ])opula- 
tion  by  a  restricted  peninsular  ])opulation 
is  explained  by  the  assumption  that  for- 
mosus descended  from  the  part  of  the  Sce- 
loporus  stem  that  had  been  adapting  to  the 
mountain  habitat  between  the  central 
plains  and  the  western  deserts.  As  the 
Pacific  slopes  became  more  and  more  arid 
following  glacial  retreat,  the  desert-adapt- 
ed orcutti  stock  displaced  the  mountain- 
adapted  fonnosus  stock. 

From  the  Pacific  slopes  in  Guerrero,  the 
formosus  stock  speciated  southward,  pro- 
ducing formosus  (with  subspecies  in  Guer- 
rero and  the  central  uplands  of  Oaxaca), 
malachiticus  (along  the  Pacific  slopes  from 
Chiapas  to  Panama),  lunaei  (in  the  up- 
lands of  central  Guatemala),  lundeUi  (in 
the  central  regions  of  the  Yucatan  Penin- 
sula), and  tanneri  in  Oaxaca  (Smith  and 
Larsen,  1975). 

Farther  north  along  the  Pacific  Coast, 
the  orcutti  stock  produced  clarki  (from 
central  Arizona,  through  the  center  of 
Sonora  and  down  the  Pacific  Coast  of 
Sinaloa  to  Nayarit)  and  melanorhinus 
(along   the    Pacific    slopes    from   Nayarit 


Fig.  15.     Eai'ly   radiation  in   Group   III. 


March  1975 


LARSEN,  TANNER:    SCELOPORINE  LIZARDS 


17 


Fig.  16.     Second   radiation  in  Group  III. 


through  Jalisco,  Colima,  Michoacan,  Guer- 
rero, and  Oaxaca  to  Chiapas).  Hall's  com- 
ments about  the  separation  of  nelsoni  and 
pyrocephalus  along  the  Rio  Grande  de 
Santiago  are  also  appropriate  for  clarki 
and  melanorhinus .  Apparently  this  river 
was  a  geographic  barrier  for  two  groups 
speciating  in  opposite  directions. 

Another  branch  from  the  orcutti  stock 
produced  the  nuigister  complex.  The  sub- 
sequent subspeciation  of  magister  accord- 
ing to  Phelan  and  Brattstrom  (1955)  was 
from  central  California  southward  into 
Baja  California  and  southeastward  into 
Arizona  and  New  Mexico.  However,  or- 
cutti has  34  chromosomes,  magister  zos- 
teromus  (and  all  other  peninsular  sub- 
species of  magister)  has  30,  and  m.  magis- 
ter has  26.  This  supports  Hall's  ph^logeny 
with  early  speciation  in  Baja  Cahfornia 
and  subsequent  emergence  of  two  stems 
{orcutti  and  magister). 

A  third  and  final  branch  from  the  or- 
cutti stock  moved  eastward  through  the 
interglacial  deserts  of  Arizona,  New  Mex- 
ico, and  Texas.  This  branch  (olivaceus) 
became  trapped  in  the  Gulf  Coastal  Plain 
refuge  during  the  second  glacial  period 
(Fig.  15).  Speciation  proceeded  from  oli- 
vaceus (central  Texas,  Tamaulipas,  Nuevo 
Leon,  and  adjacent  states)  southward 
across  the  Central  Meseta  to  spinosus  (oc- 
cupying the  entire   Neo volcanic   Plateau 


from  Puebla  and  Veracruz  on  the  east  to 
the  tip  of  Durango  on  the  west),  horridus 
(with  subspecies  along  the  entire  southern 
flank  of  the  distribution  of  spinosus),  and 
edwardtaylori   (in  Oaxaca)    (Fig.   16). 

A  secondary  speciation  from  olivaceus 
(to  cautus)  has  been  questioned  by  Hall 
(because  of  intergrades),  but  he  (pers. 
comm.,  1973)   did  make  this  observation: 

Most  interestingly  there  seems  to  be  al- 
most no  question  that  cautus  and  olivaceus 
intergrade  south  and  west  of  Monterrey 
(Nuevo  Leon)  with  gene  flow  occurring 
presently  through  the  dry  valleys  and 
passes.  There  might  be  an  absolute  classic 
circle  of  subspecies  whose  terminal  popula- 
tions are  fully  sympatric. 

The  last  major  speciation  wdthin  Scelo- 
porus  started  with  cyanogcjiys  in  the  Gulf 
Coastal  Plain  refuge  (Fig.  15).  The  first 
branch  produced  jarrovi  (in  the  northern 
plateaus  and  adjacent  escarpments  from 
Arizona  on  the  northwest  to  Veracruz  on 
the  southeast),  which  in  turn  produced 
ornatus  (in  the  ranges  of  southern  Coahu- 
ila),  lineolatcralis  (restricted  to  the  moun- 
tains of  eastern  Durango),  and  dugesi 
(with  subspecies  in  the  mountains  of  Gua- 
najuato, Michoacan,  Colima,  Jalisco,  and 
Nayarit.) 

The  second  branch  from  cyanogenys 
moved  westward  to  produce  poinsetti 
(which    occupies    most    of    the    northern 


18 


GREAT  BASIN  NATURALIST 


Vol.  35,  No.  1 


Plateau  through  southern  New  Mexico, 
southwestern  Texas,  and  the  Mexican 
states  of  Chihuahua,  Coahuila,  and  Du- 
rango).  The  third  branch  extended  across 
Mexico  in  a  southwesterly  direction  and 
resulted  in  hullcri  (in  the  mountains  of 
Jalisco) . 

The  final  radiation  from  the  cyano- 
genys  stock  extended  southward  and  re- 
sulted in  serrifer  (occupying  most  of  the 
Gulf  Coastal  Plain  in  Tamaulipas,  San 
Luis  Potosi,  Veracruz,  Tabasco,  Cam- 
peche,  and  Yucatan),  mucronatus  (a 
mountain  form  in  the  Oaxaca  Upland  and 
other  mountains  in  the  state  of  Guerrero, 
Veracruz,  Puebla,  Mexico,  and  Hidalgo), 
and  torquatus  (which  inhabits  a  large 
area  in  central  Mexico,  including  parts  of 
Hidalgo,  Veracruz,  Mexico,  Distrio  Fede- 
ral, Puebla,  Morelos,  Guanajuato,  Micho- 
acan,  Nuevo  Leon,  Jalisco,  San  Luis  Poto- 
si, and  Zacatecas). 

Conclusions 

When  presenting  his  arrangement, 
Smith  (1939)  said,  "Material  from  cer- 
tain areas  is  still  lacking,  and  more  direct 
evidence  of  relationships  is  frequently  to 
be  desired.  The  conclusions  now  ])resented 
are  accordingly  tentative."  Smith's  state- 
ment may  still  apply.  Problem  areas  in- 
clude Baja  California  and  the  grammicus 
complex.  Also  several  new  species  and 
subspecies  are  being  considered  by  various 
workers.  New  kinds  of  data  are  now  be- 
ing researched  (microdermatoglyphics, 
for  example).  However,  a  point  has  been 
reached  at  which  different  sets  of  data 
reinforce  similar  conclusions.  With  over 
80  characters,  the  new  groups  and  sub- 
groups are  distinct  at  the  .999  level  of 
confidence  (Larsen  and  Tanner,  1974). 
With  such  a  high  level  of  confidence,  we 
conclude  that  Figure  5  is  a  natural  ar- 
rangement of  species  and  that  future  ad- 
justments may  be  minor. 

When  phylogeny  and  zoogeography  are 
considered  simultaneousl}-,  several  trends 
are  evident  in  the  evolution  of  SccJoporus: 

(1)  the  size  altered  from  small  to  large; 

(2)  the  scales,  once  small,  smooth,  and 
granular,  changed,  becoming  large,  carin- 
ate,  mucronate,  and  imbricate;  (3)  ini- 
tial movement  and  speciation  was  from 
north  to  south,  and  several  secondary  ra- 
diations were  from  southern  centers  north- 
ward  and   from   northern   centers   south- 


ward; (4)  the  geography  of  Baja  Cali- 
fornia created  several  relicts;  (5)  habitat 
preference  changed  from  ground  to  rocks, 
cliffs,  and  trees;  and  (6)  the  ancestral 
stock,  which  originally  was  subtropical, 
adapted  to  arid  conditions,  and  then  sever- 
al groups  returned  to  tropical  or  sub- 
tropical climates. 

Cope  (1900)  called  SccJoporus  the  piece 
de  resistance  for  the  theory  of  derivation 
of  species.  This  genus  seem  to  show  such 
principles  as  parallelism,  convergence,  di- 
vergence, genetic  drift,  geographical  bar- 
riers, adaptive  radiation,  centrifugal  spe- 
ciation, and  waif  and  relict  population 
development.  In  fact,  the  cape  region  of 
Baja  California  may  provide  examples  of 
speciation  by  continental  drift.  Sceloporus 
also  exhibits  a  high  degree  of  chromoso- 
mal variation,  including  examples  of  Rob- 
ertsonian  fission  and  fusion,  and  several 
formulae  for  sex  determination.  This  ge- 
nus is  extremely  well  suited  for  illustra- 
tion and  discussion  of  evolutionary  theory. 

We  conclude  that  Sceloporus  has  re- 
cently speciated  in  an  explosive  manner. 
Because  of  this  ra]:)id  adaptive  radiation, 
it  is  difficult  to  determine  phylogenetic 
relationships  with   classical  techniques. 

We  are  ])ersuaded,  however,  that  the 
genus  Sceloporus  does  contain  three  dis- 
tinct monophyletic  groups.  Grou]:)  I  is  dis- 
tinct from  the  other  tw^o  groups  in  having 
(1)  a  postfemoral  dennal  pocket  and  less 
than  7  ^'entrals  betw'een  the  femoral  pore 
series  or  (2)  (if  the  postfemoral  dermal 
pocket  is  absent)  a  vestigial  gular  fold 
and  no  postrostrals.  The  rest  of  the  spe- 
cies in  the  genus  Sceloporus  lack  either  a 
jiostfemoral  dermal  pocket  or  a  vestigial 
gular  fold.  If  they  lack  the  vestigial  gular 
fold,  postrostrals  are  ])resent  and  there  are 
more  than  8  ^'entrals  between  the  femoral 
pore  series.  In  considering  the  systemat- 
ics  of  the  entire  complex,  we  believe  that 
it  is  now  feasible  to  recognize  for  Group 
I  (Table  1)  the  Cope  (1888)  monotypic 
generic  designation  of  Lysoptychus  (L. 
lateralis:=Sceloporus  couchi  Baird,  1858). 

We  have  not  by  our  methods  been  able 
to  arrive  at  a  satisfactory  taxonomic  divi- 
sion of  Groups  II  and  III,  even  though 
these  groups  become  sej)arable  and  distinct 
by  use  of  multivariate  analysis.  We  be- 
lieve that  Groups  II  and  III  represent  a 
large  assemblage  of  species  that  have 
evolved  more  recently  but  that  although 
the    characters    between    the    groups    are 


March  1975 


LARSEN,  TANNER:    SCELOPORINE  LIZARDS 


19 


showing  indications  of  evolutionary  separ- 
ation, they  have  not  reached  a  point  of 
distinction  that  permits  the  development 
of  a  workable  taxonomic  key.  We  there- 
fore choose  at  this  time  to  retain  them  in 
the  genus  Sceloporus. 

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NEW  SYNONYMY  AND  NEW  SPECIES  OF  AMERICAN 
BARK  BEETLES  (COLEOPTERA:  SCOLYTIDAE)^ 

Stephen  L.  Wood" 

Abstract. —  New  synonymy  is  proposed  as  follows:  Pityokteines  Fuchs  {  =  Orthotomides  Wood), 
Araptus  foveifrons  (Schedl)  {  =  inter jectus  Wood),  n.  comb.,  Cryptocarenus  seriatus  Eggers  {  =  boliv- 
ianus  Eggers),  Hylastes  gracilis  LeConte  {=asper  Swaine),  Procryphalus  utahensis  Hopkins  {=aceris 
Hopkins),  Scolytodes  maurus  (Blandford)  {=Hexacolus  ellipticus  Eggers),  Scolytus  hermosus  Wood 
{=sylvaticus  Bright),  Xyleborus  adelographus  Eichhoff  {  =  vitiosus  Schedl),  Xyleborus  catulus 
Blandford  {=intricatus  Schedl),  Xyleborus  nepos  Eggers  {=signatus  Schedl),  Xyleborus  titubanter 
Schedl  {=dissidens  Wood).  The  following  species  and  subspecies  are  named  as  new  to  science: 
Cnesinus  electus  (Costa  Rica),  C.  pilatus  (Mexico),  Hylastes  asperatus  (New  Mexico),  Scolytus  toru- 
lus  (Costa  Rica),  S.  laetus  (Mexico),  Scolytodes  amabilis  and  S.  lepidus  (Mexico),  S.  obesus  (Pana- 
ma), Pseudothysanoes  concentralis  and  P.  tumidulus  (Mexico),  Thysanoes  tuberculatus  (Mexico), 
Pityokteines  mystacinus  (Washington),  Ips  pilifrons  subsp.  thatcheri  (Nevada),  Araptus  attemiatus 
(Mexico),  A.  fossifrons  (Mexico,  Guatemala),  Amphicranus  parilis  (Mexico),  Corthylus  cecropii 
(Costa  Rica). 


On  the  following  pages  several  newly 
discovered  cases  of  synonymy  and  1 7  spec- 
ies and  1  subspecies  new  to  science  are 
presented  for  American  Scolytidae.  The 
specific  synonymy  is  presented  in  alpha- 
betical order  for  convenient  reference.  The 
species  new  to  science  represent  the  gen- 
era Cnesinus  (2),  Hylastes  (1),  Scolytus 
(2),  Scolytodes  (3),  Pseudothysanoes  (3), 
Thysanoes  (1),  Pityokteines  (1),  Araptus 
(2),  Amphicranus  (1),  and  Corthylus 
( 1 ) .  The  new  subspecies  is  of  Ips  pili- 
frons. The  new  species  are  from  the  fol- 
lowing countries:  United  States  (2),  Mex- 
ico (10),  Costa  Rica  (3),  Panama  (1), 
Mexico  and  Guatemala  ( 1 ) .  The  new 
subspecies  is  from  the  United  States  (Ne- 
vada). 

New  Synonymy 
Pityokteines  Fuchs 

Pityokteines  Fuchs,  1911,  Morphologische  Studien 
iiber  Borkenkafer:  I.  Die  Gattungen  Ips  De- 
Geer  und  Pityogenes  Bedel,  p.  33  (Type- 
species:  Ips  curvidens  GeiTnar,  subsequent 
designation  by  Hopkins,  1914,  Proc.  U.S.  Nat. 
Mus.  48:127) 

Orthotomides  Wood,  1951,  J.  Ent.  Soc.  Kansas 
24:32  (Type-species:  Orthotomicus  lasiocarpi 
Swaine,  original  designation) .  New  synonymy 

The  discovery  of  mystacinus  Wood, 
described  below,  closes  the  gap  in  charac- 
ters between  Pityokteines  and  Orthoto- 
mides to  such  an  extent  that  the  latter 
name  must  be  placed  in  synonymy  even 
though  lasiocarpi  (Swaine)  entirely  lacks 
the  long  female  frontal  hair  characteristic 
of  Pityokteines.    Female  mystacinus  have 


part  of  the  frontal  vestiture  elongate  and 
also  have  two  sutures  visible  on  the  pos- 
terior face  of  the  antennal  club. 

This  generic  synonymy  necessitates  the 
transfer  of  lasiocarpi  from  Orthotomides 
to  Pityokteines. 

Araptus  foveifrons  (Schedl),  n.  comb. 

Tharnnophthorus  foveifrons  Schedl,  1963,  Ent. 
Arb.  Mus.  Frey  14:161  (Holotype,  male; 
Guadalajara.  Jalisco,  Mexico;  Schedl  Coll.) 

Araptus  interjectus  Wood,  1974,  Brigham  Young 
Univ.  Sci.  Bull.,  Biol.  Ser.  19(1)  :44  (Holo- 
type, male;  Volcan  de  Agua,  Guatemala; 
Wood  Coll.).  New  synonymy 

When  the  North  and  Central  American 
Araptus  were  reviewed  for  the  mono- 
graph, two  closely  related  species  were 
found,  both  of  which  fit  the  description 
of  Tharnnophthorus  foveifrons  Schedl.  It 
was  assumed  that  the  common,  widely 
distributed  species  of  these  two  was  fovei- 
frons; the  rare  one  was  named  interjectus 
Wood.  Recently,  I  had  the  opportunity 
to  examine  the  male  holotype  of  foveifrons 
and  to  compare  it  to  the  male  holotype 
of  interjectus.  They  represent  the  same 
species.  The  common  species  with  which 
it  had  been  confused  is  named  below  as 
Araptus  fossifrons. 

Cryptocarenus  seriatus  Eggers 

Cryptocarenus  seriatus  Eggers,   1933,  Orig.  Mem. 

Trav.    Lab.    Ent.    Paris    1(1):  10     (Holotype, 

female;  Nouveau  Chantier,  French  Guayana; 

Paris  Mus.) 
Cryptocarenus     bolivianus     Eggers,      1943,     Mitt. 

Miinchner    Ent.    Ges.    33:356    (Holotype,    fe- 


■■Part   of   this   research   was   sponsored   hy   the  National   Science  Foundation. 

-Department    of   Zoology,    Brigham   Young   University,    Prove,    Utah  84602.  Scolytidae  rontrihutioii  No,  60 


21 


22 


GREAT  BASIN  NATURALIST 


Vol.  35,  No.  1 


male;  Cochabamba,  Bolivia;  U.S.  Nat.  Mus.). 
New  synonymy 

The  holotypes  of  Cryptocarenus  seriatus 
Eggers  and  boUvianus  Eggers  were  com- 
pared directly  to  my  material  fromi  Bra- 
zil, Venezuela,  and  other  areas.  All  rep- 
resent the  same  biological  species. 

Hylastes  gracilis  LeConte 

Hylastes  gracilis  LeConte,  1868,  Trans.  Amer. 
Ent.  Soc.  2:174  (Lectotype,  female;  Tahoe 
Valley,  California;  Mus.  Comp.  Zool.,  de- 
signated by  Wood,  1971,  Great  Basin  Nat. 
31:145) 

Hylastes  asper  Swaine,  1917,  Dom.  Canada  Dept. 
Agric.  Ent.  Br.  Bull.  14(1):  19  (Holotype, 
female;  Larimer  Co.,  Colorado;  Canadian 
Nat.  Coll.).  New  synonymy 

The  holotype  of  Hylastes  asper  Swaine 
is  rather  different  from  what  the  descrip- 
tion might  lead  one  to  believe.  It  is  vir- 
tually identical  to  the  type  of  longus  Le- 
Conte and  well  within  the  range  of  vari- 
ability of  gracilis  LeConte.  All  three  types 
were  compared  to  my  material  and,  in 
my  opinion,  represent  the  same  species. 
The  species  I  have  previously  treated  as 
asper  is  described  below. 

Procryphalus  utahensis  Hopkins 

Procryphalus  utahensis  Hopkins,  1915,  U.S.  Dept. 
Agric.  Kept.  99:33  (Holotype,  female;  Alta, 
Utah;  U.S.  Nat.  Mus.) 

Procryphalus  aceris  Hopkins,  1915,  U.S.  Dept. 
Agric.  Kept.  99:33  (Holotype,  female;  Al- 
bany, Oregon;  U.S.  Nat.  Mus.).  New  synon- 
ymy 

Procrophalus  aceris  Hopkins  was  named 
from  a  series  of  about  six  specimens  taken 
from  a  limb  that  had  been  cut  in  the 
vicinity  of  Albany,  Oregon.  The  limb 
was  given  to  Hopkins  who  identified  it 
as  Acer  macrophyllum  (personal  com- 
munication from  W.  J.  Chamberlin).  Be- 
cause the  strial  punctures  were  slightly 
larger  than  usual,  I  did  not  associate  these 
specimens  with  utahensis  Hopkins  until 
an  identical  series  was  taken  from  Salix 
scouleriana  at  Dixie  Pass,  Oregon.  This 
series  was  compared  directly  to  the  type 
series  of  utahensis  and  aceris  and  to  other 
series  from  British  Columbia,  Quebec,  Cal- 
ifornia, Colorado,  South  Dakota,  and  Utah. 
Only  one  species  is  represented  by  this 
material;  all  specimens  are  from  Salix. 
Chamberlin,  myself,  and  many  others 
have  searched  both  standing  and  cut  Acer 
macrophyllum     on     numerous     occasions 


without  finding  indications  of  bark  beetle 
activity.  The  host  identification  by  Hop- 
kins must  have  been  erroneous. 

Scolytodes  maurus  (Blandford) 

Prionosceles  maurus  Blandford,  1897.  Biol.  Centr. 
Amer.,  Coleopt.  4(6):  178  (Syntypes;  Cerro 
Zunil  and  Pantaleon,  Guatemala;  British 
Mus.  Nat.  Hist.) 

Hexacolus  ellipticus  Eggers,  1934,  Ent.  Blatt. 
30:80  (Holotype,  male;  Turrialba,  Costa  Ri- 
ca; Institut  fiir  Pflanzenschutzforschung 
Kleinmachnow).  New  synonymy 

The  syntypic  series  of  maurus  (Bland- 
ford) and  the  male  holotype  of  ellipticus 
Eggers  were  compared  to  my  material 
taken  from  southern  Mexico  to  Panama. 
Only  one  common  species  is  represented 
by  this  material.  In  all,  158  specimens 
were  examined. 

Scolytus  hermosus  Wood 

Scolytus  hermosus  Wood,  1968,  Great  Basin  Nat. 
28:12  (Holotype,  male;  2  km  N  Tlaxcala 
state  line  18  km  N  Tlaxco,  Puebla,  Mexico; 
Wood  Coll.) 

Scolytus  sylvaticus  Bright,  1972,  Canadian  Ent. 
104:1489  (Holotype,  male;  Cerro  Potosi,  Nu- 
evo  Leon,  Mexico;  Canadian  Nat.  Coll.). 
New  synonymy 

The  holotype  and  allotype  of  sylvaticus 
Bright  and  three  other  specimens  from  the 
type  locality  of  that  species  were  com- 
pared directly  to  the  type  series  of  hermos- 
us Wood.  While  the  holotype  has  abdom- 
inal sternum  2  less  strongly  concave  than 
normal,  the  other  Cerro  Potosi  specimens 
and  Bright's  illustration  of  a  male  from 
the  type  series  (Fig.  1,  p.  1490)  are  nor- 
mal specimens  of  hermosus.  For  this  rea- 
son, sylvaticus  must  be  placed  in  synon- 
ymy. 

Xylehorus  adelographus  Eichhoff 

Xyleborus  adelographus  Eichhoff,  1868,  Berliner 
Ent.  Zeitschr.  11:400  (Syntypes,  female;  Bra- 
zil; Brussels  Mus.) 

Xyleborus  vitiosus  Schedl,  1940,  An.  Esc.  Nac. 
Cienc.  Biol.,  Mexico  1:367  (Holotype,  female; 
Mexico?;  Schedl  Coll.).  New  synonymy 

The  syntype  of  adelographus  Eichhoff 
in  the  Brussels  Museum  is  2.8  mm  in 
length;  the  holotype  of  vitiosus  Schedl  is 
3.5  mm  (not  3.7  mm  as  given  by  Schedl). 
Seven  specimens  in  my  collection  from 
Colombia  are  3.3-3.4  mm,  and  one  from 
southern  Brazil  is  3.5  mm.  The  specimens 
from  Colombia  are  very  slightly  stouter, 


March  1975 


WOOD:    AMERICAN    SCOLYTIDAE 


23 


the  dechvity  along  the  suture  is  flat,  and 
the  punctures  on  the  declivital  striae  are 
slightly  more  deeply  impressed.  The  holo- 
type  of  vitiosus  and  my  Brazilian  speci- 
men of  adelographus  are  identical  in  every 
respect  except  for  size.  These  specimens 
are  almost  identical  to  the  syntype  of 
adelographus  and  are  regarded  as  con- 
specific.  Because  extensive  collecting  has 
not  produced  any  specimens  of  this  spec- 
ies from  Mexico  or  Central  America,  be- 
cause the  Mexican  origin  of  the  holotype 
of  vitiosus  is  questioned  on  its  locality  la- 
bel, and  because  the  holotype  of  vitiosus 
resembles  specimens  from  southern  Brazil 
much  more  closely  than  it  does  represent- 
atives of  this  species  from  Colombia,  this 
species  should  be  dropped  from  the  faunal 
Hst  of  Mexico  until  more  definite  evidence 
of  its  occurrence  in  Mexico  is  found. 

Xyleborus  catulus  Blandford 

Xyleborus    catulus    Blandford,    1898,    Biol.    Centr. 

Amer.,   Coleopt  4(6):216    (Holotype,   female; 

Volcan    de    Chiriqui,    Panama;    British    Mus. 

Nat.  Hist.) 
Xyleborus    intricatus    Schedl,    1949,    Rev.    Brasil. 

Biol.   9:274    (Holotype,   female;    St.   Catarina, 

Brazil;   Schedl  Coll.).  New  synonymy 

Several  specimens  in  the  California 
Academ}^  of  Sciences  were  identified  by 
Schedl  as  Xyleborus  intricatus  Schedl. 
Since  these  specimens  are  of  catulus 
Blandford,  a  loan  of  the  holotype  of  in- 
tricatus was  requested.  The  specimen  re- 
ceived was  labeled  as  a  "paratj^pe"  of 
intricatus  from  Nova  Teutonia,  Santa  Ca- 
tarina, Brazil;  however,  this  species  was 
based  on  a  unique  female,  and  paratypes 
were  not  designated  in  the  original  des- 
cription. Because  this  specimen  is  also  of 
catulus,  intricatus  is  tentatively  placed  in 
synonymy  until  the  holotype  is  located. 

Xyleborus  nepos  Eggers 

Xyleborus  nepos  Eggers,  1923,  Zool.  Meded.  7:198 
Lectotype,  female;  Java;  U.S.  Nat.  Mus., 
designated  by  Anderson  and  Anderson.  1971, 
Smithsonian    Contrib.    Zool.    94:21) 

Xyleborus  signatus  Schedl,  1949,  Rev.  Brasil  Biol. 
9:278  (Holotype,  female;  Mexico;  Schedl 
Coll.).  New  synonymy 

The  holotype  of  signatus  Schedl  was 
named  from  a  unique  female  labeled 
"Mexico."  This  specimen  is  identical  to  a 
long  series  of  nepos  Eggers  from  Indonesia 
and  the  Philippine  Islands,  some  of  which 
were  compared  directly  to  the  lectotype 


of  nepos.  Because  extensive  collecting  in 
Mexico  has  not  produced  another  speci- 
men of  this  species,  the  holotype  of  signa- 
tus is  considered  a  mislabeled  specimen 
and  should  be  dropped  from  the  ]\Iexican 
faunal  list  until  more  definite  evidence 
of  its  occurrence  there  is  found. 

Xyleborus  titubanter  Schedl 

Xyleborus  titubanter  Schedl,  1948,  Rev.  de  Ent. 
19: '578  (Holotype,  female;  Mexico;  Schedl 
Coll.) 

Xyleborus  dissidens  Wood,  1972.  Brigham  Young 
Univ.  Sci.  Bull.,  Biol.  Ser.  19(1)  :41  (Holo- 
type, female;  9  km  NE  Teziutlan,  Puebla, 
Mexico;  Wood  Coll.).  New  synonymy 

The  holotypes  of  titubanter  Schedl  and 
dissidens  Wood  were  compared  directly 
to  one  anotlier  and  were  found  to  repre- 
sent the  same  species. 


Taxa  New  To  Science 
Cnesinus  electus,  n.  sp. 

This  species  is  distinguished  from  the 
female  holotype  of  bisulcatus  Schedl  by 
the  more  narrowly  spaced  eyes  (above), 
by  the  smaller,  more  abruptly  elevated 
epistomal  process  (difference  very  slight), 
by  the  very  slightly  larger,  deeper  prono- 
tal  punctures,  by  the  larger,  more  widely 
spaced  strial  punctures,  and  by  the  coarser 
declivital  setae  with  a  complete  row  of 
setae  on  interstriae  1 . 

Female. —  Length  2.5  mm,  (paratypes 
2.3-2.6  mm),  2.8  times  as  long  as  wide; 
color  dark  brown,  elytra  dark  reddish 
brown. 

Frons  strongly,  transversely  impressed 
at  level  of  antennal  bases,  convex  above, 
somewhat  flattened  below  this  point,  with 
epistomal  process  poorly  developed  and 
devoid  of  granules  or  tubercles;  eyes  ap- 
proximate above,  separated  by  a  distance 
equal  to  0.5  times  greatest  width  of  an 
eye  (about  twice  this  wide  in  bisulcatus) ; 
surface  rugulose  and  rather  coarsely 
punctured  above  eyes,  rugose-reticulate 
below  upper  level  of  eyes,  with  a  few 
fine  punctures  along  lateral  margins  and 
on  epistoma;  vestiture  limited  to  lateral 
and  epistomal  areas.  Eyes  very  large; 
very  coarsely  faceted. 

Pronotum  1.1  times  as  long  as  wide; 
widest  just  in  front  of  middle,  almost 
straight  from  widest  point  to  sharply  de- 
fined basal  angles,  somewhat  constricted 


24 


GREAT  BASIN  NATURALIST 


Vol.  35,  No.  1 


just  behind  narrowly  rounded  anterior 
margin;  surface  dull,  subshining,  more 
distinctly  shining  toward  base,  punctures 
almost  round  at  base,  becoming  increas- 
ingly elongate  anteriorly,  occasionally 
confluent  in  anterior  area;  glabrous. 

Elytra  1.7  times  as  long  as  wide;  sides 
almost  straight  and  parallel  on  basal  two- 
thirds,  rather  narrowly  rounded  behind; 
striae  1  rather  strongly,  others  moder- 
ately, not  abruptly  impressed,  punctures 
small,  except  on  1  not  confluent,  distinct- 
ly, rather  strongly  impressed,  separated 
by  distances  slightly  less  than  their  own 
diameters;  interstriae  feebly  convex,  shin- 
ing, punctures  varying  from  minute  to 
two-thirds  size  of  those  of  striae,  in  in- 
definite uniseriate  rows.  Declivity  con- 
vex, except  moderately  impressed  between 
interstriae  3;  striae  feebly  impressed, 
punctures  rather  small  but  impressed;  in- 
terstriae 1  abruptly  elevated  to  apex, 
about  half  as  high  as  wide,  others  almost 
flat,  2  about  one  and  one-half  times  as 
wide  as  1,  twice  as  wide  as  3;  all  interstri- 
ae armed  by  uniseriate  rows  of  moderately 
large  rounded  setiferous  granules,  each 
granule  about  as  high  as  wide,  distinctly 
larger  than  in  elegans.  Vestiture  confined 
to  declivity,  except  on  interstriae  1,  con- 
sisting of  rows  of  rather  coarse,  moderately 
long,  interstrial  bristles,  and  short,  fine, 
strial  hair. 

Female. —  Similar  to  male  except  epis- 
tomal  callus  more  prominent. 

Type  Locality.—  Five  miles  or  8  km 
SE  Cartago,  Cartago,  Costa  Rica. 

Type  Material. —  The  female  holo- 
tvpe,  male  allotype,  and  7  paratypes  were 
taken  on  2-VIII-63,  at  1500  m,  No.  98, 
from  twigs  of  Miconia  (?),  by  S.  L. 
Wood.  Seven  paratypes  bear  the  same 
data  except  that  they  were  from  twigs  of 
an  unknown  tree;  five  paratypes  came 
from  same  locality  on  29-VIT-63,  from  a 
woody  vine;  and  seven  paratypes  are 
from  Peralta,  Cartago,  Costa  Rica,  lO-III- 
64,  500  m,  tree  seedling,  S.  L.  Wood. 

The  holotype,  allotype,  and  parat5q3es 
are  in  my  collection. 

Cnesinus  pilatus,  n.  sp. 

This  species  is  distinguished  from  strigi- 
coUis  LeConte  by  the  larger  size,  by  the 
stouter  body  form,  by  the  different  female 
frons,  by  the  absence  of  tubercles  on  de- 


clivital  interstriae  2,  and  by  other  charac- 
ters noted  below. 

Female. —  Length  3.1  mm,  2.1  times 
as  long  as  wide;  color  dark  brown,  elytra 
dark  reddish  brown. 

Frons  as  in  strigicollis  except  area  below 
carina  slightly  less  strongly  impressed, 
distance  between  eyes  0.8  times  as  wide 
as  width  at  level  of  antennal  insertion  (as 
in  strigicollis) ,  carina  0.5  times  as  wide  as 
distance  between  eyes,  median  impunctate 
area  above  carina  larger,  extending  above 
upper  level  of  eyes,  vestiture  in  lateral 
areas  not  extending  above  upper  level  of 
eyes  and  not  on  median  third  at  vertex 
(extending  well  above  eyes  and  almost 
to  median  line  in  strigicollis). 

Pronotum  0.9  times  as  long  as  wide; 
about  as  in  strigicollis  except  more  close- 
ly, slightly  more  coarsely  strigose. 

Elytra  1.3  times  as  long  as  wide,  1.7 
times  as  long  as  pronotum;  similar  to 
strigicollis  except  interstriae  three  times 
as  wide  as  striae,  declivity  more  broadly 
impressed,  declivital  interstriae  2  devoid 
of  granules,  vestiture  about  half  as  long, 
stouter.  Interstriae  2  on  declivity  with  a 
row  of  punctures,  each  puncture  about 
two-thirds  as  large  as  those  of  adjacent 
striae. 

Type  Locality. —  Thirteen  km  or  eight 
miles  W  El  Palmito,  Sinaloa,  Mexico. 

Type  Material.^ —  The  female  holo- 
type was  taken  at  the  type  locality  on  7- 
Vin-64,  by  H.  F.  Howden. 

The  holotype  is  in  the  Canadian  Na- 
tional Collection. 

Hylastes  asperatus,  n.  sp. 

This  is  a  difficult  species  to  recognize. 
The  very  large  pronotal  punctures  are  ir- 
regular in  size,  as  in  porculus  Erichson, 
but  much  closer;  the  discal  interstriae  are 
slightly  wider  than  in  allied  species,  more 
nearly  convex,  and  very  finely,  closely 
crenulate  (usually  not  clearly  apparent 
unless  the  light  source  is  posterior  to  the 
specimen).  This  species  has  been  treated 
as  asper  Swaine,  but  the  type  of  asper  is 
quite  different. 

Female. —  Length  4.2  mm  (paratypes 
4.0-4.6  mm),  2.7  times  as  long  as  wide; 
color  dark  brown. 

Frons  as  in  gracilis  LeConte  with  inter- 
ocular  impression  moderately  strong, 
fine,  low  carina  evidently  always  extend- 


March  1975 


WOOD:     AMERICAN    SCOLYTIDAE 


25 


ing  from  this  impression  to  base  of  episto- 
mal  lobe  where  it  forks  as  in  related  spec- 
ies. 

Pronotum  1.2  times  as  long  as  wide; 
sides  on  slightl)^  more  than  basal  half 
straight  and  parallel,  rather  broadly  round- 
ed m  front;  surface  subshining,  indica- 
tions of  reticulation  obscure  but  usually 
visible  at  high  magnification  toward  an- 
terior or  basal  areas,  punctures  coarse, 
very  close,  deep,  irregular  in  size  in  some 
specimens,  interspaces  usually  equal  to 
less  than  one-fourth  diameter  of  a  punc- 
ture; median  line  partly  impunctate,  not 
raised;  glabrous. 

Elytra  1.8  times  as  long  as  wide,  1.8 
times  as  long  as  pronotimi;  outline  as  in 
gracilis;  striae  moderately  impressed, 
punctures  rather  coarse,  deep;  interstriae 
as  wide  as  striae,  punctures  moderately 
coarse,  close,  confused,  their  anterior  mar- 
gins elevated  into  fine,  crenulate,  trans- 
verse ridges  of  variable  height  (this  char- 
acter approached  in  allied  species,  but  not 
to  this  degree).  Declivity  convex,  steep; 
striae  narrowly,  deeply  impressed,  punc- 
tures somewhat  obscure;  interstriae  about 
twice  as  wide  as  striae,  armed  by  fine, 
confused  tubercles.  Vestiture  confined  to 
declivity,  scalelike. 

Male. —  Similar  to  female  except 
slightly  stouter,  and  last  visible  abdominal 
sternum  medially  impressed  and  pubes- 
cent. 

Type  Locality. —  New  Mexico. 

Type  Material. —  The  female  holo- 
type,  male  allotype,  and  18  para  types 
were  mounted  on  cards  all  on  one  pin 
bearing  the  label  "New  Mexico,  F.  H. 
Snow."  One  paratype  in  the  Canadian 
National  Collection  labeled  "Las  Vegas 
Hot  Springs,  New  Mexico,  7000  ft.,  Aug. 
'82,  F.  H.  Snow"  may  be  from  the  same 
series.  One  paratype  from  each  of  the 
following  Arizona  localities  is  labeled: 
Hanagan  Camp  Ground,  Greenlee  Co.,  12- 
Vn-68,  D.  E.  Bright;  Santa  Catalina  Mts., 
5-Vin-68,  D.  E.  Bright. 

The  holotype,  allotype,  and  18  para- 
types  are  in  my  collection;  the  three  re- 
maining paratypes  are  in  the  Canadian 
National  Collection. 

Scolytus  torulus,  n.  sp. 

This  species  is  distinguished  from  dimi- 
diatus  Chapuis  by  the  smaller  average  size, 
by  the  subvertical,  somewhat  convex  ab- 


dominal sternum  2,  by  the  absence  of  a 
tuft  of  hair  on  sternum  2  immediately 
posterior  to  the  spine,  by  the  very  differ- 
ent male  frons,  and  by  other  characters 
described  below. 

Male. —  Length  2.4  mm  (paratypes 
2.0-2.4  mm),  2.1  times  as  long  as  wide; 
color  very  dark  brown  to  black,  elytra 
slightly  lighter. 

Frons  rather  weakly  convex  above,  a 
distinct,  moderately  deep,  transverse  im- 
pression just  above  epistoma,  deepest  in 
median  area,  a  distinct  but  weak,  rounded, 
median  elevation  immediately  above  im- 
pressed area;  surface  strongly  reticulate 
above,  more  shining  and  obscurely  acicu- 
late  in  impressed  area,  punctvires  fine, 
rather  deep,  sparse  in  median  area,  more 
numerous  laterally;  vestiture  of  fine  hair 
on  margins  and  impressed  area,  a  few  of 
them  rather  long. 

Pronotimi  as  in  dimidiatus  except  punc- 
tures in  lateral  areas  considerably  larger. 

Elytra  1.1  times  as  long  as  wide,  1.2 
times  as  long  as  pronotum;  surface  simi- 
lar to  dimidiatus  but  much  more  finely 
punctured,  few  to  many  longitudinal  lines 
or  striations  sometimes  present.  Sparse 
setae  shorter  and  much  stouter  than  in 
dimidiatus. 

Sternum  2  subvertical,  moderately  con- 
vex, spine  similar  to  dimidiatus  but  smal- 
ler; surface  dull,  very  coarsely,  deeply, 
closely  punctured;  sterna  3-5  similar  but 
more  finely  punctured;  vestiture  of  very 
fine,  short  hair,  without  a  specialized  tuft 
posterior  to  spine  on  2. 

Female. —  Similar  to  male  except  frons 
without  impression  or  elevation,  not  strig- 
ose,  vestiture  similar  to  but  finer  and 
about  one-third  as  abundant  as  in  dimidi- 
atus female;  spine  on  sternum  2  about 
half  as  large  as  in  male. 

Type  Locality. —  Rincon  de  Osa,  Punt- 
arenas,  Costa  Rica. 

Type  Material. —  The  male  holotype, 
female  allotype,  and  10  paratypes  were 
taken  at  the  type  locality  on  ll-Vin-66, 
30  m.  No.  72,  from  a  leguminous  tree,  by 
me. 

The  holotype,  allotype,  and  paratypes 
are  in  my  collection. 

Scolytus  laetus,  n.  sp. 

This  species  is  distinguished  from  toru- 
lus    Wood    by    the    flattened    sternum    2 


26 


GREAT  BASIN  NATURALIST 


Vol.  35,  No.  1 


which  is  much  more  finely,  sparsely  punc- 
tured, with  longer,  more  abundant  hair, 
and  by  characters  of  the  frons  described 
below. 

Male. —  Length  2.4  mm  (paratypes 
2.4-2.5  mm),  2.4  times  as  long  as  wide; 
color  very  dark  brown. 

Frons  as  in  torulus  except  elevation 
wdder,  area  above  elevation  more  dis- 
tinctly, more  broadly  impressed,  vestiture 
on  lateral  margins  more  abundant,  much 
longer. 

Pronotum  and  elytra  as  in  torulus. 

Sternum  2  vertical,  almost  flat,  surface 
dull,  punctures  small,  not  clearly  evident, 
spine  as  in  torulus,  vestiture  much  longer, 
some  setae  as  long  as  spine. 

Female. —  Similar  to  male  except  frons 
as  in  female  torulus  but  with  vestiture 
slightly  more  abundant  and  longer,  par- 
ticularly in  lower  areas;  sternum  un- 
armed, spine  absent,  vestiture  finer,  more 
abundant. 

Type  Locality. —  Forty-eight  km  or 
30  miles  N  Rosamorada,  Nayarit,  Mexico. 

Type  Material. —  The  male  holotype, 
female  allotype,  and  three  male  para- 
types were  taken  at  the  type  locality  on 
15-Vn-65,  1000  m.  No.  257,  from  Inga 
paterno,  by  me. 

The  holotype,  allotype,  and  paratypes 
are  in  my  collection. 

Scolytodes  amabilis,  n.  sp. 

In  general  body  features,  this  species 
is  very  similar  to  clusiavorus  Wood,  but 
the  female  frons  is  much  more  similar  to 
volcanus  Wood.  It  is  distinguished  from 
those  species  as  indicated  below. 

Female. —  Length  1.6  mm  (paratypes 
L4-1.6  mm),  2.5  times  as  long  as  wide; 
color  very  dark  brown,  almost  black. 

Frons  essentially  convex,  with  a  very 
shallow,  subconcave  area  on  median  third 
just  below  upper  level  of  eyes,  this  im- 
pression continued  on  median  fifth  as  an 
abrupt,  shallow  sulcus  to  epistoma,  median 
half  of  lateral  areas  bordering  sulcus  with 
numerous  beadlike  granules,  remaining 
areas  somewhat  dull,  with  fine  punctures; 
surface  of  sulcus  shining,  almost  smooth; 
rather  sparse  vestiture  limited  to  margins, 
of  fine,  long  hair,  those  on  dorsal  margin 
attaining  level  of  antennal  insertion, 
shorter  laterally  and  below. 

Pronotum    and    elytral    outlines    as    in 


clusiacolens  Wood;  pronotum  surface  re- 
ticulate, punctures  as  in  clusiavorus;  strial 
and  interstrial  punctures  similar  to  but 
slightly  larger  than  in  clusiavorus.  Strial 
setae  almost  obsolete;  interstrial  setae  al- 
most obsolete  on  even-numbered  inter- 
striae,  fine,  rather  short,  and  widely 
spaced  on  odd-numbered  interstriae. 

Male. —  Similar  to  female  except  frons 
convex,  reticulate,  vestiture  very  sparse, 
short,  inconspicuous. 

Type  Locality. —  Mt.  Tzontehuitz, 
Chiapas,  Mexico. 

Type  Material.—  The  female  holo- 
type, male  allotype,  and  15  paratypes 
were  taken  at  the  type  locality  on  29-V- 
69,  at  3000  m,  by  D.  E.  Bright.  Twenty- 
one  paratypes  bear  the  same  data  except 
23-VI-69;  two  of  them  were  taken  from 
Quercus  sp. 

The  holotype,  allotype,  and  most  of  the 
paratypes  are  in  the  Canadian  National 
Collection;  the  remaining  paratypes  are 
in  my  collection. 

Scolytodes  obesus,  n.  sp. 

This  species  is  distinguished  from  the 
remotely  related  immanus  Wood  by  the 
smaller  size,  by  the  stouter  body  form, 
by  the  pair  of  carinae  on  the  female 
frons,  and  by  other  characters. 

Female. —  Length  1.8  mm,  2.1  times 
as  long  as  wide;  color  yellowish  brown, 
anterior  third  of  pronotum  darker. 

Frons  shallowly,  broadl}^  concave  from 
level  of  antennal  insertion  to  upper  level 
of  eyes  (upper  area  concealed  by  prono- 
tum), surface  smooth  and  shining  except 
subreticulate  near  margins;  epistomal  area 
from  margin  to  level  of  antennal  inser- 
tion longitudinally  divided  into  equal 
thirds  by  a  pair  of  rather  strongly  ele- 
vated carinae;  premandibular  epistomal 
lobe  large,  conspicuous,  pubescent;  vesti- 
ture mostly  confined  to  margins  of  upper 
half  of  frontal  area,  consisting  of  a  row  of 
long,  subplumose  setae,  longest  setae  equal 
to  about  one-half  to  two-thirds  distance 
between  eyes. 

Pronotum  0.97  times  as  long  as  wide; 
widest  near  base,  sides  weakly,  arcuately 
converging  on  basal  two-thirds,  rather 
broadly  rounded  in  front;  anterior  third 
moderately  declivous,  finely  asperate; 
posterior  areas  reticulate,  very  finely, 
rather    closely    punctured.     Glabrous    ex- 


March  1975 


WOOD:     AMERICAN    SCOLYTIDAE 


27 


cept  for  an  occasional  coarse  seta  on  as- 
perate area. 

Elytra  1.15  times  as  long  as  wide,  1.25 
times  as  long  as  pronotum;  sides  almost 
straight  on  basal  half,  slightly  wider  at 
base  of  declivity,  rather  broadl}'  rounded 
behind;  disc  confined  to  basal  half;  striae 
not  impressed,  punctures  small,  rather 
shallowly  impressed,  spaced  by  distances 
equal  to  diameter  of  a  puncture;  inter- 
striae  four  times  as  wide  as  striae,  smooth, 
shining,  punctures  small  to  minute,  weak- 
ly impressed;  interstriae  10  weakly  carin- 
ate  to  level  of  sternum  5.  Declivity  con- 
vex, moderately  steep;  sculpture  as  on 
disc.  Vestiture  of  sparse,  moderately  long 
bristles  on  odd-nvmibered  interstriae. 

Protibia  slender,  lacking  minute  tooth 
on  posterior  face  near  tarsal  insertion. 

Type  Locality. —  Barro  Colorado  Is- 
land, Panama  Canal  Zone. 

Type  Material. —  The  female  holo- 
type  was  taken  at  the  type  locality  on  7- 
VIII-67,  L.  and  C.  W.  O'Brien. 

The  holotype  is  in  my  collection. 

Scolytodes  lepidus,  n.  sp. 

This  species  is  distinguished  from  amoe- 
nus  Wood  by  the  slightly  larger  average 
size,  by  the  slightly  larger  elytral  punc- 
tures, by  the  presence  of  a  few  elytral 
setae,  particularly  along  sides,  and  by  the 
very  different  female  frons. 

Female. —  Length  2.0  mm  (paratypes 
1.8-2.3  mm),  2.2  times  as  long  as  wide; 
color  dark  brown,  base  of  pronotum  usu- 
ally pale. 

Frons  broadly  flattened  from  epistoma 
to  vertex,  almost  smooth,  upper  half  and 
sides  below  coarsely,  closely  punctured 
and  pubescent,  median  third  on  lower 
half  slightly  elevated,  smooth,  shining, 
impmictate,  glabrous;  vestiture  long, 
moderately  abundant,  more  widely  dis- 
tributed than  m  amoenus. 

Pronotum  1.0  times  as  long  as  wide;  as 
in  amoenus  except  discal  area  reticulate 
(smooth  to  subreticulate  in  amoenus)  and 
moderately  pubescent  at  lateral  margins 
(almost  glabrous  in  amoenus). 

Elytra  1.2  times  as  long  as  wide,  1.3 
times  as  long  as  pronotum;  as  in  amoenus 
except  strial  and  interstrial  punctures 
larger,  more  completely  confused  and 
lateral  areas  with  sparse  setae  (entirely 
glabrous  in  amoenus). 


Male. —  Similar  to  female,  with  frons 
similar  to  male  amoenus  except  more 
coarsely  punctured,  more  protuberant  in 
median  area,  with  no  gramiles. 

Type  Locality.^ —  Thirty-three  km  or 
21  miles  N  Juchitlan,  Jalisco,  Mexico. 

Type  Material. —  The  female  holo- 
type, male  allotype,  and  25  paratypes 
were  taken  at  the  type  locality  on  3-Vn- 
65,  at  1300  m.  No.  177,  from  Ficus,  by 
me,  from  the  same  branches  that  con- 
tamed  the  type  series  of  amoenus. 

The  holotype,  allotype,  and  paratypes 
are  in  my  collection. 

Scolytodes  genialis,  n.  sp. 

This  species  is  distinguished  from  lepi- 
dus Wood  by  the  absence  of  pmictures  in 
the  asperate  area  of  the  jDronotum  and  by 
the  much  finer  punctures  on  the  pronotal 
disc  and  on  the  elytra. 

Female. —  Length  1.8  mm  (paratypes 
1.8-2.1  mm),  2.3  times  as  long  as  wide; 
color  light  brown,  anterior  half  of  prono- 
tLun  darker. 

Frons  as  in  lepidus  except  vestiture  fin- 
er, very  slightly  shorter.  Pronotmn  as  in 
lepidus  except  punctures  very  fine,  shal- 
low. Elytra  as  in  lepidus  except  strial 
punctures  fine,  shallow,  in  definite  rows, 
interstrial  punctures  very  small,  confused, 
striae  1  not  impressed  on  declivity,  vesti- 
ture on  sides  of  elytra  minute. 

Type  Locality. —  Laguna  Santa  Maria, 
Nayarit,  Mexico. 

Type  Material.- —  The  female  holo- 
type, male  allotype,  and  six  paratypes 
were  taken  at  the  type  locality  on  7-VII- 
65,  at  900  m.  No.  197,  irom  Ficus  with 
yellow  bark,  by  me.  Four  paratypes  are 
from  24  km  or  15  miles  S  Mazamitla, 
Jalisco,  Mexico,  22-VL65,  2500  m,  No. 
97,  Ficus  with  yellow  bark,  by  me. 

The  holotype,  allotype,  and  paratypes 
are  in  my  collection. 

Pseudot/iysanoes  concentralis,  n.sp. 

This  species  is  distinguished  from  quer- 
cinus  Wood  by  the  concentric,  carinate 
pronotal  asperities,  by  the  more  strongly 
impressed  Irons,  by  the  greatly  reduced 
elytral  punctures,  and  by  the  wider  elyt- 
ral scales. 

Female. —  Length  1.0  mm  (paratypes 


28 


GREAT  BASIN  NATURALIST 


Vol.  35,  No.  1 


1.0-1.2  mm),  2.6  times  as  long  as  wide; 
color  dark  brown. 

Frons  rather  strongly  concave  on  medi- 
an two-thirds  of  area  below  upper  level 
of  eyes,  surface  subrugose,  with  sparse 
granules  except  almost  smooth  on  lower 
half  of  concavity;  vestiture  short,  sparse. 
Antennal  scape  as  wide  as  long,  as  long 
as  pedicel,  bearing  a  small  tuft  of  long 
hair;  club  about  as  in  quercinus. 

Pronotum  0.73  times  as  long  as  wide; 
outline  as  in  quercinus;  simimit  at  mid- 
dle; anterior  slope  on  median  third  armed 
by  six  concentric,  uniform,  transverse 
carinae,  other  asperities  absent,  carina  1 
submarginal,  6  at  summit;  posterior  areas 
shining,  almost  smooth,  punctures  al- 
most obsolete.  Vestiture  consisting  of  a 
row  of  scales  posterior  to  each  of  first  five 
carinae,  and  rather  sparse,  coarse  hair 
in  remaining  areas. 

Elytra  1.8  times  as  long  as  wide,  2.4 
times  as  long  as  pronotum;  outline  as  in 
quercinus;  striae  not  impressed,  punc- 
tures small,  shallow;  interstriae  almost 
smooth,  about  twice  as  wide  as  striae, 
punctures  almost  obsolete;  surface  usually 
covered  by  a  thin  incrustation.  Declivity 
convex,  steep,  sculpture  as  on  disc  but 
punctures  even  more  obscure.  Vestiture 
mostly  abraded,  consisting  of  widely 
spaced,  short,  interstrial  scales,  each  scale 
one  and  one-half  to  two  times  as  long  as 
wide. 

Male. —  Length  0.8  mm;  similar  to 
female  except  smaller,  slightly  stouter; 
frontal  concavit}'  not  as  deep;  pronotal 
asperities  not  fused  or  clearly  concentric. 

Type  Locality.- —  Ten  km  or  6  miles 
SE  Totolapan,  Oaxaca,  Mexico. 

Type  Material. —  The  female  holo- 
type,  male  allotype,  and  24  paratypes 
were  taken  at  the  type  locality  on  21 -VI- 
67,  1000  m.  No.  73,  from  a  leguminous 
roadside  shrub  with  a  yellow  flower  that 
was  presumed  to  be  Cassia  sp.,  by  me. 

The  holot^^pe,  allotype,  and  paratypes 
are  in  my  collection. 

Pseudothysanoes  tumidulus,  n.  sp. 

This  species  is  distinguished  from  the 
distantly  allied  graniticus  Wood  by  the 
different  female  frons,  by  the  smaller  ely- 
tral  scales,  and  by  the  very  different  male 
declivity  as  described  below. 

Male. —    Length    1.7    mm    (paratypes 


1.5-2.0  mm),  2.2  times  as  long  as  wide; 
color  very  dark  brown. 

Frons  flattened  on  lower  two-thirds, 
with  small  concavity  on  median  fifth, 
ascending  slightly  to  epistomal  margin, 
convex  above;  subshining  and  almost 
smooth  on  flattened  area,  more  coarsely 
punctate-granulate  above;  vestiture  con- 
fined to  epistomal  area  and  to  convex  area. 

Pronotimi  similar  to  graniticus  except 
anterior  margin  narrowly  rounded  and 
armed  by  six  teeth  and  posterior  areas 
rather  strongly  reticulate. 

Elytra  1.3  times  as  long  as  wide;  sides 
straight  and  parallel  on  basal  three- 
fourths,  rather  abruptly  rounded,  then 
broadly  rounded  behind;  striae  not  im- 
pressed, punctures  small,  moderately  deep, 
spaced  by  distances  equal  to  diameter  of 
a  puncture;  interstriae  three  times  as  wide 
as  striae,  almost  smooth,  punctures  min- 
ute, granulate,  uniseriate  except  confused 
near  declivity.  Declivity  with  basal  mar- 
gin abrupt,  basically  convex  except  upper 
half  flattened  to  striae  4,  a  moderate  bul- 
la just  below  middle  from  interstriae  2-4; 
strial  punctLu-es  smaller  and  not  as  deep 
as  on  disc,  closer,  in  indistinct  rows,  of 
same  size  and  shape  as  confused  interstrial 
granules,  interstrial  punctures  obsolete; 
bulla  covered  by  same  surface  sculpture 
as  elsewhere.  Vestiture  of  interstrial  rows 
of  scales,  each  slightly  longer  than  wide 
on  disc  except  scales  confused,  more 
abundant,  much  longer,  and  more  slender 
at  base  of  declivity;  declivity  glabrous; 
rows  of  fine,  recumbent  strial  hair  on 
disc. 

Female. —  Similar  to  male  except  frons 
broadly,  shallowly  concave  from  epistoma 
to  well  above  eyes,  vestiture  on  upper  area 
slightly  more  abundant  (less  abundant 
and  shorter  than  in  graniticus) ;  anterior 
margin  of  pronotum  more  broadly  round- 
ed, unarmed;  declivity  convex,  sculpture 
as  on  disc  except  strial  punctures  obso- 
lete; rows  of  interstrial  scales  continued 
to  apex,  each  scale  equal  in  length  to 
three-fourths  distance  between  rows,  more 
closely  spaced  within  a  row,  each  about 
three  to  four  times  as  long  as  wide. 

Type  Locality. —  Highway  120,  129 
km  NE  San  Juan  del  Rio,  Queretaro, 
Mexico. 

Type  Material. —  The  male  holotype, 
female  allotype,  and  27  paratypes  were 
taken  at  the  type  locality  on  9-VL71,  at 


March  1975 


WOOD:     AMERICAN    SCOLYTIDAE 


29 


2500  m,  from  mistletoe  on  oak,  b}^  D.  E. 
Bright. 

The  holotype,  allotype,  and  most  of  the 
paratypes  are  in  the  Canadian  National 
Collection;  the  remaining  paratypes  are 
in  my  collection. 

Thysanoes  tuberculatus,  n.  sp 

This  species  is  distingLiished  from  gran- 
ulifer  Wood  by  the  smaller  size  and  by 
the  ver}"  different  el}^tral  declivit}^  as 
described  below. 

Male. —  Length  1.8  mm,  2.7  times  as 
long  as  mde;  color  rather  dark  yellowish 
brown. 

Frons  and  pronotum  as  in  granulifer 
and  berchemiae  Blackman. 

Elytra  1.6  times  as  long  as  wide;  disc 
as  in  granulifer  except  granules  slightly 
larger.  Declivity  steep,  convex;  striae  1 
and  2  with  punctures  minute,  visible  al- 
most to  apex;  suture  slightly  elevated,  with 
a  row  of  small  granules  on  basal  half;  in- 
terstriae  2  with  two  or  three  tubercles  on 
less  than  basal  fourth,  broadly  impressed 
below  and  entirely  devoid  of  punctures 
and  granules;  interstriae  3  moderately  el- 
evated on  middle  third  and  anned  wdth 
a  row  of  six  to  eight  rather  coarse  tuber- 
cles, lower  third  without  punctures  or 
granules;  lateral  interstriae  each  with  a 
row  of  tubercles  on  basal  area  but  none 
of  them  attaining  normal  apex  for  these 
interstriae.  Vestiture  of  rows  of  interstrial 
scales,  scales  on  disc  largely  abraded,  lit- 
tle if  any  longer  than  wide,  longer  at  base 
of  declivity;  those  on  3  up  to  three  times 
as  long  as  on  disc  and  four  times  as  long 
as  wide,  slightly  shorter  on  other  inter- 
striae; declivital  interstriae  2,  4,  lower 
two-thirds  of  1,  and  lower  half  of  3  glab- 
rous. 

Type  Locality. —  Eighty-five  km  or 
53  miles  S  Valle  Nacional,  Oaxaca,  Mexi- 
co. 

Type  Material. —  The  male  holotype 
was  taken  at  the  type  locality  on  24-V-71, 
at  3300  m,  D.  E.  Bright. 

The  holotype  is  in  the  Canadian  Na- 
tional Collection. 

Pityokteines  mystacinus,  n.  sp. 

This  species  is  distinguished  from  minu- 
tus  (Swaine)  b}^  the  smaller  size,  by  the 
very  different  ornamentation  of  hair  on 
the   female   frons,   by   the   finer  pronotal 


and  elytral  pmictures,  and  by  other  char- 
acters. It  probably  is  much  more  closely 
related  to  lasiocarpi  (Swaine). 

Female. —  Length  2.1  mm  (paratypes 
1.7-2.1  mm),  3.0  times  as  long  as  wide; 
color  brown. 

Frons  similar  to  lasiocarpi  except  more 
broadly  convex,  surface  not  as  smooth, 
somewhat  dull,  punctures  average  smaller 
and,  on  lower  third,  becoming  almost  ob- 
solete; vestiture  on  lower  third  abundant, 
rather  long,  epistomal  brush  very  broad, 
rather  dense;  setae  on  upper  half  of  fron- 
tal area  sparse,  short.  Antennal  club  al- 
most as  in  lasiocarpi. 

Pronotum  as  in  lasiocarpi  except  anteri- 
or margin  more  narrowly  rounded;  vesti- 
ture uniformly  short  as  in  lasiocarpi.  Ely- 
tra as  in  lasiocarpi  except  punctures  on 
disc  slightly  larger,  very  slightly  more 
confused  on  basal  half,  and  declivital 
striae  1  more  strongly  impressed,  with 
punctures  on  striae  1  slightly  larger,  sub- 
apical  transverse  elevation  at  apex  of  sul- 
cus more  distinct  (but  still  rather  ob- 
scure) ;  position,  number,  and  size  of  tub- 
ercles as  in  lasiocarpi.  Vestiture  similar 
in  abundance,  but  ver}^  slightly  longer 
than  a  lasiocarni. 

Male. —  Similar  to  female  except  upper 
half  of  frons  more  strongly  convex,  vesti- 
ture on  lower  third  greatly  reduced  in 
abundance  and  length;  elytral  declivity 
with  sulcus  slightly  deeper  (about  as  in 
male  lasiocarpi)   but  narrower. 

Type  Locality. —  Mount  Rainier  Na- 
tional Park,  Washington. 

Type  Material. —  The  female  holo- 
type, male  allotype,  and  three  female 
paratypes  were  taken  at  the  tvpe  localitv 
on  21-Vin-62,  silver  fir,  D.  E.  Bright.  ^ 

The  holotype,  allotype,  and  one  para- 
type  are  in  the  Canadian  National  Col- 
lection and  two  paratypes  are  in  my  col- 
lection. 

Ips  pilifrons  thc/tcheii,  n.  subsp. 

This  geographical  race  of  pilifrons  is 
distinguished  from  p.  pilifrons  Swaine,  of 
northern  Colorado,  b}-  the  characters  of 
the  frons  described  below.  Both  subspecies 
are  replaced  throughout  all  of  Utah  by  p. 
utahensis  Wood  in  which  frontal  charac- 
ters are  entirely  different.  The  following 
comparisons  are  based  on  females  having 
the  maximum  frontal  elevation  and  piles- 


30 


GREAT  BASIN  NATURALIST 


Vol.  35,  No.  1 


ity  for  their  race;  occasional  specimens  of 
p.  pilifrons  almost  overlap  the  maximum 
development  of  p.  tJiatcheri. 

Female. —  Length  4.6  mm  (parat3^pes 
3.9-4.9  mm),  2.5  times  as  long  as  wide; 
color  very  dark  brown. 

Frons  similar  to  p.  pilifrons  except  less 
strongly,  less  extensively  elevated,  eleva- 
tion occupying  lower  80  percent  of  median 
distance  from  epistoma  to  upper  level  of 
eyes  (115  percent  in  p.  pilifrons)  and 
pubescent  area  occupying  less  than  50 
percent  of  median  area  between  eyes  (80 
percent  in  p.  pilifrons) ;  lateral  areas  much 
more  sparsely,  more  finely  granulate  than 
in  p.  pilifrons.  Pronotum,  elytra,  and 
other  features  essentially  as  in  p.  pilifrons. 

Male. —  Similar  to  male  p.  pilifrons 
except  frontal  vestiture  less  abundant  and 
shorter,    particularly    along   epistoma. 

Type  Locality. —  Mt.  Wheeler,  Neva- 
da. 

Type  Material. —  The  female  holo- 
type,  male  allotype,  and  48  paratypes 
were  taken  at  the  type  locality  on  10- 
Vin-74,  at  10,000  ft.,  from  Picea  engel- 
manni^  by  me. 

The  holotype,  allotype,  and  paratypes 
are  in  my  collection. 

This  subspecies  is  named  for  Dr.  T.  O. 
Thatcher  who  discovered  it  more  than  30 
years  ago. 

Araptus  attenuatus,  n.  sp. 

This  species  is  distinguished  from  placa- 
tus  Wood  by  the  much  coarser  strial 
punctures,  by  the  much  longer  female 
frontal  vestiture,  and  by  other  characters 
cited  below. 

Female. — •  Length  1.6  mm  (paratypes 
1.4-1.6  mm),  3.0  times  as  long  as  wide; 
color  dark  brown. 

Frons  feebl}'  convex,  flattened  on  medi- 
an half  in  some  specimens;  subshining, 
rather  finely,  closely  punctured  in  periph- 
eral areas,  central  area  minutely  irregular, 
often  with  a  few  punctures;  a  weak  medi- 
an carina  on  lower  half;  vestiture  of 
fine,  long,  white,  subplumose  setae  in 
punctured  area  at  sides  and  above,  lon- 
gest setae  equal  to  two-thirds  diameter  of 
frons,   shorter  toward   epistoma. 

Pronotum  1.2  times  as  long  as  wide;  as 
in  placatus  except  anterior  margin  more 
narrowly  rounded,  posterior  areas  more 
distinctly    reticulate,    wdth    punctures    al- 


most twdce  as  large;  minute  setae  present 
on  disc. 

Elytra  1.8  times  as  long  as  wide,  1.6 
times  as  long  as  pronotum;  similar  to 
placatus  except  strial  punctures  almost 
twice  as  large,  declivity  more  narrowly 
convex,  interstriae  2  not  impressed,  1 
more  feebly  elevated.  Minute  strial  setae 
visible  on  posterior  half  of  disc  and  decliv- 
ity except  1  and  2,  interstrial  setae  con- 
fined to  declivity,  absent  on  2,  similar  to 
placatus  except  usually  finer. 

Male. —  Similar  to  female  except  frons 
more  distinctly  convex,  punctures  subru- 
gose,  obscurely  acicutate,  a  distinct,  sub- 
tuberculate,  median  prominence  at  upper 
level  of  eyes  and  continuing  toward  ver- 
tex, its  summit  transversely  etched. 

Type  Locality. —  Forty-eight  km  or  30 
miles  W  Bajia  de  los  Angeles,  Baja  Cali- 
fornia Norte,  Mexico. 

Type  Material. —  The  female  holo- 
type, male  allotype,  and  four  paratypes 
were  taken  at  the  type  locality  on  Sl-IIL 
74,  Hopk.  58650,  by  M.  M.  Furniss,  from 
host  plant  No.  88.  Twenty-two  paratypes 
are  from  32  km  or  20  miles  N  Punta 
Prieta,  Baja  California  Norte,  Mexico,  29- 
in-73,  Pedialanthus  niacrocarpus,  J.  Doy- 
en. 

The  holotype,  allotype,  and  several  para- 
types are  in  my  collection;  two  paratypes 
are  in  the  Canadian  National  Collection; 
the  remaining  paratypes  are  in  the  Uni- 
versity of  California  (Berkeley)  Collec- 
tion. 

Araptus  fossifrons,  n.  sp. 

This  common  species  was  thought  to  be 
foveifrons  (Schedl)  until  the  type  of 
Schedl's  species  became  available  for 
study;  however,  this  species  has  the  elj'tral 
punctures  much  finer  and  the  declivital 
sulcus  much  more  weakly  impressed. 

Male. —  Length  2.2  mm  (paratypes 
1.8-2.4  mm),  2.5  times  as  long  as  wide; 
color  rather  dark  reddish  brown. 

Frons  as  in  foveifrons  except  lower  area 
more  finely  punctured.  Elytral  disc  with 
punctures  much  finer,  interstrial  punc- 
tures usually  more  widely  spaced  (vari- 
able). Elytral  declivity  with  striae  1 
rather  strongly  impressed,  interstriae  2 
not  impressed,  with  a  row  of  punctures. 
Vestiture  as  in  foveifrons. 

Female. —  Similar  to  male  except  frons 
weakly    convex,    a    weak    median    carina 


March  1975 


WOOD:     AMERICAN    SCOLYTIDAE 


31 


from  vertex  to  epistoma,  surface  closely, 
rather  coarsely  punctured,  with  fme 
abundant,  moderately  long  hair  imiformly 
distributed  from  epistoma  to  above  eyes; 
frons  concealed  in  only  available  female 
of  foveifrons. 

Type  Locality. —  Lago  Amatitlan, 
Guatemala. 

Type  Material. —  The  male  holotype, 
female  allotype,  and  48  paratypes  were 
taken  at  the  type  locality  on  lO-VI-64, 
700  m,  No.  702,  from  the  fruiting  body  of 
a  climbing  (Cucurbitaceae)  vine,  by  me. 
Other  parat^'pes  were  taken  in  Mexico  as 
follows:  9  at  3  km  (2  miles)  SE  Acatlan, 
Puebla,  15-VI-67,  1500  m,  No.  37;  17  at 
9  km  (12  miles)  SE  Oaxaca,  Oaxaca,  18- 
VI-67,  No.  57;  4  from  24  km  (15  miles) 
W  Armeria,  Colima,  30-VI-65,  30  m,  No. 
141;  9  from  1  km  N  Atenquique,  Jalisco, 
24-VI-65,  1000  m.  No.  115;  2  from  8  km 
(5  miles  S  Atenquique,  Jalisco,  25-VI-65, 
1000  m.  No.  115A;  3  from  8  km  (5  miles) 
W  Juchitlan,  Jalisco,  2-VII-65,  1000  m. 
No.  174;  14  from  24  km  (15  miles)  S 
Mazamitla,  Jahsco,  22-VI-65,  2500  m. 
No.  96;  13  from  Tuxpan,  JaHsco,  23- VI- 
65,  1300  m,  No.  99;  3  from  8  km  (5  miles) 
N  Ruiz,  Nayarit,  14-Vn-65,  100  m.  No. 
245;  all  from  fruiting  pods  of  climbing 
vines  or  a  small  tree,  by  me. 

The  holotype,  allotype,  and  paratypes 
are  in  mv  collection. 


Amphicranus  parilis.  n.  sp. 

This  species  is  distinguished  from  //'//- 
formis  Blandford  by  the  much  smaller 
size,  by  the  absence  of  minute  crenula- 
tions  on  the  base  of  the  pronotal  disc,  and 
by  differences  in  the  elytral  declivity  in- 
dicated below. 

Male. —  Length  2.0  mm,  4.0  times  as 
long  as  wide;  color  pals  yellowish  brown, 
elytral  declivity  brown. 

Frons  and  pronotum  as  in  filiformis  ex- 
cept as  noted  in  diagnosis.  Elytra  as  in 
filiformis  except  less  strongly  explanate, 
sutural  emargination  only  slightly  deeper 
than  wide  (twice  as  deep  as  wide  in  fili- 
formis), declivital  spine  2  smaller,  less 
strongly  pointed. 

Type  Locality. —  Six  km  or  4  miles  W 
Tepic,  Nayarit,  Mexico. 

Type  Material. —  The  male  holotype 
was  taken  at  the  type  locality  on  1  3-VIL 


65,  1000  m.  No.  240,  from  a  tree  branch, 
by  me. 

The  holot}pe  is  in  my  collection. 

Cor  thy  III  s  cecropii,  n.  sp. 

This  abberrant  species  is  distinguished 
from  all  other  species  in  the  genus  by  the 
small  antennal  club,  without  sutures,  by 
the  absence  of  female  pronotal  asperities, 
and  by  the  minute  to  obsolete  punctures 
of  the  pronotum  and  elytra. 

Female. —  Length  2.4  mm  (paratypes 
2.0-2.5  mm),  2.3  times  as  long  as  wide; 
color  black. 

Frons  uniformh%  deeply  concave  from 
eye  to  eye,  from  epistoma  to  vertex;  sur- 
face densely,  imiformly,  very  finely  punc- 
tured over  entire  surface;  vestiture  very 
fine,  rather  abundant,  uniformly  rather 
short  over  concave  area,  margin  above 
eyes  with  a  dense  row  of  Aery  long  hair, 
a  small  tuft  of  longer  hair  at  level  of  an- 
tennal insertion  on  lateral  half.  Anten- 
nal club  1.6  times  as  long  as  wide;  asym- 
metrically obovate,  aseptate,  entire  surface 
minutely  pubescent;  posterior  face  with 
a  small  tuft  of  hair  extending  about  half 
of  club  length  beyond  apex. 

Pronotum  1.1  times'  as  long  as  wide; 
sides  weakly  arcuate  on  posterior  half, 
broadly  rounded  in  front,  a  distinct,  sub- 
marginal,  transverse  constriction;  anterior 
margin  unarmed;  summit  indefinite,  near 
middle;  asperities  absent;  surface  reticu- 
late, anterior  half  with  sparse,  minute 
granules,  ])OSterior  half  with  sparse  min- 
ute punctures.  Acute  lateral  margins 
more  strongly  developed  than  in  other 
species.  Glabrous. 

Elytra  1.3  times  as  long  as  wide,  1.2 
times  as  long  as  pronotum;  sides  almost 
straight  and  parallel  on  basal  two-thirds, 
obtusely  subangulate  behind;  disc  reticu- 
late, a  few  obscure,  irregular  lines  indi- 
cated, punctures  minute,  mostly  obsolete, 
apparently  confused.  Declivity  occupying 
slightly  more  than  posterior  third,  rather 
steep,  convex;  sculpture  as  on  disc,  except 
a  few  irregularly  placed  fine  granules 
usually  present.  Vestiture  confined  to  de- 
clivity, consisting  of  sparse,  short,  fine 
bristles  apparently  on  odd-numbered  inter- 
striae. 

Male. —  Similar  to  female  except  frons 
convex,  a  distinct,  transverse  impression 
above  epistoma,  surface  smooth,  shining. 


32 


GREAT  BASIN  NATURALIST 


Vol.  35,  No.  1 


impunctate;  anterior  margin  of  pronotum 
distinctly  produced  toward  median  line 
and  armed  b}'  two  slender  teeth;  anterior 
slope  of  pronotum  much  steeper,  asperate. 

Type  Locality. —  Tapanti,  Cartago, 
Costa  Rica. 

Type  Material. —  The  female  holo- 
type,  male  allotype,  and  15  paratypes 
were  taken  at  the  type  locality  on  24-X- 
63,  1300  m.  No.  242",  from  fallen  Cecropia 
peltata  petioles,  by  me.    Eight  paratypes 


are  from  Turrialba,  Cartago,  Costa  Rica, 
5-VII-63,  700  m.  No.  19,  Cecropia  petioles, 
by  me.  Three  paratypes  are  from  6  km 
S  San  Vito,  Puntarenas,  13-III-68,  H. 
Ilespenheide.  Three  specimens  not  in- 
cluded in  the  type  series  are  from  El 
Laurel  (Experiment  Station),  12  km  SW 
Caracas,  Venezuela,  l-V-70,  1300  m.  No. 
475,  Cecropia  petioles,  by  me. 

The  holotype,   allotype,   and  paratypes 
are  in  my  collection. 


GENETICS,   ENVIRONMENT,   AND   SUBSPECIES   DIFFERENCES: 
THE  CASE  OF  POLITES  SABULETl 

(Lepidoptera:    Hesperiidae) 

Arthur  M.  Shapiro^ 

Abstract. —  Polites  sabuleii  is  an  example  of  an  insect  having  a  univohine,  monophenic  high- 
elevation  subspecies  and  a  multivoltine  lowland  one  that  produces  similar  phenotypes  only  in  cold 
weather.  When  reared  under  conditions  that  induce  the  warm-weather  phenotype  in  lowland  stocks, 
the  montane  subspecies  P.  s.  tecumseh  continues  to  produce  its  usual  phenotype,  indicating  that  it  has 
become  genetically  fixed. 


One  variant  of  the  persistent  "genetics- 
environment"  duality  in  biology  concerns 
the  nature  of  subspecies  differences.  The 
problem,  as  it  applies  to  butterflies,  was 
well  simimarized  in  Klots's  (1951)  dis- 
cussion of  geographic  variation: 

To  what  degree  much  of  the  recorded  geo- 
graphic variation  is  a  matter  of  tempera- 
ture and  humidity  differences  is  something 
which  we  can  only  infer.  In  Papilio  glau- 
cus  .  .  .  spring  specimens  tend  to  be  small 
and  pale.  .  .  .  As  we  go  northward  we  find 
that  in  central  Canada,  where  there  is  only 
one  generation  a  year,  the  whole  popula- 
tion looks  similarly  small  and  pale.  In  Can- 
ada this  population  has  been  named  as  a 
geographic  subspecies,  ''canadensis,'''  i.e.  a 
part  of  the  species  limited  to  a  certain  area 
and  showing  distinctive  characteristics.  The 
temptation  is  strong  to  attribute  the  whole 
thing  to  lowered  temperatures  alone.  But 
suppose  we  brought  a  batch  of  eggs  of 
canadensis  down  to  Florida,  and  reared  the 
butterflies  in  the  conditions  under  which 
the  very  large,  richly  colored  subspecies 
australis  develops  there.  Would  our  cana- 
densis eggs  develop  as  australis  ...  or  would 
they  develop  into  the  same  small  pale 
specimens  that  their  parents  were? 

Twenty-three  years  later  Ehrlich,  Holm, 
and  Parnell  (1974)  could  only  write  that 

many  butterflies  have  spring  generations 
that  are  smaller  and  darker  than  their 
summer  generations,  the  difference  pre- 
sumably being  due  to  the  seasonal  variation 
in  the  environment.  However,  in  some 
northern  parts  of  their  range,  [they]  have 
only  a  single  summer  generation,  which 
is  small  and  dark  and  resembles  the  spring 
generation  of  southern  localities.  In  the 
northern  populations,  the  individuals  are 
presumed  to  have  genotypes  that  produce 
the  dwarfing  and  darkening.  Although  the 
critical  transfer  experiments  have  not  been 
done,  the  greater  constancy  of  the  northern 
forms  in  the  face  of  environmental  changes 
supports  these  presumptions  [emphasis 
added] . 

The  same  problem  was  recognized  in 


plants  as  far  back  as  the  1920s,  in  the  clas- 
sic work  of  Turesson  (1922,  1925,  1929) 
later  brilliantly  expanded  by  Clausen, 
Keck,  and  Hiesey  (1940,  1947,  1948,  and 
other  papers).  This  work  firmly  estab- 
lished the  concept  of  the  ecotype  in  plant 
ecology  and  genetics,  a  concept  more  or 
less  readily  generalizable  to  animals  in 
cases  like  those  discussed  by  Klots  and 
Ehrlich  et  al.  Turesson  and  Clausen  et  al. 
were  able,  by  transplant  experiments,  to 
separate  phenotypic  variation  produced 
directly  by  the  physical  environment  from 
that  produced  indirectly  through  the  se- 
lection of  climaticall}'  adapted  genotypes. 
This  paper  is  the  second  of  a  series  re- 
porting on  analogous  studies  of  North 
American  Lepidoptera. 

The  Subspecies  of  Polites  sabuleti 

Situations  of  the  sort  described  above 
are  not  limited  to  populations  separated 
by  latitude;  many  Lepidopterans — like  the 
]3lants  studied  by  Clausen  and  his  col- 
leagues— have  altitudinal  variants,  often 
described  taxonomically  at  the  subspecies 
level,  and  these  are  especially  interesting 
because  of  the  short  ground  distances  be- 
tween the  high-  and  low-elevation  popula- 
tions and  the  possibility  of  investigating 
the  nature  of  their  contacts,  if  any. 

Polites  sabuleti  Boisduval  is  a  small, 
largely  tawny  skipper  (Hesperiidae), 
widely  distributed  in  western  North 
America.  Three  named  subspecies  occur 
in  California:  P.  s.  sabuleti,  P.  s.  tecumseh 
Grinnell,  and  P.  s.  chusca  Edwards.  The 
last  is  a  very  pale  desert  population  and 
has  not  been  examined  in  this  study.  P.  s. 
sabuleti  and  P.  s.  tecumseh  are  parapatric 
in  northern  and  central  California,  occur- 
ring at  low  and  high  elevations  respec- 
tively. 


'Department   of  Zoology,   University  of  California,   Davis,   California    95616. 


33 


34 


GREAT  BASIN  NATURALIST 


Vol.  35,  No.  1 


P.  s.  sabuleti  is  very  widespread  on 
sandy  soils,  in  saline  and  alkaline  marsh- 
lands, and  in  urban  vacant  lots.  It  is 
usually  closely  associated  with  its  normal 
larval  host,  alkali  grass  {DisticJilis  spicata 
[L.]  Greene,  Gramineae),  but  may  breed 
occasionally  on  Bermuda  grass  (Cynodon 
dactylon  [L.]  Pers.),  an  introduced  lawn 
grass  and  weed.  P.  s.  sabuleti  is  strongly 
multivoltine  at  sea  level,  with  possibly  as 
many  as  five  generations  a  year.  The 
flight  season  is  Aery  long,  ranging  from 
late  March-April  to  mid-  or  late  Novem- 
ber at  Sacramento  and  Suisun  City,  Cali- 
fornia. 

P.  s.  tecumseh  occurs  in  subalpine  mea- 
dows that  become  dry  in  summer,  and  in 
alpine  fell-fields  in  the  high  Sierra  Neva- 
da. Its  host  plant  is  not  known,  but  many 
collectors  have  noted  an  association  with 
species  of  sedges  {Carex^  Cyperaceae). 
Tilden  (1959)  reports  P.  s.  tecumseh  fly- 
ing from  Jul}'  to  September  in  Yosemite 
National  Park,  which  he  interprets  as  in- 
dicating two  broods.  The  more  complete 
data  given  in  Garth  and  Tilden  (1963) 
do  not  support  .this  interpretation,  as  there 
is  no  wide  spread  of  dates  within  a  given 
year  at  a  single  locality.  Farther  north, 
at  Donner  Pass  (7,000  feet)  there  is  no 
indication  of  more  than  one  brood.  Em- 
mel  and  Emmel  (1962)  found  it  there 
from  19  June  to  19  August,  1960;  Sha- 
piro found  it  at  the  same  localities  from  1 1 
July  to  24  August,  1973,  and  18  July  to 
12  August,  1974.  The  condition  of  Don- 
ner Pass  specimens  does  not  suggest  even 
a  partial  second  brood.  Like  many  mon- 
tane butterflies,  P.  s.  tecumseh  emerges 
later  at  higher  elevations;  thus,  at  10,000 
feet  it  flies  mainly  in  August  and  into 
ver}'^  early   September. 

As  with  the  altitudinal  subspecies  of 
Phyciodes  campestris  Behr  (N^mphali- 
dae)  previously  studied  (Shapiro,  1975a), 
those  of  Polites  sabuleti  are  separated  by 
a  zone  in  which  neither  seems  to  occur. 
At  the  latitude  of  Sacramento,  P.  s.  sabu- 
leti is  unknown  as  a  breeding  resident 
above  1,500  feet  and  P.  s.  tecumseh  is  un- 
recorded below  5,000  feet.  Tilden  (1959) 
confuses  the  matter  by  indicating  that  P. 
s.  sabuleti  extends  much  higher  at  Yo- 
semite, but  his  data  (given  in  Garth  and 
Tilden,  1963)  make  it  plain  that  this  re- 
fers to  the  arid  east  slope  only:  the  rec- 
ords are  from  Bridgeport  (6,743  ft.)  and 
Mono  Lake  (6,419  ft.).   On  the  west  slope 


the  lowest  Yosemite-area  record  of  P.  s. 
tecumseh  is  Gin  Flat  (7,036  ft.)  and  there 
are  no  records  of  P.  s.  sabuleti  at  all  (al- 
though it  is  abundant  on  the  floor  of  the 
San  Joaquin  Valley).  The  nature  of  the 
east-slope  contacts  has  not  been  studied, 
but  few  habitats  suitable  for  either  sub- 
species occur  on  the  abrupt  Sierran  es- 
carpment. 

Polites  sabuleti  tecumseh  differs  from 
summer  P.  s.  sabuleti  in  being  smaller, 
hairier,  and  more  heavily  marked,  especi- 
ally on  the  hindwing  ventrally.  The  dark 
markings  on  this  wing  are  often  described 
as  being  a  "colder,"  grayer  color  than  in 
P.  s.  sabuleti.  Although  a  series  of  23 
tecumseh  from  the  vicinity  of  Donner 
Pass  shows  considerable  variation,  no  sea- 
sonal pattern  is  apparent.  In  P.  s.  sabuleti 
from  Sacramento  and  Suisun  City  (over 
450  specimens  examined)  there  is  marked 
seasonal  variation:  March-May  and  Sep- 
tember-November specimens  are,  on  the 
average,  smaller,  darker,  and  hairier  than 
summer  ones,  and  some  are  superficially 
exceedingly  similar  to  P.  s.  tecumseh,  al- 
though there  are  minor  (  but  fairl}^  con- 
sistent) differences  in  certain  details  of 
the  pattern.  The  phenotypes  of  wild 
specimens  of  both  taxa  are  illustrated  in 
Figures   1-3. 

Experimental    Methods    and    Results 

Would  stock  of  P.  s.  tecumseh  reared 
under  conditions  that  produce  summer 
phenotypes  of  P.  s.  sabuleti  produce  the 
normal  tecumseh  phenotype,  or  would  it 
be  modified  in  the  direction  of  the  low- 
land, summer  one?  Ova  were  obtained 
from  a  female  tecu?nseh  collected  at  Don- 
ner Pass  (7,000  ft),  24  July  1974,  and 
from  two  female  sabuleti  collected  in  a 
salt  marsh  at  Suisun  City,  Solano  County 
(10  ft.),  6  August  1974.  The  resulting 
progeny  were  reared  side  by  side  in 
plastic  Petri  dishes  (51/2"  diameter  X  %") 
at  comparable  densities  (5-8  larvae/dish) 
under  continuous  illumination  from  a  60w 
bulb  at  25C  (77F).  All  larvae  were  fed 
fresh  cuttings  of  Bermuda  grass  (Cynodon 
dactylon),  and  mortality  in  both  stocks 
was  negligible.  Thirty  adult  P.  s.  sabuleti 
(16  cf  14  ?  )  and  thirteen  P.  s.  tecumseh 
(8  cf  5  ?  )  were  obtained.  Continuous 
light  was  selected  as  a  regime  ecologically 
nonsignificant  to  both  stocks  but  known 


March  1975 


SHAPIRO:    POLITES  SABULETI 


35 


(i-,-.:^  fei*:^ 


\K,M£AJ 


'Ls^mJ' 

4                        ,     * 

Fig.  1.     Dorsal    and   ventral    surfaces  of   summer   specimens   of  Polites  sabuleti  sabuleti  from   the 
Central   Valley   of   California. 


■4/  ^T 


Fig.  2.     Dorsal  and  ventral  surfaces  of  early  spring   and  late  fall  P.  s.  sabuleti  from  the  Central 
Valley,  approaching  the  phenotype  of  P.  s.  tecumseh. 


•f'''.^:; 


Fig.  3.     Dorsal  and  ventral  surfaces  of  P.  s.  tecumseh   from    Donner   Pass,    California    (7,000    feet), 
July-August. 


36 


GREAT  BASIN  NATURALIST 


Vol.  35,  No.  1 


to  inhibit  diapause  in  P.  sahuleti  and  re- 
lated species. 

The  two  stocks  differed  in  several  re- 
spects in  the  laboratory.  First-  and  second- 
instar  larvae  oi  P.  s.  sabuleti  were  yellow- 
ish green;  in  the  third  instar  they  turned 
purplish  brown;  and  thereafter  they  re- 
mained that  color.  P.  s.  tecumseh  larvae 
were  purplish  brown  throughout  their  de- 
velopment. At  corresponding  points  in 
the  life  cycle  the  early  stages  of  P.  s.  sabu- 
leti were  always  larger  than  their  high- 
altitude  counterparts.  The  developmental 
rates  of  the  two  stocks  differed  very  sig- 
nificantly, with  little  overlap:  from  egg  to 
adult  P.  s.  sabuleti  took  39-76  days 
(weighted  mean,  58.5  days)  and  P.  s. 
tecumseh,  70-111  days  (weighted  mean, 
86.0).   No  diapause  was  observed  in  either 


stock.  Normally,  high-elevation  or  -lati- 
tude stocks  of  Lepidoptera  develop  more 
rapidly  than  conspecific  ones  from  more 
temperate  climates  when  reared  under 
uniform  laboratory  conditions;  the  rever- 
sal of  this  situation  in  Polites  sabuleti  is 
to  my  knowledge  unique  in  Lepido])teran 
stocks  in  which  diapause  is  not  manifested 
in  culture. 

The  developmental  differences  noted 
above  were  not  mirrored  in  larval  be- 
havior or  morphology.  The  adults,  how- 
ever, were  obviously  different  and  "true" 
to  their  normal  phenotypes  (Figs.  4,  5): 
nondiapaused  tecumseh  reared  at  high 
temjieratures  retained  all  of  their  distin- 
guishing characters,  including  size.  It  thus 
appears  that  the  complex  of  characters 
present  as  a  developmental  option  in  low- 


n 


Fig.  4.     Dorsal  and  ventral  surfaces  of  representative  bred  Polites  sabuleti  sabiileti;  continuous  light, 
25C. 


Fig.  5.     Dorsal  and  ventral  surfaces  of  representative  bred  P.   s.    tecumseh;   same  conditions  as  in 
Figure  4. 


March  1975 


SHAPIRO:    POLITES  SABULETI 


37 


land,  multivoltine  populations  is  geneti- 
call}'  fixed  in  P.  s.  tecumsch,  confirming 
Ehrlich  et  al.'s  prediction. 

Discussion 

Three  sets  of  "altitudinal  subspecies" 
have  now  been  investigated  in  butterflies, 
representing  three  different  and  quite  un- 
related families.  They  are  Picris  occident- 
alis  Reakirt  and  its  alpine  representative, 
familiarly  (but  incorrectly)  known  as  P. 
o.  "calyce^'  Edwards  (Pieridae);  PJiycio- 
des  campcstris  Behr  and  its  montane  sub- 
species montana  Behr  (Nymphalidae) ; 
and  Politcs  sahidcti.  The  first  two  are  dis- 
cussed at  length  in  Shapiro  (1975a).  Each 
species  presents  a  jiicture  different  from 
the  others. 

Pieris  occidentalis  shows  xery  little,  if 
any,  genetic  differentiation  of  the  univol- 
tine,  monophenic  and  bivoltine,  diphenic 
populations  of  high  and  moderate  eleva- 
tions, respectively.  The  alpine  stock  re- 
tains the  ability  to  produce  an  estival 
phenot}'pe  and  to  develop  without  dia- 
pause, and  its  own  phenotype  is  indis- 
tinguishable from  the  vernal  one  produced 
downslope.  Their  mating  behavior  in- 
volves male  aggregations  on  mountain- 
tops,  a  behavior  pattern  conducive  to  gene 
flow,  and  in  laboratory  experiments  no 
reproductive  barriers  have  been  found  be- 
tween uni-  and  bivoltine  populations. 

Phyciodes  campestris  shows  a  superfi- 
cially similar  picture,  in  that  the  high  al- 
titude subspecies  montana,  when  reared 
under  outdoor  conditions  at  sea  level,  pro- 
duces the  foothill  phenotype  rather  than 
its  own.  However,  the  cold-season  pheno- 
type of  lowland  populations  is  quite  dif- 
ferent from  montana,  and  the  ability  to 
produce  the  montana  phenotype  appears 
to  be  restricted  to  high-elevation  popula- 
tions. In  this  case,  then,  phenotypic  plas- 
ticity is  not  reciprocal,  and  the  high- 
elevation  population  is  genetically  dif- 
ferentiated. 

Polites  sabuleti  shows  the  highest  de- 
gree of  differentiation  yet  encountered. 
The  possibility  that  it  has  achieved  repro- 
ductive isolation  (i.e.,  speciation)  between 
high-  and  low-elevation  populations  can- 
not be  discounted.  Because  of  the  differ- 
ence in  developmental  time,  no  crosses 
between  the  stocks  could  be  made.  They 
are  not  known  to  intergrade  anywhere, 
since  the  altitudinal  discontinuity  i)etween 


them  seems  to  rim  the  length  of  the  Sier- 
ra Nevada. 

These  experiments  have  been  duplicated 
with  a  latitudinal  subspecies  pair — Cali- 
fornia Pieris  occidentalis  and  its  subspe- 
cies P.  o.  nelsoni  Edwards  from  Fairbanks, 
Alaska  (Shapiro,  1975b).  Their  pheno- 
typic differences  are  clearly  heritable,  and 
the  res]:)onse  of  both  phenot}'pe  and  dia- 
pause to  photoperiod  has  been  observed  in 
Fi  and  F:.  hybrids. 

Studies  of  the  Pieris  occidentalis  and  P. 
napi  L.  species  complexes  (Shapiro,  1975 
c)  strongly  imply  that  univoltinism  is 
evolutionarily  derivative  from  multivoltin- 
ism,  accompanying  the  successful  invasion 
of  increasingly  rigorous  climates.  High- 
altitude  and  -latitude  populations  of  wide- 
spread species  are  probably  derived  from 
lowland  sources,  as  has  been  well  docu- 
mented for  the  Sierran  alj^ine  flora  (Cha- 
bot  and  Billings,  1972).  The  overall  pic- 
ture emerging  from  these  studies  supports 
the  suggestion  that  seasonal  ])henotypes 
of  multivoltine  populations  may  become 
fixed  through  selection  of  modifiers  in- 
fluencing thresholds  of  develo]:)mental  ex- 
pression ("genetic  assimilation,"  Wad- 
dington,  1953).  The  somewhat  unusual 
circumstances  in  Phyciodes  campestris 
montana  will  be  explored  in  another  pa- 
per. 

Acknowledgments 

Collection  of  livestock  for  this  study  was 
funded  by  Grant  D-804  from  the  Com- 
mittee on  Research  of  the  Academic  Sen- 
ate, U.  C.  Davis.  Steven  R.  Sims,  John 
H.  Lane,  and  Adriejine  R.  Shapiro  assisted 
in  field  work.  Aspects  of  this  research 
benefited  from  conversations  from  Mich- 
ael Rosenzweig,  William  E.  Bradshaw, 
and  E.  W.  Jameson,  Jr. 


LiTER' 


Ci 


Chabot,  B.  F.,  and  W.  D.  Billings.  1972.  Ori- 
gins and  ecology  of  the  Sierran  alpine  flora 
and  vegetation.  Ecol.  Monographs  42:163- 
199. 

Clausen,  J.,  D.  D.  Keck,  and  W.  Hiesey.  1940. 
Experimental  studies  on  the  nature  of  spe- 
cies. I.  Effects  of  varied  environments  on 
western  North  American  plants.  Carnegie 
Inst.  Wash.,  publ.  520. 

.      1947.     Heredity    of    geographically    and 

ecologically  isolated  races.  Amer.  Nat.  81: 
114-133. 

.  1948.  Experimental  studies  on  the  na- 
ture of  species.  III.  Environmental  responses 


38 


GREAT  BASIN  NATURALIST 


Vol.  35,  No.  1 


of  climatic  races  of  Achillea.  Carnegie  Inst. 
Wash.,  publ.  581. 

Ehrlich,  p.  R.,  R.  W.  Holm,  and  D.  R.  Parnell. 
1974.  The  process  of  evolution,  2d  ed.  Mc- 
Graw-Hill, New  York.    378  pp. 

Emmel,  T.  C,  and  J.  F.  Emmel.  1962.  Eco- 
logical studies  of  Rhopalocera  in  a  high  Sier- 
ran  community — Donner  Pass,  California.  I. 
Butterfly  associations  and  distributional  fac- 
tors. J.  Lepid.  Soc.  16:23-44. 

Garth,  J.  S.  and  J.  W.  Tilden.  1963.  Yosem- 
ite  Butterflies.  J.  Res.  Lepid.  2:1-96. 

Klots,  a.  B.  1951.  A  field  guide  to  the  but- 
terflies of  North  America,  east  of  the  Great 
Plains.     Houghton   Mifflin,    Boston.     349   pp. 

Shapiro,  A.  M.  1975a.  Ecotypic  variation  in 
montane  butterflies.  Wasmann  J.  Biol.,  in 
press. 


.  1975b.  Photoperiodic  control  of  devel- 
opment and  phenotype  in  a  subarctic  popula- 
tion of  Pieris  occidcntalis  (Lepidoptera:  Pi- 
eridae).    Canad.    Entomol.,   in   press. 

.     1975c.     Developmental    and    phenotypic 

responses  to  photoperiod  in  uni-  and  bivoltine 
Pieris  napi  (Lepidoptera:  Pieridae)  in  Cali- 
fornia. Trans.  Roy.  Ent.  Soc.  London,  in  press. 

Tilden,  J.  W.  1959.  The  butterfly  associations 
of  Tioga  Pass.  Wasmann  J.  Biol.  17:249- 
271. 

TuRESsoN,  G.  1922.  The  species  and  the  variety 
as    ecological    units.    Hcreditas    3:100-113. 

.     1925.     The   plant   species   in   relation   to 

habitat  and   climate.   Hereditas  6:147-236. 

.      1929.     Zur   Natur  und   Begrenzung  der 

Artenlieiten.     Hereditas     12:323-334. 

Waddington,  C.  H.  1953.  Epigenetics  and  evo- 
lution.  Symp.   Soc.   Exptl.   Biology   7:186-199. 


LIFE  HISTORY  AND  ECOLOGY  OF  MEGARCYS  SIGNATA 

(PLECOPTERA:    PERLODIDAE),   MILL   CREEK,   WASATCH 

MOUNTAINS,   UlAH^ 

Mary  R.  Gather-  and  Arden  R.  Gaufin- 

Abstract. —  During  an  investigation  of  some  of  the  stoneflies  (Plecoptera)  of  Mill  Creek,  Wasatch 
Mountains,  Utah,  Megarcys  signata,  a  large  omnivorous  stonefly,  was  found  to  have  a  univoltine  life 
history  and  a  slow  seasonal  life  cycle. 

Temperature  appears  to  affect  the  growth  rate  of  Megarcys  signata.  Warmer  stream  temperatures 
accompany  the  acceleration  of  the  growth  rate,  whereas  cooler  stream  temperatures  apparently  retard 
the  growth  rate. 

Periods  of  maximum  absolute  growth  rate  con-espond  with  maximum  carnivorous  feeding  from 
August  to  September  and  March  to  April.  Ghironomidae,  Ephemeroptera,  and  Plecoptera,  in  that  order, 
were  the  most  abundant  prey  in  the  foreguts.  Young  n^-mphs  ingested  considerable  amounts  of  diatoms, 
filamentous  algae,  and  detritus  but  not  as  much  animal  matter  as  did  older  nymphs. 

Megarcys  signata  was  unifonnly  distributed  throughout  Mill  Creek,  except  at  the  lowest  station, 
where  few  nymphs  were  found. 

Emergence  occurred  in  May  and  June,  the  peak  occurring  in  June.  The  mean  size  of  females 
and  males  decreased  as  emergence  progressed. 


This  report  is  part  of  an  eighteen-month 
study  of  some  of  the  stoneflies  of  Mill 
Creek,  Wasatch  Mountains,  Utah.  Be- 
cause a  detailed  description  of  the  study 
area  and  the  methods  and  materials  is 
given  in  another  paper  (Cather  and  Gau- 
fin  1975,  only  a  summary  is  included 
here. 

Mill  Creek  Canyon  is  located  11  km 
southeast  of  Salt  Lake  City,  Utah,  in  the 
Wasatch  Mountains  of  the  Middle  Rocky 
Mountain  Province.  Six  stations  were  se- 
lected along  a  12  km  length  of  the  stream 
in  the  Wasatch  National  Forest  with  ele- 
vations ranging  from  1,605  to  2,280  m. 
The  sampling  stations  were  nimibered 
consecutively.  Station  I  denoting  the  high- 
est elevation.  The  three  lower  stations 
(1,605-1,785  m)  are  easily  accessible  all 
year,  but  the  three  upper  stations  (1,995- 
2,280  m)  are  accessible  only  in  the  simi- 
mer  and  fall.  Average  minimum  and 
maximum  daily  flows  were  0.3  m^/sec 
and  1.2  mVsec,  respectively,  during  the 
study  period.  Depth  averaged  11-45  cm, 
and  current  averaged  0.2-0.7  m/sec  during 
the  fall  when  measurements  were  taken. 
The  substrate  of  the  sampling  area  at  all 
stations  ranged  from  coarse  sand  to  small 
cobbles.  Minimum  water  and  air  tempera- 
tures recorded  during  adult  emergence 
were  3  C  and  9  C,  respectively.  Maximum 
water  and  air  temperatures  during  this 
period  were  13  C  and  26  C,  respectively. 
Water  chemistry  was  similar  at  all  sta- 
tions. Dissolved  oxygen  ranged  from  6.0 
to  8.5  mg/1   (70-120  percent  saturation). 


calcium  bicarbonate  109-189  mg/1,  calci- 
vmi  carbonate  0-2.4  mg/1,  pH  7.5-8.3, 
total  hardness  100-340  mg/1,  and  conduc- 
tivity 312-859  mhos/cm. 

Methods  and  Materials 

Nymphs  of  Megarcy  signata  were  col- 
lected at  least  monthly  from  June  1971  to 
December  1972  at  each  of  six  stations. 
Additional  nymphs  were  collected  in 
spring  1973  for  food  habit  studies.  Hand- 
screens  of  mesh  sizes  7  and  9  sq/cm  were 
used,  the  smaller  handscreen  being  used 
during  the  majority  of  the  study  in  an 
attempt  to  collect  the  smaller  instars.  An 
area  of  about  80  cm-  of  the  stream  bottom 
was  disturbed  in  an  attempt  to  collect  at 
least  100  nymphs  monthly.  All  nymphs 
were  preserved  in  80  percent  ethanol. 

Adults  were  collected  weekly  through- 
out the  emergence  period  and  biweekly 
during  peak  emergence  using  a  sweep  net 
and  handpicking  from  vegetation,  rocks, 
and  bridges.  All  adults  were  preserved  in 
80  percent  ethanol. 

The  interocular  distance  of  all  nymphs 
and  adults  was  measured  to  the  nearest 
0.1  mm  using  an  ocular  micrometer  in  a 
dissecting  microscope  for  determining 
growth  rates  and  to  see  if  the  mean  size  of 
the  adults  decreased  as  emergence  pro- 
gressed. The  nymphs  were  identified  as 
males  and  females  when  possible. 

Foregut  analyses  were  conducted  on  200 
nymphs  collected  from  the  field.  Nymphs 
were  selected  from  an  upper   (I)    and  a 


^Study  supported  by  Environmental  Protection  Agency  Traineesliip  Grant  No.  5T2-WP-542-03. 
^Department  of  Biology,   University  of  Utah,  Salt  Lake  City. 


39 


40 


GREAT  BASIN  NATURALIST 


Vol.  35,  No.  1 


lower  (IV)  station  during  each  season 
(Station  V  had  to  be  substituted  for  Sta- 
tion I  during  winter  and  spring).  The 
method  used  followed  Swapp  (1972).  All 
prey  animals  were  enumerated  and  identi- 
fied to  order  except  where  family  or 
generic  determination  was  possible.  Three 
types  of  the  family  Chironomidae  were 
recognized  and  designated  as  species  a,  b, 
and  c  (based  on  morphology  of  head 
capsule).  These  are  discussed  under  re- 
sults and  discussion  but  are  not  separated 
in  Table  1.  All  algae  were  determined  to 
genus  where  possible.  Percentage  compo- 
sitions of  algae  and  detritus  were  esti- 
mated when  present,  and  dominant  items 
were  recorded.  A  volume  analysis  was 
not  conducted. 

Identification  of  nymphs  and  adults  fol- 


lowed Gaufin  et  al.    (1966),  and  nomen- 
clature followed  lilies   (1966)  and  Zwick 

(1973). 

Results  and  Discussion 

The  only  systellognathan  stonefly  pres- 
ent in  Mill  Creek  in  numbers  large  enough 
for  analysis  is  Megarcys  signata.  This 
species  exhibits  a  slow  seasonal  type  of 
life  cycle.  Emergence  and  oviposition  oc- 
cur in  May  and  June  with  hatching  soon 
after.  Small  nymphs  (0.5  mm  interocular 
distance)  appear  in  July  at  the  lower  sta- 
tions. Nymphs  of  comparable  size  gen- 
erally appear  for  the  first  time  in  August, 
September,  and  October  at  the  upstream 
stations.  Rapid  growth  occurs  from  Aug- 
ust to  emergence  (Figs,  la,  b,  c).  The  size 


lOOn 


75 


50- 


25 


2.5- 

-.  2.0- 

E 

o 

].o^ 

0.5 


Month 


Fig.  1.  Growth  of  Megarcys  signata.  Arrows  indicate  emergence.  I.  O.  indicates  interocular  dis- 
tance: (a)  Monthly  mean  size  as  a  percentage  of  total  mean  size  at  Station  IV;  (b)  Monthly  mean 
cumulative  growth  at  Station  IV;  (c)  Monthly  mean  absolute  growth  rate  (data  pooled  from  all  sta- 
tions).  A   indicates  change  in   I.   O.   distance. 


March  1975 


GATHER,    GAUFIN:     UTAH    PLECOPTERA 


41 


frequencies  of  nymphs  and  the  mean 
cumulative  growth  at  each  station  are 
shown  in  Figs.  2  and  3,  respectively. 

A  comparison  of  the  cumulative  growth 
at  all  stations  reveals  that  the  most  rapid 
growth  occurs  at  the  lowest  (warmest) 
stations  (Fig.  4).  Thus,  there  seems  to  be 
a  direct  correlation  between  growth  and 
temperature.  Baumann  (1967)  found  no 
direct  correlation  in  this  species  in  Mill 
Creek.  Seasonally  the  most  rapid  growth 
occurs  during  the  fall  and  early  winter 


(September- January).  Growth  apparently 
slows,  but  does  not  stop,  during  the  winter 
(January-March),  increases  from  March 
to  April,  and  then  decreases  from  April  to 
May,  prior  to  emergence.  Sheldon  (1972) 
reported  similar  results  in  his  study  on  the 
Arcynopteryx  species  complex  he  studied 
in  California.  However,  Schwarz  (1970) 
reported  no  growth  at  times  during  the 
winter  in  other  Systellognatha.  The  corre- 
lation between  growth  and  food  habits  will 
be  discussed  later. 


2.0' 
101 


II        J 

O       1,0 


2.0- 

1.0- 


103     2 

A 


33 


•l  'h 


34 


27 


4 

3        0 


50 


J         A        30        ND         J         F         MAM        J         J         ASO 

Month 


"n       ^ 


Sfa  2.0-|       b 

IV 

1.0-1 


1    2.0- 


O     1.0- 


10' 


AC  15        9        1 

38    33       <^h°^n  r   ^^ 


ly       ,3        1/ 


?       cf: 


J         A        S        5        S         5        ]  F        M        A        ?ir 

Month 


9         6 


10     5    'l'^ 


2        3 

I     ' 

50 

5     6     N     F 


Fig.  2.  [Frequency  distribution  of  nymphal  size  classes  of  M.  signata.  Number  of  individuals 
shown  above  each  polygon;  males  and  females  as  indicated:  (a)  Stations  I,  II,  and  III;  (b)  Stations 
IV,  V,  and  VI. 


42 


GREAT  BASIN  NATURALIST 


Vol.  35,  No.  1 


Megarcys  signata  seems  to  show  no 
preference  for  either  the  upstream  or 
downstream  stations.  This  species  is  even- 
ly distributed  throughout  the  stream  ex- 
cept at  Station  VI.  Here  the  substrate  is 
almost  entirely  cemented.  Where  the 
water  is  deep  enough  for  this  large  insect, 
the  current  is  too  slow;  where  the  current 
is  fast  enough,  the  water  is  too  shallow 
and  the  substrate  too  homogeneous.  More 
individuals  were  collected  at  Station  IV 
than  at  any  other  station. 


The  emergence  of  M.  signata  began  in 
early  May  at  the  lower  stations  and  lasted 
until  late  June  at  the  higher  stations  (Fig. 
5).  Peak  emergence  was  in  June.  Bau- 
mann  (1967)  found  this  species  emerg- 
ing from  late  April  to  mid- July  in  Mill 
Creek.  Emergence  is  progressively  later 
as  the  elevation  increases.  Baumann 
(1967),  Hynes  (1970),  and  Nebeker 
(1971)  reported  similar  results.  They 
were  first  collected  at  Stations  III  and  V 
in  early  May,  when  the  water  tempera- 


Sta 


Sta 


2.0- 


O      ).0- 


2.0- 


2.0- 
1.0- 


o 
-  la 


--HH 


J         ASONDJ  FMAMJ         JASOND 

Month 


T~y 


~r~T 


T— r 


Fig.  3.  Mean  cumulative  growth  of  nymphs  of  M.  signata.  Vertical  line  represents  size  range  of 
nymphs;  shaded  area  represents  standard  deviation;  unshaded  area  represents  standard  eiTor  of  the 
mean;  solid  line  connects  means;  rectangle  represents  emergence  period:  (a)  Stations  I,  II,  and  III; 
(b)  Stations  IV,  V,  and  VI. 


March  1975 


GATHER,    GAUFIN:     UTAH    PLECOPTERA 


43 


J   a 


1 5" 


^^ 


Fig.  4.  Comparison  of  mean  cumulative 
growth  of  nymphs  of  (D)  M.  signata  at  Stations 
I-V.  Station  I  represented  by  (o);  Station  II  by 
(•);  Station  III  by  (■);  Station  IV  by  (A); 
Station  V  by  (A);  and  Station  VI  by  (D):  (a) 
July,  1971  through  June,  1972;  (b)  July,  1972 
through  December,  1972. 

tures  were  3  C  and  5  C,  respectively. 
Emergence  ended  in  late  May  (water  tem- 
perature 9  C)  at  Station  V,  but  lasted  until 
mid- June  (12  C)  at  Station  IV  and  late 
June  (5.5-7  C)  at  the  three  upper  stations. 
Megarcys  signata  is  a  secretive  insect  that 
hides  in  cracks  under  bridges  or  among 
vegetation  to  escape  warm  summer  tem- 
peratures. Clusters  of  these  stoneflies 
usually  containing  one  female  and  several 
males  were  often  collected  in  these  hiding 
places.  Brinck  (1949)  reported  the  same 
phenomenon  in  related  species.  The  col- 
lection data  probably  reflect  this  secretive 
habit  in  that  M.  signata  should  have  been 
collected  earlier  at  Station  IV. 

Females  and  males  generally  emerged 
together  in  a  1:1  ratio.  Harper  and  Pilon 
(1970)  reported  similar  findings.   Only  55 


100- 

0 

? 

100 

0 

d" 

100 

0 

^^        9 

rf- 

100- 

0 

■  d- 

100- 

0- 

<f 

100 

a                  .« 

100 

100' 

20      30        10      20      30 


Fig.  5.     Emergence  at  each   station  of  M.   sig- 
nata. 

females  and  49  males  were  collected.  Fe- 
males outnumbered  males  in  both  May 
and  June  (Fig.  6).  There  was  no  size 
overlap  between  the  females  and  males. 
The  mean  size  of  females  and  males  de- 
creased as  emergence  progressed  from  May 
to  June  at  all  stations  (data  pooled).  Khoo 
(1968),    Schwarz     (1970),    and    Sheldon 


50 

0- 

— 

39 
2<f 

50- 

0 

—- 

2d' 

50- 

0- 

.Is 

2^ 

^^r^-Jl? 

^<J 

38<f 

50- 

0 

— 

4d- 

50 

0 

__35 

50 

0 

!? 

50 

|. 

2,0      2  2      2.4      2  6  18      2.0      2.2      2.4      2,6 

I.O  (mm) 


Fig.  6.  Frequency  distribution  of  adults  of 
M.  signata.  Number  of  individuals  shown  above 
each  polygon. 


44 


GREAT  BASIN  NATURALIST 


Vol.  35,  No.  1 


(1972)  reported  a  similar  phenomenon. 
The  mean  size  of  females  decreased  from 
2.4  mm  to  2.3  nun,  while  that  of  males 
decreased  from  1.9  mm  to  1.8  mm  (Fig. 
7).  The  data  were  rather  inconclusive 
because  only  small  numbers  of  adults  were 
collected  in  May.  More  intensive  collect- 
ing may  reveal  this  trend  more  strongly. 
Sheldon  (1972)  reported  similar  results  in 
the  Arcynopteryx  species  complex  he 
studied  and  suggested  that  the  decrease  in 
mean  size  also  influenced  fecundity;  that 
is,  the  smaller  females  carried  fewer  eggs. 
Warmer  stream  temperatures  and  in- 
creasing photoperiod  may  act  as  emer- 
gence cues  before  growth  and  egg  develop- 
ment are  completed  (Khoo  1968,  Hynes 
1970). 

A  total  of  200  nymphs  of  M.  signata 
were  dissected  and  the  foreguts  examined 
for  a  preliminary  food-habit  analysis.  The 


nymphs  were  selected  from  upper  and 
lower  stations  and  from  each  season  of  the 
year.  Table  1  gives  the  results  of  this 
analysis.  Of  those  foreguts  examined,  39 
were  empty  (some  of  these  stoneflies  were 
beginning  to  molt;  others  had  just 
molted).  Chironomids  seemed  to  be  the 
preferred  food  item.  Three  types  of  chi- 
ronomids based  on  differences  in  the  head 
capsules  were  recognized.  Of  these  three 
types,  one  was  much  more  frequently 
found  in  the  foreguts.  A  total  of  442  were 
found  in  39  percent  of  the  insects  ex- 
amined. The  other  two  types  were  found 
only  occasionally.  Mayflies  were  the  sec- 
ond most  abundant  food  item  ingested  (37 
percent  of  all  individuals).  Baetis  spp. 
were  most  frequently  recognized.  Stone- 
flies,  notably  chloroperlids,  comprised  a 
considerable  portion  of  the  gut  contents, 
also  (27  percent  of  the  individuals).   Rich- 


T.\BLE    1.     Percentage   of   dissected    nymphs    of  Megarcys  signata  containing  specific  food  items. 


Season 

Summer 

Fall 

Winter 

Spring 

Date 

7/20/72 

2/4/72 

9/8/72 

10/15/71 

2/18/72 

3/17/73 

Station  No. 

I 

I 

V 

V 

Head  Capsule  Size  Range 

for  Sample    (mm) 

0.4-1.1 

0.5-1.4 

1.7-2.6 

1.9-2.5t 

No.  Foreguts  Examined 

25 

25 

19 

31 

Empty  Foreguts  (%) 

12 

8 

26 

13 

Class  Insecta 

Ephemeroptera 

5 

0 

21 

63 

Plecoptera 

9 

4 

29 

49 

Trichoptera 

4 

0 

0 

3 

Diptera 

Chironomidae 

32 

17 

79 

26 

Other  Diptera 

0 

0 

0 

0 

Unidentified 

0 

0 

7 

7 

Division  Cyanophyta 

Oscillatoria   (?)    spp. 

4;9* 

9 

0 

0 

Division  Chlorophyta 

0 

0 

0 

Mougeotia   spp. 

0 

30;35* 

14;21* 

0 

Enteromorpha  spp. 

18;14 

Desmids 

Closterium   spp. 

9 

4 

0 

0 

Division   Chiysophyta 

Diatoms 

Navicula  spp. 

100 

69;17* 

71;21 

22 

Gomphonema  spp. 

18 

26 

29;14* 

0 

Cymbella   spp. 

64 

39 

0 

0 

Fragilaria    spp. 

4 

22;4* 

0 

0 

Nitzschia  spp. 

14 

•43 

0 

0 

Synedra  spp. 

0 

9 

0 

0 

Surirella  spp. 

23 

0 

0 

0 

Diatnma  spp. 

0 

0 

0 

0 

Misc.   diatoms 

27 

48 

0 

0 

Unidentified  sp.   1 

81  ;9* 

35;65* 

43  ;7* 

0 

Unidentified  sp.  2 

50 

0 

7 

0 

Unidentified  filamentous  algae 

14 

0 

0 

0 

Unidentified   plant   fragments 

0 

0 

0 

0 

Detritus  (sand  grains,  silt. 

plant  remains,  diatom  frustules) 

4;77* 

4;26* 

43;29* 

85 

f Measurements  from  3/17/72 
•Dominant 


March  1975                          gather,  gaufin: 

UTAH    PLECOPTERA 

45 

Table  1    (Continued) 

Season 

Summer 

Fall 

Winter 

Spring 

Date 

8/3/72 

8/11/71 

10/8/71 

3/12/73 

9/1/71 

10/15/71 

1/7/72 

3/17/73 

Station  No. 

IV 

IV 

IV 

IV 

X 

Head  Capsule  Size  Range 

for  Sample    (mm) 

0.5-1.2 

1.0-1.5 

1.4-2.3 

1.8-2.6+ 

No.  Foreguts  Examined 

25 

25 

25 

25 

Empty  Foreguts    (%) 

12 

36 

36 

16 

20 

Class  Insecta 

Ephemeroptera 

58 

19 

75 

48 

37 

Plecoptera 

9 

19 

81 

29 

27 

Trichoptera 

0 

0 

13 

0 

2 

Diptera 

Chironomidae 

45 

4 

88 

81 

44 

Other  Diptera 

0 

0 

13 

0 

1 

Unidentified 

18 

13 

4 

4 

7 

Division  Cyanophyta 

Oscillator ia   (?)    spp. 

0 

38* 

0 

0 

2;5* 

Division  Chlorophyta 

Mougeotia  spp. 

23 

0 

0 

0 

3 

Enter omorpha  spp. 

0 

0 

0 

0 

8;9* 

Desmids 

Closterium  spp. 

4 

4 

0 

0 

3 

Division  Chrysophyta 

Diatoms 

Navicula  spp. 

0 

'75* 

63* 

0 

33;22* 

Gomphonema   spp. 

0 

50 

38 

0 

20;2* 

Cymbclla    spp. 

0 

4 

0 

0 

15 

Fragilaria  spp. 

0 

0 

13 

4 

6;0.5* 

Nitzschia  spp. 

0 

0 

0 

0 

8 

Synedra  spp. 

0 

0 

0 

0 

1 

Surirella  spp. 

0 

0 

0 

0 

3 

Diatoma  spp. 

0 

0 

13 

0 

1 

Misc.    diatoms 

0 

0 

0 

19 

12 

Unidentified  sp.   1 

0 

19 

31 

0 

25;11* 

Unidentified  sp.  2 

0 

0 

0 

0 

7 

Unidentified   filamentous    algae 

0 

0 

0 

0 

2 

Unidentified  plant  fragments 

4 

0 

0 

19 

3 

Detritus  (sand  gi-ains,  silt, 

plant  remains,  diatom  fiiistules) 

14 

63  ;4* 

0 

19 

30;17* 

f Measurements  from  3/17/72 
*  Dominant 


ardson  and  Gaufin  (1971)  determined 
that  M.  signata  fed  primarily  on  Ephe- 
meroptera,  Chironomidae,  and  Simuliidae. 
In  his  study,  Swapp  (1972)  found  that 
the  similar  species,  M.  suhtruncata  (Han- 
son) and  Skivala  parallela  (Frison),  in- 
gested mayflies  and  chironomids  and  that 
S.  parallela  ingested  caddisflies  as  well. 
However,  he  found  only  one  S.  parallela 
of  200  specimens  that  contained  other 
stoneflies.  The  food  habits  of  M.  signata 
are  similar  to  those  of  Skwala  curvata 
(Hanson)  as  investigated  by  Sheldon 
(1972),  with  the  exception  of  the  number 
of  Trichoptera  eaten.  In  Mill  Creek  M. 
signata  ingested  very  few  caddisflies. 

Filamentous  algae,  diatoms,  and  detritus 
also  composed  a  significant  percentage  of 


the  gut  contents.  Richardson  and  Gaufin 
(1971),  Sheldon  (1972),  and  Swapp 
(1972)  report  that  all  the  Isogeninae  spe- 
cies they  studied  are  herbivores  to  a  con- 
siderable extent.  Hynes  (1941)  and 
Brinck  (1949)  agree  that  so-called  car- 
nivorous species  also  feed  on  vegetable 
matter.  There  is  no  way  to  determine  if 
these  items  are  actively  ingested,  present 
in  the  prey  stomachs,  or  eaten  in  the 
search  for  prey.  The  corollary,  that  the 
prey  species  may  have  been  ingested  while 
M.  signata  were  grazing  on  periphyton, 
may  also  be  true  (Sheldon  1972).  Navi- 
cula spp.  and  an  unidentified  species  of  a 
filamentous  diatom  were  the  most  nu- 
merous of  the  algae  found  in  the  guts. 
Detritus  in  the  form  of  sand  grains,  silt. 


46 


GREAT  BASIN  NATURALIST 


Vol.  35,  No.  1 


Month 

Fig.   7.     Size  range  of  adults  of  M.  signata. 

diatom  frustules,  leaf  fragments,  and  other 
plant  remains  were  found  in  30  percent  of 
the  insects  examined.  Other  diatoms  en- 
countered were  Gomphonema  spp.,  Cym- 
bella  spp.,  and  Nitzschia  spp.  These  are 
common  stream  dwellers  of  Mill  Creek, 
occurring  on  the  rocks  in  shallow  water. 

There  are  some  significant  seasonal  dif- 
ferences in  the  food  items  found  in  the 
guts.  In  the  summer  (July-early  Septem- 
ber) significantly  more  M.  signata  (58 
percent)  ingested  mayflies  than  in  the  fall 
at  Station  IV,  but  at  Station  I  the  num- 
bers feeding  on  Ephemeroptera  were  low 
(5  percent).  Stoneflies  were  found  in  only 
9  percent  of  the  guts  examined  at  Stations 
I  and  IV.  Chironomids,  on  the  other  hand, 
were  present  in  32  and  45  percent  of  the 
guts  at  Stations  I  and  IV,  respectively. 
The  number  of  chironomid  head  capsules 
found  was  also  greater  than  in  the  fall. 
At  Station  IV  diatoms  were  notably  lack- 
ing in  the  gut  contents,  although  mayflies 
and  chironomids  were  present.  This  could 
be  because  the  swift  current  dislodges  the 
algae  from  the  substrate  and  because  the 
deeper  water  and  suspended  solids  from 
spring  runoff  shut  out  sufficient  light. 
However,  at  Station  I,  where  the  water 
was  shallower,  diatoms  were  found  in  the 
majority  of  the  guts  examined   {Navicula 


spp.  were  found  in  all  stoneflies  ex- 
amined). Filamentous  algae  and  detritus 
were  also  present  in  many  guts.  Entero- 
morpha  spp.,  a  filamentous  green  algae, 
was  the  dominant  plant  material  in  14  per- 
cent of  the  guts,  while  detritus  was  the 
dominant  item  in  77  percent  of  the  guts. 
This  herbivorous  material  may  have  come 
from  the  stomachs  of  the  chironomid  prey 
(mayflies  present  in  only  5  percent  of  the 
stoneflies)  since  Oliver  (1971)  states  that 
some  chironomids  feed  on  algae  and 
detritus.  However,  many  guts  contained 
detritus  as  the  dominant  material  without 
chironomids  being  present.  It  is  conceiv- 
able that  during  the  summer  the  newly 
hatched  nymphs  of  M.  signata  could  feed 
at  least  partially  on  diatoms,  filamentous 
algae,  and  detritus.  At  this  stage  they  are 
probably  not  as  effective  a  predator  as  at 
later  life  stages.  Coffman,  Cimmiins,  and 
Wuycheck  (1971)  found  a  similar  pattern 
in  other  groups  of  insects,  in  that  young 
individuals  consumed  primarily  detritus 
but  shifted  to  algal  or  animal  ingestion  as 
they  matured.  The  lowest  absolute  growth 
rate  occurs  in  the  summer  from  July  to 
August  despite  increased  carnivorous  feed- 
ing. Growth  increases  sharply  from  Au- 
gust to  September,  however. 

In  the  fall  (mid-October)  at  the  two 
stations  analyzed,  Ephemeroptera  were 
found  in  19  percent  of  the  guts  at  Station 
IV  and  in  none  of  the  guts  at  Station  I. 
This  can  probably  be  explained  on  the 
basis  of  emergence  of  many  of  the  mayfly 
species.  One  mayfly  adult  was  found  in- 
gested, however.  Plecoptera  were  also 
relatively  rare  in  the  fall,  many  having 
emerged  already.  Many  summer  and  fall 
emergers  may  still  be  in  the  egg  stage  or 
too  small  to  be  prey.  Chironomidae  were 
least  numerous  at  this  time  of  year  also 
(present  in  only  17  percent  of  the  insects 
examined).  Diatoms  and  detritus  were  the 
most  numerous  items  found  in  the  guts 
during  the  fall.  Since  M.  signata  emerges 
mostly  in  the  summer,  the  fall  specimens 
represent  some  of  the  smaller  sizes.  They 
would  not  require  as  much  food  and  thus 
are  adapted  to  the  relative  paucity  of 
prey  species.  At  Station  IV  even  diatoms 
except  Navicula  spp.  were  not  numerous 
in  those  stonefhes  examined.  The  swift 
current  could  be  a  factor  in  removing 
many  from  the  substrate.  At  this  station 
more  individuals  had  fed-  on  mayflies  and 
stoneflies  than  at  Station  I.  Another  dif- 
ference   was    that    more    empty    foreguts 


March  1975 


GATHER,   GAUFIN:     UTAH    PLEGOPTERA 


47 


were  found  in  those  stoneflies  dissected 
from  Station  IV  (maximum  number 
reached)  than  at  Station  I  (9  vs.  2).  The 
lack  of  sufficient  prey  could  be  a  limiting 
factor.  However,  the  period  October- 
November  represents  one  of  the  highest 
absolute  growth  rates  during  the  year. 
The  growth  rate  data  were  obtained  by 
pooling  all  samples  from  all  stations,  how- 
ever. 

During  the  winter  (January-February) 
mayflies,    stoneflies,   caddisflies,   and   chi- 
ronomids    in    the    guts    increased    signifi- 
cantly.   Sheldon   (1972)    reported  similar 
results  in  some  of  the  Arcynopteryx  spe- 
cies complex  he  studied.    In  the  winter  at 
Station  IV,  there  were  prey  species  present 
in  more  of  the  guts  than  at  any  other  time 
during  the  year.    This  represents  an  in- 
crease in  the  absolute  growth  rate  but  a 
decrease  from  the  previous  month.    The 
calculated  absolute  growth  rates  from  No- 
vember to  December  and  from  December 
to  January  are  questionable,  since  only  3 
M.  signata  nymphs  were  collected  in  De- 
cember.   This  increase  in  growth  rate  is 
probably  correlated  with  the  availabilit}^ 
of  food,  because  only  the  family  Capniidae 
are  emerging.   Chironomids  were  the  most 
numerous,  being  present  in  88  percent  of 
the  guts  examined.  These  slower-moving 
insects  would  be  easier  prey  than  the  fast- 
er mayflies  and  stoneflies.    Stoneflies  also 
increased  dramatical!}^  in  the  guts  despite 
the   fact  that  the  numbers   available   de- 
creased due  to  winter  emergence.   By  win- 
ter, however,  the  summer  and  fall  emerg- 
ers,    such    as    the    family    Chloroperlidae, 
may  have  attained  a  sufficient  size  to  be 
suitable  prey.  Surprisingly,  the  increase  in 
numbers  of  prey  found  in  the  guts  also 
coincides  with  one  of  the  periods  of  the 
least  amount  of  growth.    The  low  winter- 
stream     temperature     could     stress     the 
nymphs  enough  that  some  growth  may  be 
sacrificed  even  though  an  adequate  food 
supply  may  be  available.    The  number  of 
fliatoms  and  other  algae  decreased  signifi- 
(antly,    probably    due    to    the    decreased 
water  temperature  and  lack  of  sufficient 
solar    radiation.     Nevertheless,    Nayicula 
spp.  and  Gomphoncma  spp.  were  still  nu- 
merous.    Enteromorpha   spp.   were   domi- 
nant in  21   percent  of  the  guts  examined 
(luring  this  period.    The  stoneflies  dissect- 
(h1  from  Station  V  were  collected  in  Feb- 
ruary,   however,    when    stream    tempera- 
tures were  beginning  to  warm  slightly  and 
solar   radiation   was   increasing.    Detritus 


was  present  in  43  percent  of  the  guts  and 
comprised  the  dominant  item  in  29  per- 
cent of  those  examined.  The  mmiber  of 
empty  foreguts  increased  to  a  maximum 
again  during  this  time. 

The  early   spring    (March)    was  also  a 
time    of    increased    carnivorous    feeding. 
This  also  coincides  with  a  significant  in- 
crease in  the  absolute  growth  rate.   At  this 
time  many  stoneflies  and  mayflies  are  ap- 
proaching  their   maximum   size   prior   to 
emergence.    The  quiescent  stage  just  be- 
fore emergence  may  also  make  them  easi- 
er prey.    The  percentage  of  mayflies  and 
stoneflies  in  the  guts  increased  at  Station 
V  but  decreased  at  Station  IV.    One  ex- 
planation   for    this    may   be    that    during 
spring  runoff,   the  water  level  is   deeper 
at  Station  IV  than  at  Station  V.   However, 
the  occurrence  of  chironomids  decreased 
significantly  at  Station  V  but  remained 
high  at  Station  IV.    The  reason  for  this 
was    not    determined.     Some    emergence 
could  have  occurred  at  Station  V,  but  be- 
cause the  stations  are  in  close  proximity 
this   would   not  seem  to  be   the   answer. 
Sheldon      (1972)      stated     that     mature 
nymphs  of  the  Arcynopteryx  species  com- 
plex he  studied  decrease  their  consumption 
of  animal  food  in  the  spring  (April).    At 
this  time  M.  signata  in  Mill  Creek  under- 
goes   a    noticeable    decrease    in    absolute 
growth  rate  prior  to  emergence.    Diatoms 
and  other  algae  were  still  relatively  rare 
in  the  guts  examined.   Presumably  spring 
runoff   created   spates   which  might  have 
removed  these  forms  from  the  substrate. 
Detritus  was  found  in  85  percent  of  the 
guts  at  Station  V.   Increased  runoff  due  to 
snow  melt  contributed  large  amounts  of 
allochthonous  detritus  to  the  stream.    The 
greater  depth  of  water  at  Station  IV  may 
have  prevented  the  concentration  of  detri- 
tus  and   effectively   removed   it.     On   the 
other  hand,   at   Station  V  the  channel  is 
wide  enough  that  shallow  areas  are  avail- 
able for  detritus  to  collect. 

In  discussing  food  habits  it  is  important 
to  remember  that  the  partitioning  of  re- 
sources is  accomplished  by  the  w^ide  distri- 
bution in  size  range,  which  decreases  in- 
traspecific  competition  (TTartland-Rowe, 
1964;  Radford  and  Hartland-Rowe,  1971). 
In  most  samples  a  difference  in  inter- 
ocular  measurement  of  M.  signata  aver- 
aged 0.6-1.0  mm  from  the  smallest  to 
largest  individuals.  This  was  a  significant- 
ly larger  size  range  than  in  the  euholog- 
nathan  species  studied. 


48 


GREAT  BASIN  NATURALIST 


Vol.  35,  No.  1 


Acknowledgments 

This  study  is  part  of  a  Ph.D.  research 
project  by  the  senior  author.  The  authors 
wdsh  to  thank  Bill  P.  Stark  for  his  helpful 
suggestions. 


Literature  Cited 

Baumann,  R.  W.  1967.  A  study  of  the  stone- 
flies  (Plecoptera)  of  the  Wasatch  Front,  Utah. 
Unpublished  M.S.  thesis,  Univ.  Utah,  Salt 
Lake  City.    114  pp. 

Brinck,  p.  1949.  Studies  on  Swedish  stoneflies. 
Opusc.  Entomol.  11:1-250. 

C.\THER,  M.  R..  AND  A.  R.  G.\UFiN.  1975.  Com- 
parative ecologv  of  three  Zapada  species  of 
Mill  Creek.  Wasatch  Mountains,  Utah  (Ple- 
coptera: Nemouridae).  Submitted  to  Amer. 
Midi.  Nat. 

CoFFMAN,  W.  P.,  K.  W.  Cummins,  and  J.  C. 
WuYCHECK.  1971.  Energy  flow  in  a  wood- 
land stream  ecosystem.  I.  Tissue  support 
trophic  structure  of  the  autumnal  communitv. 
Arch.   Hydrobiol.    68:232-276. 

Fenneman,  N.  M.  1931.  Physiography  of  west- 
ern United  States.  McGraw-Hill  Book  Co., 
New  York.    534  pp. 

Gaufin.  a.  R.,  a.  V.  Nebeker,  and  J.  Sessions. 
1966.  The  stoneflies  of  Utah.  Univ.  Utah 
Biol.   Ser.   14:1-93. 

Harper,  P.  P.,  and  J.  G.  Pilon.  1970.  Annual 
patterns  of  emergence  of  some  Quebec  stone- 
flies.   Can.   J.  Zool.  48:681-694. 

Hartland-Rowe,  R.  1964.  Factors  influencing 
the  life  histories  of  some  stream  insects  in 
Alberta.  Verb.  Internat.  Verein.  Limnol.  15: 
917-925. 


Hynes,  H.  B.  N.  1941.  The  taxonomy  and 
ecology  of  the  nymphs  of  British  Plecoptera 
with  notes  on  the  adults  and  eggs.  Proc.  Roy. 
Entomol.   Soc.    London   91:459-557. 

.      1970.     The    ecology    of    stream    insects. 

Ann.  Rev.  Entomol.   15:25-42. 

Illies,  J.  1966.  'Katalog  der  rezenten  Plecoptera. 
Das  Tierreich,  82.  Walter  de  Gruyter  and 
Co.,  Berlin.    632  pp. 

Khoo,  S.  G.  1968.  E.xperimental  studies  on  dia- 
pause in  stoneflies.  L  Nymphs  of  Capnia  bi- 
frons  (Newman).  Proc.  Roy.  Entomol.  Soc. 
London  43:40-48. 

Nebeker,  A.  V.  1971.  Effect  of  temperature  at 
different  altitudes  on  the  emergence  of 
aquatic  insects  fi-om  a  single  stream.  J.  Kans. 
Entomol.  Soc.  44:26-35. 

Oliver,  D.  R.  1971.  Life  history  of  the  Chiro- 
nomidae.    Ann.    Rev.   Entomol.    16:211-230. 

Radford,    D.    S..   and   R.    Hartland-Rowe.     1971. 
The  life  cycles  of  some  stream  insects   (Eph- 
emeroptera,  Plecoptera)  in  Alberta.  Can.  En 
tomol.   103:609-617. 

Richardson,  J.  W.,  and  A.  R.  Gaufin.  1971. 
Food  habits  of  some  western  stonefly  nymphs. 
Trans.  Amer.  Entomol.  Soc.  97:91-121. 

ScHW.'^RZ,  p.  1970.  Autoekologische  Untersuch- 
ungen  zum  Legenszyklus  von  Setipalpia-Ar- 
ten  (Plecoptera).  Arch.  Hydrobiol.  67:103-140. 

Sheldon,  A.  L.  1972.  Comparative  ecology  of 
Arcynopteryx  and  Diura  (Plecoptera)  in  a 
California  stream.  Arch.  Hydrobiol.  69:521- 
546. 

Swapp,  T.  E.  1972.  Food  habits  and  life  history 
of  stoneflies  of  the  Cle  Elum  River,  Washing- 
ton. Unpublished  M.S.  thesis.  East.  Washing- 
ton  State   Coll..   Bellingham.   58   pp. 

ZwicK,  P.  1973.  Insecta:  Plecoptera.  Phylo- 
genetisches  system  und  katalog.  Das  Tier- 
reich, 94.  Walter  de  Gioiyter  and  Co.,  Berlin. 
465  pp. 


RECORDS  OF  STONEFLIES  (PLECOPTERA)  FROM  NEVADA^ 

Mary  R.  Gather,'  Bill  P.  Stark,'  and  Arden  R.  Gaufin" 

Abstract. —  Distributional  data  are  presented  for    13    species   of   Nevada    stoneflies   including   eight 
species  new  to  the  state  list.    A  checklist  of  30  species  confirmed  for  Nevada  is  included. 


The  Nevada  stonefly  fauna  has  received 
scant  attention  from  collectors  working  the 
intermountain  region.  A  review  of  the 
literature  re\eals  22  species  recorded  from 
the  state,  most  of  these  without  specific 
distributional  data. 

We  are  reporting  eight  additional  spe- 
cies, Pteronarcys  princeps  Banks,  Isoperla 
fulva  Claassen,  Isoperla  patricia  Frison, 
Hesperoperla  pacifica  (Banks),  Utaperla 
sopladora  Bicker,  AUoperla  severa  Hagen, 
Sweltsa  color  a  dens  is  (Banks)  and  Malen- 
ka  tina  (Bicker).  Distributional  data  are 
presented  for  these  species  as  well  as  new 
records  for  species  previously  reported. 

We  thank  B.  W.  Baumann,  U.  S.  Na- 
tional Museum,  for  species  records. 

Pteronarcella  hadia  (Hagen). — New 
Becords:  Humboldt  Co.,  Cottonwood  Creek 
near  Paradise  Valley,  30-VII-68,  nymphs. 

Pteronarcys  californica  Newport. — New 
Records:  Washoe  Co.,  Beno,  18-VII-51,  T. 
H.  Zehrbach,  1    d"- 

Pteronarcys  princeps  Banks. — Distribu- 
tion: Elko  Co..,  Franklin  Biver,  Highwav 
11,  19-IX-57,  G.  F.  Edmunds,  Jr.,  and  B. 
K.  Allen,  nymph;  Lander  Co.,  Big  Creek, 
9-IX-54,  T.  C.  Frantz,  nymphs;  Big  Creek, 
Big  Creek  Campground,  14-VI-74,  B.  P. 
Stark,  18  d"  H  ?  ,  nymphs. 

Skwala  parcdlela  (Frison). — New  rec- 
ords: Elko  Co.,  Columbia  Creek,  20-IX-57, 
G.  F.  Edmunds,  Jr.  and  B.  K.  Allen, 
nymphs;  Secret  Creek  at  Secret  Pass,  15- 
VI-74,  B.  P.  Stark,  exuvium;  same  loca- 
tion, 4- VII- 74,  M.  and  E.  Cather,  nymph; 
Washoe  Co.,  Beno,  9-III-41. 

Isoperla  fulva  Claassen. — Distribution: 
Elko  Co.,  Secret  Creek,  Highway  11,  15- 
VI-74,  B.  P.  Stark,  cf  d'  ?  ?  ;  Secret  Creek, 
Secret  Pass,  15-VI-74,  B.  P.  Stark,  dd 
?  9  . 

Isoperla  patricia  Frison. — Distribution: 
Humboldt  Co.,  5  miles  north  of  Paradise 
Valley,   18-VI-67. 


Doroneuria  baumanni  Stark  and  Gau- 
fin. — New  Becords :  Lander  Co.,  Big 
Creek,  Big  Creek  Campground,  14-VI-74, 

B.  P.  Stark,  nymphs;  Nye  Co.,  South  Twin 
Biver,  lO-VII-54,  T.  C.  Frantz,  nymphs; 
Ophir  Creek,  4-X-54,  T.  C.  "Frantz, 
nymphs. 

Hesperoperla  pacifica  (Banks). — Distri- 
bution:  Elko  Co.,  Willow  Creek,  10-1-65, 

C.  Murvosh,  nymph;  Secret  Creek,  High- 
way 11,  15-VI-74,  B.  P.  Stark,  2  9  ;  Secret 
Creek,  Secret  Pass,  15-VI-74,  B.  P.  Stark, 
1  d"  1  9  ;  Lamoille  Creek,  2  miles  above 
Camp  Lamoille,  6- VII- 74,  M.  and  E. 
Cather,  1    d,  nymphs. 

Utaperla  sopladora  Bicker. — Distribu- 
tion: Elko  Co...  Lamoille  Creek,  2  miles 
above  Camp  Lamoille,  6-VII-74,  M.  and 
E.  Cather,  mature   d   nymph. 

AUoperla  severa  Hagen. — Distribution: 
Elko  Co.,  Secret  Creek,  Highway  11,  15- 
VL74,  B.  P.  Stark,  11  d  10  9;  Secret 
Creek,  Secret  Pass,  15-VL74,  B.  P.  Stark, 
1   d^  2   9. 

Sweltsa  coloradensis  (Banks). — Distri- 
bution: Elko  Co.,  Secret  Creek,  Secret 
Pass,  15-VL74,  B.  P.  Stark,  1  d';  Lamoille 
Creek,  2  miles  above  Camp  Lamoille,  6- 
Vn-74,  M.  and  E.  Cather,  2  c^  1  9 , 
nymph;  Lander  Co.,  Big  Creek,  Big  Creek 
Campground,  14-VL74,  B.  P.  Stark,  13 
d'  9  9  ;  White  Pine  Co.,  Lehman  Cave, 
Baker,  11-VL61,  S.  G.  Jewett,  Jr.,  6  d 
3    9. 

Triznaka  pintada  (Bicker). — New  rec- 
ords: Clark  Co.,  Deer  Creek,  5-VIII-69. 

Malenka  tina  (Bicker). — Distribution: 
Elko  Co.,  Secret  Creek,  Secret  Pass,  15-VI- 
74,  B.  P.  Stark,  1    d'  2    9  . 

Nevada    List 

The  following  is  a  checklist  of  the  30 
species  confirmed  as  occurring  in  Nevada. 

Pteronarcidae 
Pteronarcella  badia  (Hagen) 
P.  regularis  (Hagen) 


^Study   supported   by   NSF   Grant   No.    6986-609   and   EPA   Grant  No.  3033-364. 
^Department   of   Biology,    University   of   Utah,   Salt   Lake  City,    Utah  84112. 


49 


50 


GREAT    BASIN    NATURALIST 


Vol.  35,  No.  1 


Pteronarcys  californica  Newport 
P.  princeps  Banks 

Peltoperlidae 
Sierraperla  cor  a  (Needham  and  Smith) 
Soliperla  thyra  (Needham  and  Smith) 

Perlodidae 
Kogotus  modesius  (Banks) 
Skwala  parallela  (Prison) 
Isoperla  fulva  Claassen 
/.  rnarmorata  Needham  and  Claassen 
/.  patricia  Prison 

Perlidae 
Doroneuria  baumanni  Stark  and  Gaufin 
Hesperoperla  pacifica  (Banks) 

Chloroperlidae 
Utaperla  sopladora  Ricker 
Alloperla  sever  a  Hagen 
Suwallia  paUidula  (Banks) 
Sweltsa  coloradensis  (Banks) 
S.  pacifica  (Banks) 
Triznaka  pintado  (Ricker) 
Chloroperla  cydippe  Newman 

Needham  and  Claassen    (1925)    recorded  this 

species    from    Washoe    Co.,    Reno.     We   have  not 

examined   the   specimens   but   they    probably  are 
Hastaperla  chilnualna  Ricker. 

Nemouridae 
Malenka  californica  (Claassen) 
M.  coloradensis  (Banks) 
M.  tina  (Ricker) 
Podmosta  delicatula  (Claassen) 


Prosloia  besameisa  (Ricker) 
Soyedina  nevadensis  (Claassen) 
Zapada  cinctipes  (Banks) 

Capniidae 
Capnia  lacustra  Jewett 
Uiacapnia  tahoensis  (Nebeker  and  Gaufin) 

Leuctridae 
Paraleuctra  occidentalis  (Banks) 

References 

Baumann,  R.  W.  1970.  The  genus  Nemoura 
(Plecoptera)  of  the  Rocky  Mountains.  Un- 
published dissertation,  Univ.  Utah,  Salt  Lake 
City.  1 72  p. 

Gaufin,  A.  R.  1964.  Systematic  list  of  Plecop- 
tera of  Intermountain  Region.  Proc.  Utah 
Acad.   Sci.,  Arts,   Let.   41:221-227. 

Jewett,  S.  G.,  Jr.  1965.  Pour  new  stoneflies 
from  California  and  Oregon  (Plecoptera). 
Pan-Pac.  Entomol.  41:5-9. 

Logan,  E.  R.  and  S.  D.  Smith.  1966.  New 
distributional  records  of  Intermountain  stone- 
flies  (Plecoptera).  Biol.  Soc.  Nev.  Occas. 
Pap.  9:1-3. 

Nebeker,  A.  V.  and  A.  R.  Gaufin.  1965.  The 
Capnia  colunibiana  complex  of  North  Ameri- 
ca (Capniidae:  Plecoptera).  Trans.  Amer. 
Entomol.   Soc.   91:467-487. 

Needham,  J.  G.  and  P.  W.  Claassen.  1925. 
Plecoptera  or  stoneflies  of  America  north  of 
Mexico.    Thomas  Say  Pound.  2:1-397. 

Ricker,  W.  E.  1952.  Systematic  studies  in  Ple- 
coptera. Ind.  Univ.  Publ.  Sci.  Ser.   18:5-200. 


GROWTH  OF  PLECOPTERA  (STONEFLY)  NYMPHS 
Ar  CONSTANT,  ABNORMALLY  HIGH  TEMPERATURES^ 

Joseph  M.  Branham-,  Arden  R.  Gaufin-,  and  Robbin  L.  Traver- 

Abstract. —  Six  species  of  Plecoptera  were  maintained  at  four  different  temperatures,  which  were 
constant  and  higher  than  occurred  in  the  natural  habitat,  and  three  species  at  two  different  day  lengths. 
Each  animal  was  weighed  each  day  or  each  week.  Weight  of  two  species  in  the  wild  was  monitored 
from  periodic  collection. 

The  weight  of  each  anunal  fluctuated  rhythmically,  changing  about  five  percent  every  five  days. 
These  short-term  fluctuations  probably  resulted  from  changes  in  water  content.  Molting  occurred 
when  a  peak  weight  was  predicted  from  the  cycle  and  involved  temporary  gain  of  about  20  per- 
cent in  weight.  Grow'th  probably  stopped  for  some  time  before  molt  and  was  most  rapid  just  after- 
ward.   Many   animals   died   at  molt. 

The  time  before  death  was  less  for  univoltine  species  than  for  those  with  longer  life  cycles.  Plecop- 
tera collected  in  winter  from  water  near  0  C  lived  for  shorter  times  than  did  those  collected  in 
autumn  from  water  near  10  C.  Two  species  died  sooner  at  higher  temperatures  and  one  died  sooner 
with  shorter  day  lengths. 

Growth  in  the  laboratory  was  generally  slower  than  in  nature.  One  species  grew  faster,  while 
three  grew  more  slowly  at  higher  temperatures.  One  species  grew  faster  under  long-  than  short-day 
conditions. 

Premature  emergence,  expected  at  the  higher  temperatures,   did  not  occur,  except  in  one  animal. 


Calefaction,  or  abnormal  warming,  can 
alter  the  life  span  of  some  aquatic  insects. 
Some  die  soon  after  being  experimentally 
exposed  to  higher  than  normal  tempera- 
tures. Others  acclimatize  and  may  emerge 
prematurely  (Nebeker  and  Lemke,  1968; 
Nebeker,  1971).  Effects  of  constant,  high- 
er -  than  -  normal  temperatures  on  the 
growth  and  development  of  aquatic  in- 
sects are,  however,  little  understood. 

Temperature  changes  could  be  impor- 
tant cues  to  growth  and  seasonal  emer- 
gence. Species  living  in  or  near  springs, 
where  the  temperature  is  relatively  con- 
stant and  does  not  become  as  cold  as  in 
neighboring  streams,  frequently  emerge 
unseasonably  (Thorup,  1963).  Alternate- 
ly, temperature  changes  are  necessary  for 
normal  development  of  some  insects  (Hod- 
son  and  Rawy,  1956).  Such  changes  lead 
to  considerable  biochemical  restructuring 
on  a  seasonal  basis  (Somero  and  Hochach- 
ka,  1971)  and  can  also  affect  gene  ex- 
pression and  the  phenotype  of  insects 
(Sanderson,  1910;  Wigglesworth,  1965; 
Waddington,  1957).  The  elimination  of 
temperature  changes  could,  therefore,  in- 
terfere with  normal  development. 

Abnormally  high  temperatures  could 
have  long-term  cumulative  effects  on 
growth  and  development  (Sander,  1910; 
Richards,  1956,  1957).  Growth  could  be 
faster  than  normal  because  chemical  rates 
generally    accelerate,    or    slower    because 


metabolic  equilibria  are  upset  (Ludwig, 
1910) .  The  effect  on  growth  could  in  turn 
influence  development,  emergence,  and 
adult  function. 

Understanding  the  effects  of  tempera- 
ture on  growth  and  development  of  aqua- 
tic insects  would  be  useful  in  appraising 
the  effects  of  thermal  pollution,  because 
larval  insects  are  an  important  component 
of  aquatic  environments,  particularly 
streams  (Hynes,  1970).  The  purpose  of 
this  research  was  to  elucidate  the  growth 
patterns  of  some  species  of  Plecoptera 
maintained  in  the  laboratory  at  different 
temperatures.  Stonefly  nymphs  were 
chosen  because  they  develop  in  winter 
and  spring  when  the  stream  temperatures 
are  often  near  freezing  and  because  they 
are  abundant.  Three  carnivorous  and 
three  herbi\'orous  species  were  collected 
in  autumn  1972  and  established  in  the 
laboratory  at  four  different  temperatures 
and  two  different  day  lengths.  Some  were 
examined  and  weighed  daily;  others, 
weekly.  Growth  patterns  of  individuals 
were  plotted,  and  statistical  comparisons 
were  made  between  groups  of  animals  un- 
der different  conditions. 

Materials  and  Methods 

Some  features  of  the  species  studied  are 
outlined  in  Table  1.  All  were  collected 
in    October,    when    stream    temperatures 


^This  investigation  was  supported  by  Research  Contract  No.   14-12-438   from   the   National   Water   Quality   Laboratory,   Federal 
Water  Pollution  Control  Administration,   to  the  University  of  Montana    Biological    Station,    Bigfork,    Montana. 
'Department  of  Biology,   University  of  Utah,  Salt  Lake  City,  Utah  84112. 


51 


52 


GREAT  BASIN  NATURALIST 


Vol.  35,  No.  1 


Table  1.     Collection  sites  and  activity  cycles  of  species  studied. 


Family 


Time  of  Years 

Nonnal  in  Life 

Emergencef        Span 


Collection  Sites 


Nemoura  cinctipes 

Nemouridae 

Jan.-May 

Pteronarcella  badia 

Pteronarcidae 

May-Sept. 

Pteronarcys  californica 

Pteronarcidae 

Apr. -Aug. 

>1 

Arcynopteryx  signata 

Perlodidae 

June-July 

Acroneuria  pacifica 

Perlidae 

Mar.-Sept. 

>1 

Claassenia  sabulosa 

Perlidae 

June-Aug. 

Mill   Creek,   Salt  Lake  Co. 
Upper    Provo    R.,    Wasatch    Co. 
Lower  Provo  R.,  Utah  Co. 
Mill  Creek,  Salt  Lake  Co. 
S.   Fork  Provo  R.,   Utah  Co. 
Weber  R.,  Summit  Co. 


[•Gaufin,  Nebeker,  and  Sessions,   1966 


were  10-12  C,  and  the  experimental 
groups  were  established  in  the  first  part 
of  November,  except  where  noted  below. 

The  specimens  were  maintained  separ- 
ately in  perforated  plastic  (polyethylene) 
drinking  cups  9.5  cm  in  depth  and  7  cm 
in  top  diameter,  tapering  to  5  cm  at  the 
bottom.  The  cups  were  suspended  and 
w'ere  about  half  submerged  in  stainless 
steel  aquaria  approximately  100  cm  long, 
17  cm  wide,  and  15  cm  deep  and  having 
a  25.5  U  volume.  Nine  such  aquaria  were 
suspended  in  two  refrigerated  water  baths; 
four  in  a  bath  at  9.5  C  and  five  in  another 
bath  at  13  C.  Each  aquarium  was  heated 
with  a  thermostatically  regulated  element. 
Tap  water  flowed  through  each  aquarium 
at  about  0.4  L/min.  (or  about  one  aquar- 
ium volume/hour).  Each  aquariimi  was 
aerated  with  filtered  compressed  air  and 
agitated  wdth  a  paddle  wheel  that  mixed 
the  water  in  the  aquarium  and  caused  a 
predominantly  up-and-down  oscillation  of 
water  in  the  cups.  Concentration  of  dis- 
solved oxygen  remained  greater  than  90 
percent  of  saturation  in  the  cups.  Temper- 
ature was  maintained  with  a  standard  er- 
ror of  ±  0.02  C  and  a  range  of  about  ± 
0.5  C,  at  10,  12,  14,  or  16  C.  These  aquaria 
were  lighted  from  about  one  meter  above 
with  two  fluorescent  bulbs  on  a  12-hr. 
light,  12-hr.  dark  cycle.  Ambient  room 
light,  which  was  not  excluded,  varied 
somewhat. 

In  a  separate  experiment  to  estimate  the 
effect  of  light  periodicity,  three  species 
were  maintained  in  cups  suspended  in 
plastic  aquaria  flushed  with  aerated  run- 
ning water  (1  vol./hr.)  but  with  no  pad- 
dle wheel  (dissolved  O^,  >  90  percent 
saturation).  These  were  in  a  walk-in  cold 
room  that  maintained  the  water  tempera- 
ture at  11.5  C  within  the  same  limits  as 
in  the  other  aquaria.  They  were  illumin- 
ated with  fluorescent  lights  (2  bulbs)  that 
delivered  about  400  Lux  at  the  water  sur- 


face on  either  a  12L,  12D  (long- day)  or 
a  6L,  18D  (short-day)  periodicity  (Beck, 
1968). 

One  or  several  small  stones  and  several 
decaying  leaves  (cotton wood,  Populus  an- 
gustifolia;  or  maple,  Acer  grandidenta- 
tum)  were  kept  in  each  cup.  Herbivorous 
species  were  also  supplied  a  few  pellets  of 
Purina®  rabbit  chow  every  few  days,  and 
the  carnivorous  ones  were  kept  supplied 
with  a  mixture  of  small  aquatic  organ- 
isms, including  amphipods,  chironomids, 
ephemeroptera,  oligochaetes,  and  flat- 
worms  collected  from  near  a  local  fish 
hatchery. 

Changes  in  weight  were  monitored 
either  each  day  between  1  and  3  pm  or 
each  week  on  Thursday  between  1  and 
5  PM.  Each  animal  was  gently  blotted 
with  a  Kim  wipe®  (Kimberly-Clark  Cor- 
pororation),  air-dried  for  half  a  minute, 
and  weighed  to  the  nearest  0.1  mg  on  a 
Mettler  H6T  balance.  The  standard  er- 
ror of  the  method,  determined  by  re- 
peatedly weighing  the  same  animals  about 
20  times  in  an  hour,  was  about  0.002  of 
the  mean  for  animals  about  200  mg,  about 
0.005  of  the  mean  for  ones  about  150  mg, 
and  about  0.01  of  the  mean  for  animals 
about  30  mg.  Repeated  weighing  revealed 
a  gradual  but  statistically  significant  de- 
crease in  weight  of  each  animal  even 
though  they  were  retiu-ned  to  the  water 
between  each  determination,  so  the  stan- 
dard error  of  single  weighings  each  day  or 
week  may  have  been  less  than  for  about 
20  weighings  an  hour. 

Natural  growth  rates  were  calculated 
for  Pteronarcella  badia  and  Arcynopteryx 
signata  from  approximately  monthly  col- 
lections of  22  specimens  of  each  species 
from  the  initial  site.  The  animals  were 
weighed  as  above,  within  several  hours  of 
collection,  and  growth  Was  determined  as 
the  rate  of  change  of  the  average  weight. 

The  data  were  evaluated  by  regression 


'  March  1975 


BRANHAM,  ET  AL.:    PLECOPTERA  GROWTH 


53 


analysis  (Rao,  1958;  Bailey,  1959;  Alder 
and    Roessler,    1968),    using    a    Hewlett 
Packard     9100     B     computer     and     pro- 
grammed    procedures     supplied    by    the 
manufacturer.  Growth  trends  of  each  mdi- 
vidual  were  analyzed  from  weight  data  by 
determining  the  correlation  coefficient  (r) 
and,  from  it,  the  possibility  that  changes 
in  weight  were  correlated  linearly  with 
time,  according  to  the  calculated  slope  (m) 
which   intercepted   the   axis   representing 
weight  at  the  hypothetical  initial  weight 
(b)  independent  of  fluctuations  at  the  be- 
ginning of  the  observations.    Growth  was 
judged  to  be  positive,  negative,  or  insignif- 
icant from  the  correlation  coefficient   (P 
>    .05),    and    its   magnitude    was    deter- 
mined from  the  slope.    Absolute  growth 
values,  in  weight  units,   were   converted 
to  relative  ones  for  comparisons  between 
animals  by  determining  the  rate  of  change 
as    a   percentage   of   the   averaged    initial 
weight  (m  x  100/b).   The  effects  of  vari- 
ous conditions  were  then  evaluated  by  t- 
test  or  regression  analysis,  using  the  rela- 
tive growth  rates. 

Short-term    Fluctuations   in   Weight 

The  weight  of  all  stoneflies  examined 
every  day  increased  and  decreased 
rhythmically,  typically  varying  3  to  25 
percent  of  the  body  weight  about  every 
4-6  days  (Table  2;  Figs.  1,  2,  and  3).  The 
magnitude  of  weight  difference  between 
high  and  low  periods  exceeded  the  prob- 
able error  due  to  the  method,  and  the 
trends  apparent  in  the  plotted  data  indi- 
cate that  the  fluctuations  were  not  merely 
artifacts  of  the  method.  There  was  no 
apparent  correlation  between  the  rhyth- 
mic pattern  and  environmental  conditions, 
nor  were  the  cycles  of  different  animals 

Table  2.  Periodic  variations  seen  with  daily 
weighings  (Average  ±  SE)  calculated  from  the 
first  six  cycles. 

Period 
(days 
No.  of     between 
animals      peaks)  Amplitude* 


P.   californica 

5 

5.9  ±   .4 

3.3   ±   .3 

P.  badia 

4 

4.5  ±1.0** 

6.4  ±1.6 

A.  pacifica 

5 

5.0  ±   .4 

7.2  ±1.0 

C.   sabulosa 

1 

4.7 

5.7 

A.  signata 

1 

6.3 

12.3 

-^X 


Fig.  1.  Weight  changes  of  a  Pteronarcys  ca 
fornica  kept  under  various  conditions  This  md 
vidual  was  collected  in  December  and  acclunatu 
d  to  laboratory  conditions  for  15  days  without  too. 
It  was  then  weighed  daily  while  being  starved  A 
14  days),  fed  rabbit  chow  B  (15  days),  and  the 
leaves  and  rabbit  chow  C  (21  days),  while  being 
ept  at  10  C.  It  was  then  changed  to  16  L  an 
weighed  daily  while  being  fed  leaves  and  rabbit 
how  D  (50  days)  and  finally  weighed  once  eac 
week  for  10  weeks  (E).  It  did  not  molt  and  w 
s  still  alive  after  the  170-day  observation  period. 

in   phase   with   each  other,   even   though 
they  were  in  the  same  aquarium. 

It  was  hypothesized  that  periodic  feed- 
ing behavior  caused  the  observed  rhyth- 
mic weight  changes.  To  test  this,  ten 
Pteronarcys  californica    (collected  5   De- 


•Difference    between    peaks    and    troughs,    expressed    as   per- 
centase    of    average    weight.  >•,       • 

•Significantly   (p   <    .05)    different  only  from  P.  calijormca 
•Significantly  (p  <   .001)  different  only  from  P.  cahlonuca 


Fig  2.  Daily  weight  changes  before  and  a: 
er  molting  (Pteronarcys  californica  and  Acroneun 
pacifica)  and  before  emergence  (Arcynopteryx  si 
nata) .  The  molted  cuticle  was  found  and  the  an 
mal  weighed  about  six  hours  after  the  molt  occui 
ed. 


54 


GREAT   BASIN    NATURALIST 


Vol.  35,  No.  1 


Fig.  3.     Daily  weight   changes   before   death. 

cember  1972)  were  starved  for  15  days 
without  weighing;  then  weighed  every  day 
for  14  days,  while  starving;  then  fed  either 
Purina  rabbit  chow  or  decaying  leaves 
and  weighed  daily  for  15  days;  and,  fin- 
ally, fed  the  other  food  and  weighed  daily. 
Starvation  did  not  eliminate  the  rhythmic 
weight  changes  (Fig.  1),  although  all  the 
animals  were  losing  weight.  The  periodi- 
city remained  unchanged  upon  feeding 
either  or  both  foods.  The  amplitude  was 
not  affected  significantly  (P  >  .05)  when 
leaves  were  supplied  but  was  significantly 
increased  over  the  starvation  level  after 
feeding  rabbit  chow,  either  alone  or  with 
leaves  (Table  3). 

The  rhythmic  weight  changes  could 
have  been  associated  with  molting.  Weight 
changes  during  molting  were  observed  in 
a  few  specimens.  Of  the  16  animals 
weighed  daily,  five  molted  one  time  suc- 
cessfully, one  emerged,  and  six  died  molt- 
ing. One  Ps.  californica  and  one  Acro- 
neuria   pacifica   were   weighed    during   a 


successful  molt.  The  Ps.  californica  was 
24.4  percent  heavier  at  the  time  of  molt- 
ing than  just  before,  and  the  A.  pacifica 
was  19.0  percent  heavier.  The  ones  that 
died  molting  were  all  considerably  heavi- 
er than  they  had  been  (Table  8) .  Molting 
generally  occurred  when  the  pattern  of 
weight  change  indicated  that  a  peak 
weight  was  due  to  occur.  After  molting, 
the  period  between  weight  maxima  was 
usually  shorter  and  the  amplitude  greater 
than  it  had  been  before  the  molt;  but  af- 
ter several  cycles,  the  pattern  began  to 
resemble  the  premolt  condition  again. 

The  one  animal  on  a  daily  weighing 
schedule  that  emerged  (an  A?-,  signata, 
Fig.  2)  also  molted  and  emerged  at  a  peak 
in  the  weight  cycle.  The  adult  was  25 
percent  lighter  than  the  nymph.  Its  gut 
was  empty. 

The  frequency  of  weighing  affected 
the  weight  of  stoneflies.  Animals  weighed 
a  number  of  times  in  an  hour  lost  weight 
during  the  course  of  the  observations.  Five 
animals  that  had  been  weighed  every  day 
gained  weight  appreciably  when  the 
weighing  frequency  was  reduced  to  once 
a  week.  The  grovvi;h  rate  (m)  was  usu- 
ally slightly  greater  with  .the  less  frequent 
weighings,  but  the  change  in  rate  was 
not  statistically  significant  (Fig.  1).  Hand- 
ling could  have  caused  the  animals  to 
contract  and  expel  water  or  gut  contents, 
and  it  likely  interfered  with  feeding  pat- 
terns. 

Long-term  Growth  Patterns 

The  stoneflies  under  observation  lived 
for  various  periods.  Those  that  lived  a 
month  or  more  either  gained,  lost,  or  re- 
mained   the    same    weight    during    their 


Table  3.     Effects  of  food  on  periodic  variation,  seen  with  daily  weighings  of  P.  californica  (aver- 
age ±  SE). 


Period 

(days 

between 

peaks) f 


Amplitude 
(difference  be- 
tween peaks  & 
ti-oughs,  expressed 
as  a   %   of  aver- 
age wt.) 


fEstimated  from  three  peaks 

IfThree   that   did   not  respond   to   cither   food   were   e.xcluded   fr 
JOnly    those    fed    rabbit   chow    after   leaves   responded 
•Significantly   different    (p   >    .01)   from  starved   group  only 

♦•Significantly  different   (p  >   .001)   from  starved  group  only 


om  tnese  cali 


Gain  after 
feeding   (%   of 
previous  wt.) 


Starved 

10 

5.1    ±   1.0 

2.2   ±   .3 

Fed  leaves 

3it 

5.6  ±   1.0 

3.1    ±   .3 

4.3  ±   .3 

Fed  rabbit  chow 

^\ 

5.3  ±   1.2 

5.3   ±   .4** 

11.8   ±   .6 

Fed  both 

7+f 

5.1    ±   1.2 

4.4  ±   .5* 

8.7   ±   .5t 

March  1975 


BRANHAM,  ET  AL. :    PLECOPTERA  GROWTH 


55 


Table  4.     Average    (X    ±    SE    (N)  )    Weeks  of  life. 


Long  Day 

Short  Day 

IOC 

12  C 

14  C 

16C 

(11.5  C) 

(11.5  C) 

p.  californica\  25.2±3.0(5) 

17.2±2.1(6) 

24.5  ±4.9(6) 

23.8±4.6(6) 

P.    badia            15.0  +  2.5(5) 

17.7±1.3(6) 

11.4±2.1(6) 

10.7+1.6(6) 

6.0±1.1(11) 

6.7+   .6(11) 

A^.  cinctipes         4.7±    .9(6) 

6.2±1.2(6) 

2.5  ±   .3(6) 

4.7 ±    .9(6) 

A.  pacifica         18.9±5.8(5) 

16.5  +  3.5(4) 

18.0±4.3(6) 

14.2±4.3(5) 

15.6±1.9(11) 

11.1±2.6(11) 

C.  sabulosa         17.0±   .1(2) 

17.5±1.5(2) 

9.7±1.1(3) 

A.  signata            9.5  +  3.1(4) 

9.3±1.7(4) 

2.7+   .6(7) 

2.8+   .4(12) 

3.4±   .9(7) 

2.7±   .6(7) 

fEight   Ps.    californica   and   three  Ac.  pacifica  were   still   nlivo  at   the   tune  of  wiiting,    33   weeks  after  the  observations  began. 


life.  Some  individuals  lost  weight  for  part 
of  their  life  and  gained  in  another  part. 
It  was  important  to  consider  the  history, 
length  of  life,  and  ultimate  fate  as  well 
as  the  overall  growth  patterns  in  assessing 
the  effects  of  temperature  on  growth  rate. 

Some  species  lived  longer  in  the  labora- 
tory than  did  others  (Table  4).  In  gen- 
eral the  smaller,  univoltine  species,  Ne- 
moura  cinctipes.  Pa.  badia,  and  Ar.  signa- 
ta, did  not  live  as  long  as  the  larger  spe- 
cies, Ac.  pacifica,  Claassenia  sabulosa,  and 
Ps.  califo7-nica,  which  probably  live  for 
several  years  as  nymphs  (Table  1).  Pter- 
onarcella  badia  and  Ar.  signata  lived  for 
significantly  (P  >  .05)  less  time  at  higher 
temperatures.  Pteronarcella  badia,  Ar.  sig- 
nata, and  Ac.  pacifica  kept  in  the  light- 
control  chambers  died  significantly  sooner 
than  their  counterparts  in  the  tempera- 
ture experiments.  The  former  were 
caught  in  winter  from  streams  near  0  C 
and  were  kept  without  stirring.  Day 
length  had  no  significant  effect  on  the 
life  span  of  Ar.  signata  and  Pa.  badia. 

Weight  changes  prior  to  death  followed 
three  distinctive  patterns,  which,  in  some 
specimens,  could  have  been  related  to 
cause  of  death  (Fig.  3,  Table  5).  Often 
death  occurred  during  a  recognizable  molt. 
Sometimes  the  growth  curve  turned  sharp- 
ly up,  as  if  molting,  but  there  was  no  ex- 
ternal sign  of  molting.  More  often,  death 
followed  a  diminuation  in  the  amplitude 
of  cyclical  weight  changes  and  was  not 
marked  by  any  sharp  change  in  weight. 
Two  Ps.  californica  that  did  not  change  in 
weight  upon  being  supplied  with  either 
food  showed  this  pattern,  and  it  may  have 
reflected  starvation.  The  third  pattern 
was  marked  by  a  sharp  loss  of  weight  at 
death.  Adults  weighed  less  after  emerg- 
ing (Fig.  2),  and  such  sharp  terminal 
weight  loss  could  have  indicated  unsuc- 
cessful emergence.  The  circumstances  of 
death  of  all  animals  that  died  are  pre- 
sented in  Table  5. 


Periodic  molting  is  a  characteristic  of 
Arthropod  growth.  However,  many  of 
the  animals  observed  here  apparently  did 
not  molt,  while  many  others  died  in  the 
process  (Table  5).  No  A^.  cinctipes  and 
only  one  Ar.  signata  were  observed  to 
molt.  The  occurrence  and  frequency  of 
molting  for  the  other  species  are  enumer- 
ated in  Table  6.  Some  molts  were  prob- 
ably overlooked,  but  the  analysis  of  growth 
curves  suggests  that  most  were  detected 
by  the  presence  of  the  cast  cuticle. 

The  times  until  first  molt,  between 
molts,  and  until  death  for  ones  that  did 
not  molt  are  compared  in  Table  6.  Time 
before  the  first  molt  (considering  ones 
that  molted  successfully  or  that  died  molt- 
ing) was  quite  variable  and  was  probably 
a  function  of  the  condition  of  the  animals 
at  the  time  of  capture.  The  time  between 
molts  was  also  highl}'  variable.  The  aver- 
age period  before  the  first  molt  was  not 
significantly  different  from  the  time  be- 
tween molts  for  any  species.  The  aver- 
age length  of  life  of  individuals  that  did 
not  molt  was  also  the  same  as  the  average 
period  before  the  first  molt  and  the  period 
between  molts.  The  average  period  be- 
fore molt  differed  between  species:  Ps. 
californica  ^=  C.  sabulosa  >  Ac.  pacifica 
>  Pa.  badia  (t-test,  differences  considered 
significant  if  p  >  .05).   There  was  no  cor- 


Table  5. 

Circumstances  of  death   (percentage 

of  N). 

N     Mt 

It 

2t 

3t     Et 

P.    californica 

31+f       8 

26 

50 

15       0 

P.  badia 

52         21 

20 

53 

3       2 

N.  cinctipes 

22           0 

0 

100 

0       0 

A.    pacifica 

40ft     26 

11 

52 

10       0 

C.   sabulosa 

8         57 

0 

28 

14       0 

A.   signata 

42           2 

11 

74 

11       2 

fl   Growtli   curve   tiuTicd   up   at   death,   as   prior   to   molting 

2  Growth    cun-e    continued    imchanged    until    death 

3  Growth  cun'c  turned  down  at  death,  as  prior  to  emergence 
M  Died  molting 

E  Emerged 

IfFom-    additional    P.    californica    and    two    .'1.    pacifica    were 
accidentally  killed. 


56 


GREAT    BASIN    NATURALIST 


Vol.  35,  No.  1 


Table  6.     Occurrence    and   frequency   of   molts 


Number  observedf  Weeks   (average   ±   SE  (range  ) 

molting  Before  first  Of  life  of  ones 

Ox     Ix     2x     3x  molt  Between  molts  not  molting 


P.  californica 

P.  badia 

A.  pacifica 

C.  sabulosa 


16  ±  3  (12-30) 

5  ±  1  (  1-18) 

9  ±  1  (  2-21) 

15  ±  1  (13-17) 


15  ±  3  (10-19) 
7  ±  1  (  1-11) 
9  ±   1    (  2-18) 


17  ±  2  (3-32) 

5  ±  1    (1-19) 
8  ±  1    (3-22) 

6  ±  3  (3-  8) 


fMolting   recognized    by   finding   a   cast   cuticle   or   found 


in  the  process  of  molting. 


relation  between  size  and  the  length  of 
time  before  or  between  molts  either  with- 
in or  between  species.  There  was  also 
no  correlation  between  the  length  of  time 
before  molts  and  the  temperature  or  light 
period  for  any  species  examined. 

Changes  in  weight  accompanied  molting 
(Table  7).  Most  animals  weighed  during 
a  molt  (alive  or  dead)  showed  a  dramatic 
increase  over  the  premolt  weight.  Usually 
this  increase  was  much  greater  than  the 
longer-term  increase  (determined  by  com- 
paring the  average  weight  the  month  be- 
fore molt  wdth  the  average  weight  for  the 
month  afterward)  and  could  have  been  as- 
sociated with  the  mechanism  of  molting. 
Animals  that  molted  several  times,  and 
thus  apparently  were  adapted  to  labora- 
tory conditions,  grew  appreciably  between 
molts.  Some  of  the  ones  that  died  with- 
out molting  grew  about  the  same  amounts 
as  did  others  between  molts,  suggesting 
that  death  could  have  resulted  from  fail- 
ure to  molt. 

For  each  animal  observed,  the  growth 


rate  (calculated  as  the  regression  coef- 
ficient [m] ) ,  for  the  four  weeks  prior  to 
molting  usually  was  different  from  that 
for  the  month  after  molt  (Table  8).  There 
was  a  great  deal  of  variability  between 
individuals:  some  lost  before  and  gained 
after;  some  gained  before  and  lost  after; 
some  gained  or  lost  more  rapidly  before 
than  after;  and  vice  versa.  One  Ps.  cali- 
fornica lost  1.0  percent/ week  for  20  weeks, 
molted,  and  then  gained  1.1  percent/week 
for  15  weeks.  The  growth  patterns  in  Fig. 
2  are  from  apparently  normal  animals 
that  lived  many  months  in  the  laboratory. 
The  number  that  molted  and  lived  at 
least  a  month  afterward  was  small,  so 
averages  and  limits  (Table  8)  do  not  in- 
dicate significant  (p  >  .05)  differences 
in  average  pre-  and  postmolt  rates.  There 
was  no  significant  correlation  with  the 
controlled  parameters  of  temperature  or 
light. 

The  overall  growth  of  each  animal  un- 
til death  was  evaluated  by  calculating  the 
correlation  coefficient  (r)  and  the  regres- 


Table  7.     Change  in  weights  with  molting. 


Percent  change  in  weight  (average  ±  SE  (range)  ). 
N    During  moltf     N     With   moltfl  N       Between  moltsf       N  Total,  ones  not  moltingft 

P.  californica    4-  26 ±  2(22-30)  6  25±5(9-41)  3  22.6±4.9(13.1-29.1)  28  19.4±  10.1  (-22.8-174.3) 

P.  badia           10  65 ±   6(36-85)  10  12±4(  5-32)  5  9.8±5.7( -1.2-20.4)  30  21.6±   6.9(-25. 1-144.4) 

A.  pacifica       12  22±  4(  3-47)  12  2±2(-9-19)  8  15.6±5.5(  0.7-48.6)  15    -3.8±    1.6(-14.4-     9.2) 

C.  sabulosa        4  36±14(18-78)  3  1±7(-8-15)  0  110.0 

fAnimals  weighed  during  a  successful  molt  or  found  dead  in  the   process   of  molting.     Weight  during  molt  as  a   percentage  of 
the  last  premolt  weight. 
tfAverage  weight   for   the   month   after   molt   as   a   percentage   of    the   average   for    the   month   preceding  the   molt. 
JCalculated    from    the    growth    curve.     Slope    multiplied    by    the   time  and  e.\pressed  as  a   percentage  of  the  intercept   (i.e.,   the 
hypothetical  initial  weight) . 
ttTotal    growth    calculated    as   m    x   weeks   of   life   x    100/b. 


Table  8.     Growth   rates    (percentage   change/week)   before  and  after  molt  (m  ±  SE  (range)  ). 


Month  before  molt 


Month  after  molt 


P.  californica 

P.   badia 

A.  pacifica 
All 

Long  day 
Short  day 
C.  sabulosa 


-.16 
2.73 


.84   (-3.35-    2.05) 
1.34   (-2.37-10.69) 


.54  ±  .41  (-2.68-  6.32) 

.96  ±  .99  (-3.68-  3.46) 

-.02  ±  .50  (-2.53-  1.69) 

.38  ±  2.07  (-2.71  -  4.33) 


2.79   ±     .81    (    .09- 
-.39   ±    1.29   (-9.76 


5.75) 
5.75) 


1.17  ±     .56  (-3.23-10.98) 

3.23  ±  "1.20  (-  .05-10.96) 

-.30  ±   1.00  (-3.23-    1.69) 

-2.04  ±     .78  (-3.57-  -1.08) 


March  1975 


BRANHAM,  ET  AL.:    PLECOPTERA  GROWTH 


57 


sion  coefficient  (m)  in  0.1  mg  change/ 
week.  The  overall  growth  rate  was  some- 
times influenced  strongly  by  the  final 
phase  of  the  growth  curve  as  the  animal 
expired.  Growth  during  the  first  two 
months  was  therefore  calculated  separate- 
ly. To  simplify  comparison  between  spe- 
cies and  experimental  groups  (Table  4) 
the  growth  rate  (m)  was  expressed  as  a 
percentage  of  the  calculated  initial  weight 
(b,  the  intercept).  The  number  of  animals 
was  so  small  and  the  range  of  values  so 
great  within  each  group  that  averages  and 
standard  errors  are  of  little  meaning.  Re- 
gression analysis  of  rates  of  change  against 
temperature,  however,  indicated  that  tem- 
perature had  a  significant  effect  on  the 
growth  rates  of  some  animals  (r  signifi- 
cant at  p  >  .05).  The  proportionate  (per- 
centage) effect  of  temperature  on  growth 
rate,  exclusive  of  experimental  error,  is 
expressed  as  the  coefficient  of  determina- 
tion (r-  X  100)  (Alder  and  Roessler, 
1968).  The  relationship  between  differ- 
ence in  growth  rate  per  degree  Celsius  is 
expressed  by  m. 

These  observations  were  apparently  val- 
id only  for  the  animals  collected  in  Octo- 
ber and  November,  when  the  stream  tem- 
peratures were  still  about  10  C.  The  few 
animals  collected  later  in  the  year  from 
colder  streams  died  in  a  shorter  time  and 
generally  grew  at  rates  different  enough 
from  the  others'  that  it  seemed  best  to  ex- 
clude them  from  the  preceding  calcula- 
tions. 

Light  periodicity  could  also  have  af- 
fected development  in  the  laboratory. 
Significantly  more  Pa.  badia  molted  under 
long-  than  short-day  conditions.  There 
was  no  significant  difference  in  length  of 
life  or  growth  rate  between  animals  of 
either  species  held  under  long-  or  short- 
day  conditions.  Acroneuria  pacifica  lived 
longer  under  long-day  conditions  (15.5  dr 
2.0  vs  10.2  ±  2.3  weeks)  but  molted 
sooner  under  short  days  (7.1  ±  1.5  vs  9.1 
±1.5  weeks).  Significantly  more  molted 
under  long  than  short  days.  Their  growth 
rate  to  death  was  significantly  greater 
under  long  days  (1.9  ±  .8  percent/week) 
than  under  short  days  (-1.0  ±  .3  per- 
cent/week) (Table  6).  These  animals 
were  not  strictly  comparable  to  the  ones 
used  for  the  temperature  experiments: 
the  water  in  their  tanks  was  not  stirred, 
and  they  were  collected  later  in  the  year 
(Pa.  badia  and  Ac.  pacifica  in  November 


and  Ar.  signata  in  February).  Pteronar- 
cella  badia  and  Ar.  signata  kept  under 
equal  periods  of  light  and  dark  died  signif- 
icantly sooner  than  ones  under  similar 
light  and  temperature  conditions  but  col- 
lected earlier  and  kept  in  stirred  water 
(Table  4). 

Laboratory  conditions  were  quite  differ- 
ent from  nature,  so  it  was  desirable  to 
compare  growth  of  laboratory  and  wild 
animals  from  natural  populations  where 
possible.  Pa.  badia  and  Ar.  signata  per- 
sisted at  the  initial  collecting  sites  in  popu- 
lations of  relatively  uniform-sized  indi- 
viduals. Their  growth  in  nature  was  quite 
linear  for  both  species  between  October 
and  May  9  (Fig.  4).  The  average  per- 
centage of  weight  increase  per  week  of 
wild  Pa.  badia  was  6.26  ±  0.5  (r  =  .998), 
and  that  of  Ar.  signata  was  10.17  ±  .07 
(r  =  .938).  Such  an  analysis  was  im- 
practical with  the  larger  species  because 
of  the  simultaneous  existence  of  several 
year  classes  and  the  effect  of  investigator 
selection.  Wild  Ar.  signata  emerged  na- 
turally in  May  (water  temperature  5-8  C) 
and  Ps.  californica.  Pa.  badia,  and  Ac.  pa- 
cifica in  May  (water  temperature  10-11 
C). 

Discussion 

Stonefly  naiads  varied  in  weight  as  time 
passed.   Some  of  the  variation  was  associa- 


Fig.  4.  Growth  of  Pteronarcella  badia  and  . 
rcynopteryx  signata  in  the  stream.  Means  (wit 
SE)  were  calculated  from  periodic  samples  of  22 
nimals  and  converted  to  percentage  of  the  averag 
initial  weight. 


^H 


GHKAT   BASIN    NATURALIST 


Vol.  55,  No.  1 


tcrj  wilh  short-torm  IJuftu^jtiori  in  state, 
vvhiJo  some  represented  long-term 
"growth."  Variation  in  gut  content  rapid- 
ly  altered  the  weight  of  experimental  ani- 
]nals--as  much  as  10  percent  or  more 
n^'ig.  1,  Table  3j.  Cyclical  weight  changes, 
on  the  order  of  5  percent  variation  every 
5  days  (Tables  2  and  3),  occurred  even 
in  the  absence  of  feeding  and  (ould  have 
resulted  from  the  amount  of  water  in  the 
animal,  in  either  its  gut  or  its  tissue.  This 
cyclical  weight  change  could  have  been 
.issociated  with  molting.  Each  animal 
I  hat  was  observerl  daily  and  that  molted 
did     so    wlieii    a    predicted     peak     wciight 

liould  have  occurred,  and  animals 
weighed  ( onsiflerably  more  wlien  mfjiling 
llian  just  before  or  after  the  molt  riabb; 
7).  Such  rai)id  wciight  gain  could  be  rc- 
spoiisibb;  for  splitting  the  f)id  cuticle  and 
rilling  out  the  new  ont!  (Wiggbisworth, 
l')f)5j. 

The  laihifc  of  molting  was  a  common 
(ause  of  death  of  animals  in  th(?  labora- 
tory. Many  animals  died  during  or  just 
after  molting.  Others  flied  at  a  time  of 
peak  weight,  considerably  above  the  pre- 
vious average,  as  if  about  to  molt  (b^ig.  1 
Table  5).  Most  of  the;  animals  died  with- 
out any  molts  <!xc(!pt  for  Ac.  pacifica  and 
('.  s((l>iil()s(/  ri'able  fi).  On  the  average, 
llios(>  tbat  did  not  molt  died  approximately 
wlicu  they  should  have  moltcnl,  as  judged 
iioiri  the  average  time  before  or  Ix'tween 
molls,  '['heir  average  growth  was  also 
about  the  same  as  the  average  growth  be- 
tween molts  of  animals  that  survived  in 
som(!  s[)(H;ies  (Table  7).  No  A^.  cinclipcs 
and  only  oru;  Ar.  siirnala  wer(>  observed  to 
molt,  and  their  averag(i  life  was  bvss  tlian 
lor  tlu!  other  species  (Table  4).  It  seems 
likely  that  failure  oi  some  as])cct  of  molt- 
ing was  a  common  cimse  of  death  of 
stonefli(!s  kept  in  the  laboratory. 

Relatively  few  animals  lived  long  enough 
with  repealed  molts  (Table  1)  to  be  con- 
si<l(»re(l  normal.  The  one  P.  hadid  that 
emerged  (kept  at  1()  i))  molted  llu^  first 
week  in  the  laboratory  (1  Nov.)  with  a 
1*)  pei-((Mit  gain  in  w(Mght,  gained  28  [)er 
((Mit.  m<)r(>  in  the  next  8  weeks,  and 
emefged  in  .fanuary,  5  months  i)rema~ 
tin-ely.  Its  growth  was  almost  linear  from 
the  l)eginning  of  the  observations  (r  =: 
.<)9H)  and  slightly  less  (m  =  5.2  percent/ 
week)  than  the  average  for  the  species  in 
nature.  Most  of  the  P.  hatlia  that  molted 
successfully  and  then  lived  for  at  least  a 


month  gained  weight  prior  to,  and  as  a  re- 
sult of,  the  molt,  but  then  lost  weight  (Ta-  \ 
bles  6,  7j.  Food  may  have  been  deficient, 
or  the  newly  molted  ones  may  have  been 
less  tolerant  of  laboratory  conditions.  Two 
P.  californica  that  lived  for  the  entire  55- 
week  observation  period  molted  twice  each. 
Both  lost  weight  the  month  before  the 
first  molt  and  showed  no  significant 
change  the  month  before  the  second  molt. 
Both  gained  considerable  weight  at  both 
molts,  remained  at  a  higher  weight,  and 
( on  tinned  to  grow  at  an  accelerated  rate 
after  the  molt  (Fig.  2,  Tables  6,  7,  and  8). 
They  apparently  did  not  feed  before  the 
molt,  expanded  in  volume  during  the  molt, 
and,  at  the  larger  volume,  hardened  and 
began  to  eat.  The  four  Ac.  pacifica  that 
lived  for  the  duration  of  the  observations 
usually  were  not  growing  significantly 
prior  to  molt,  expanded  appreciably  dur- 
ing the  molt,  but  then  declined  to  a  weight 
considerably  below  the  premolt  weight 
within  a  day  or  so.  They  then  grew  rap- 
idly for  several  weeks,  until  the  previous 
weight  was  reached,  after  which  they  did 
not  grow  significantly  again  until  after 
anolh(!r  molt.  Some  regressed  at  molting  i 
and  lost  weight  overall  (Table  7)  (Fig.  2, 
Table  9).  Probably  their  pattern  was  not 
normal  but  reflected  subsistence  in  an  un- 
natural environment  (Beck  and  Bharad- 
waj,  1972). 

The  aniicipatcHi  |)remature  emergence 
(Nebeker,  1971  j  was  not  found.  Of  the 
animals  coIUhUhI  in  October,  only  one 
(Pa.  hadia)  emerged  and  only  a  few  died 
with  a  pattern  of  weight  changes  even  sug- 
gesting em(>rgence  (Table  4).  This  could 
hav(^  l)(HMi  because  the  animals  were  col- 
lected early  in  the  autumn  and  kept  at 
constant  temperatures  and  long-daylight 
conditions  at  or  above  those  existing  at 
the  time  of  collection.  Perhaps  some  en- 
vironmental cue  was  absent.  Or  possibly 
the  small  j)lastic  cups  in  which  the  ani- 
mals were  kept  were  too  confining  to  per- 
mit normal  behavior.  A  number  of  P. 
calijornica  collected  from  streams  near  O 
('  in  January,  acclimatized  to  16  C,  and 
kept  (onnnunally  in  fish-breeding  nets 
in  the  laboratory  did  emerge  in  March, 
three  months  before  the  wild  population 
(>merg(Hl  naturally.  One  Ar.  signata,  col- 
lected in  February,  emerged  22  days  after 
being  put  in  a  cup  and  kept  at  10  C  (Fig. 
2). 

The  lenglli  of  lile  of  Pa.  hadia  and  Ar. 


March  1975 


BRANHAM,  ET  AL.:    PLECOPTERA  GROWTH 


59 


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60 


GREAT   BASIN   NATURALIST 


Vol.  35,  No.  1 


signata  in  the  laboratory  was  significantly 
correlated  inversely  with  temperature, 
but  other  factors  were  apparently  more 
important  in  determining  the  life  span  of 
the  other  species  (Table  4). 

The  average  growth  rate  of  stoneflies 
kept  in  the  laboratory  was  considerably 
less  than  that  of  wild  ones  in  the  stream, 
for  the  species  that  could  be  compared,  al- 
though the  fastest  growing  ones  in  the 
laboratory  equaled  the  rate  of  those  in  na- 
ture. The  initial  growth  rate,  for  the  first 
two  months  of  life  in  the  laboratory,  was 
greater  than  later  on  and  was  significantly 
correlated  inversely  with  temperature  for 
Pa.  badia  and  Ar.  signata,  about  30  to  40 
percent  of  the  effect  (r-  x  100)  being  at- 
tributable to  temperature  (Table  9). 
Longer-term  growth  was  less  influenced 
by  temperature  for  these  species  (Table 
9),  suggesting  that  those  individuals  most 
affected  by  temperature  did  not  survive 
much  longer  than  about  two  months. 
Growth  of  Ac.  pacifica  was  similarly  af- 
fected inversely  by  temperature,  but  to  a 
lesser  extent  (only  24  percent  of  the  ef- 
fect on  initial  growth  rate  being  attribut- 
able to  temperature.  Table  9)  and  ap- 
parently not  enough  to  cause  early  death. 

The  growth  rate  of  Ps.  californica,  par- 
ticularly initiall}^,  was  increased  at  higher 
temperatures  (Table  11).  It  was  also  the 
longest-lived  species  in  the  laboratory  (Ta- 
ble 4). 

The  wdde  range  of  responses  to  the  stress 
imposed  by  laboratory  conditions  (includ- 
ing temperature)  is  remarkable  but  not 
unexpected.  The  Plecoptera  observed  here 
came  from  mountain  streams  that  natu- 
rally change  considerably  from  season  to 
season  and  along  their  course.  Variation 
in  ability  to  respond  to  stressful,  chang- 
ing environmental  conditions  would  be  of 
advantage  to  species  living  under  such 
conditions:  it  would  reduce  the  probability 
that  all  individuals  would  be  eliminated. 
Such  wide  variation  means  that  laboratory 
experiments  such  as  this  should  use  a 
large  sample  size  of  comparable  individu- 
als. It  probably  is  not  valid  to  compare 
animals  from  different  locations  or  ones 
collected  at  different  seasons,  and  Lud- 
wig  (1928)  has  demonstrated  that  differ- 
ent stages  of  development  of  an  insect 
vary  in  sensitivity  to  temperature. 

This  study  suggests  that  some  species 
(i.e.,  Ps.  californica.  Pa.  badia,  Ac.  pacifi- 
ca, and  perhaps  C.  sabulosa)  were  better 


suited  to  long-term  observations  than  were 
the  others.  Variables  other  than  tempera- 
ture had  pronounced  effects  on  stonefly 
growth  and  fate,  and  these  should  be  elu- 
cidated before  further  studies  are  under- 
taken. The  most  important  of  these  seem, 
subjectively,  to  be  water  quality,  the  na- 
ture of  water  movement  in  the  habitat 
relative  to  energy  expenditure  by  the  ani- 
mal, suitability  of  the  habitat  to  the  ani- 
mal's behavior,  and  (most  important) 
food.  The  physiological  state  of  individu- 
als is  more  difficult  to  assess  but  must  be 
considered. 

Literature  Cited 

Alder,  H.  L.,  and  E.  B.  Roessler.  1968.  Intro- 
duction to  probability  and  statistics.  W.  H. 
Freeman,    San   Francisco,   333   pp. 

Bailey,  J.  T.  J.  1959.  Statistical  methods  in  bi- 
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Beck,  S.  D.  1968.  Insect  photoperiod.  Academic 
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Beck,  S.  D.,  and  R.  K.  Bharadwaj.  1972.  Re- 
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Gaufin,  a.  R.,  a.  V.  Nebeker,  and  J.  Sessions. 
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HoDSON,  A.  C,  and  M.  a.  al  Rawy.  1956.  Tem- 
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Hynes,  H.  B.  N.  1970.  The  ecology  of  running 
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LuDwiG,  D.  1928.  The  effects  of  temperature 
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Nebeker,  A.  V.  1971.  Effect  of  high  winter 
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Nebeker,  A.  V.,  and  A.  E.  Lemke.  1968.  Pre- 
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Rao,  C.  R.  1958.  Some  statistical  methods  for 
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Richards,  A.  G.  1956.  Temperature  in  relation 
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and  suboptimum  temperatures  on  insect  de- 
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Sanderson,  E.  D.  1910.  The  relation  of  tem- 
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SOMERO,     G.      N.     AND     P.     W:     HoCHACHKA.        1971. 

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WATER  BALANCE  AND  FLUID  CONSUMPTION  IN  THE 
SOUTHERN  GRASSHOPPER  MOUSE,  ONYCHOMYS  TORRIDUS 

Vernon  C.  Bleich^'-  and  Orlando  A.  Schwartz^'S 

Abstr.\ct. —  Weight  loss  was  rapid  and  fluid  consumption  decreased  shai-ply  when  Onychomys  tor- 
ridus  were  exposed  to  salinities  greater  than  0.3  Molar.  The  southern  grasshopper  mouse  is  physio- 
logically unspecialized  for  maintaining  water  balance  in  its  xeric  habitat.  The  southern  grasshopper 
mouse  is  capable  of  weight  maintenance  on  smaller  daily  water  rations  than  is  the  northern  grass- 
hopper mouse  {Onychomys  leucogaster) .  Differences  in  the  water  balance  of  O.  tonidus  and  O.  leu- 
cogaster  may  influence  their  local  distributions  in  areas  of  sympatiy. 


Previous  investigations  of  water  bal- 
ance in  cricetid  rodents  have  included 
limited  information  on  the  grasshopper 
mice  of  the  genus  Onychomys.  Schmidt- 
Nielsen  and  Haines  (1964)  subjected  O. 
torridus  to  several  diets  and  to  increasing 
concentrations  of  NaCl  solution  to  test  the 
species's  ability  to  maintain  water  balance 
on  various  regimens.  They  used  body 
weight  maintenance  as  the  criterion  for 
demonstrating  water  balance.  Boice  (1972) 
presented  limited  data  on  daily  water 
consumption  in  O.  leucogaster^  and  he 
cited  the  lack  of  other  water  consump- 
tion data  for  the  genus.  This  investiga- 
tion was  undertaken  to  partially  fill  that 
void  and  to  provide  additional  informa- 
tion on  the  water  balance  of  O.  torridus. 

Methods 

Six  southern  grasshopper  mice,  O.  t. 
tularensis,  were  obtained  18  km  north  of 
Reyes  Station,  San  Luis  Obispo  County, 
California.  The  mice  were  taken  to  our 
laborator}-^  and  housed  in  steel  laboratory 
cages  (16.5  x  18  x  25.5  cm).  A  substrate 
of  commercial  mineral-type  "cat  litter" 
was  provided.  The  temperature  was  con- 
trolled (x=23  C;  range  21-24  C),  and 
the  photoperiod  was  set  to  coincide  with 
natural  conditions.  The  mice  were  fed 
an  unsupplemented  diet  of  sunflower 
seeds  ad  libitum  throughout  the  study. 

Our  experimental  design  was  modified 
from  that  of  Schmidt-Nielsen  and  Haines 
(1964)  and  McManus  (1972).  Water  in- 
take was  measured  using  inverted  gradu- 
ated cjdinders  fitted  with  angled  drink- 
ing tubes.  A  series  of  controls  was  used  to 
correct  for  what  little  evaporation  oc- 
curred.     Daily    fluid    consumption    was 

^Department   of  Biolog3%    CalifoiTiia   State   University,    Long  Beach  90840 

-Present   address:    California   Department  of  Fish   and   Game,   Chino    Fish    and    Wildlife    Base,     Rftute    5,    Bird    Far 
Chino  91710 

^Present    address:    Museum    of   Natural    History,    University    of  Kansas,  Lawrence  66045 


measured  to  the  nearest  0.1   cc.    All  ani- 
mals were  weighed  daily  to  the  nearest 

The  mice  were  exposed  initially  to  ad 
libitum  tap  water  for  a  three-week  period, 
during  which  time  their  weights  stabilized. 
Daily  fluctuations  of  bod}'  weights  between 
the  second  and  third  week  on  the  tap 
water  regimen  were  negligible.  Salinity 
(NaCl)  was  then  increased  by  0.1  M 
every  other  week,  from  0.1  M  to  0.7  M. 
The  mice  were  exposed  to  each  successive 
saline  solution  for  a  period  of  one  week. 
To  allow  for  rehydration,  they  were  given 
tap  water  for  one  week  between  each 
successive  increase  in  molarity. 

Results  and  Discussion 

Mean  body  weight  and  fluid  intake 
varied  with  increasmg  salinity  (Fig.  1). 
After  the  initial  stabilization  period, 
weights  showed  an  increase  through  0.3 
M  and  then  declined  sharpl}".  Weight 
loss  continued  through  0.6  M,  reaching  a 
low  value  of  57  percent  of  the  initial 
weights.  Four  animals  died  after  exposure 
to  0.6  M  NaCl,  but  two  survived  rehydra- 
tion and  died  after  two  days'  exposure  to 
0.7  M  NaCl.  The  initial  rise  in  body 
weight  can  be  attributed  to  fluid  reten- 
tion. Beyond  molarities  of  0.3  M,  weight 
loss  was  rapid,  owing  to  dehydration  and 
decreased  food  consumption. 

Fluid  intake  increased  sharply  with  ex- 
posure to  salt  concentrations  up  to  0.2  M. 
Consumption  dropped  slightly  during  ex- 
posure to  concentrations  of  0.3  M  and  0.4 
M  and  then  dropped  sharply  through  the 
period  of  exposure  to  concentrations  of 
0.6  M. 

The   rate   of   fluid    consumption   in   O. 


62 


March  1975 


BLEICH,    SCHWARTZ:    GRASSHOPPER    MOUSE 


63 


Fig.  1.     Water  intake   and  weight   of   Onych- 
omys  torridus  as  functions  of  NaCl  molarity. 


torridus  showed  an  initial  increase,  and 
then  a  decrease,  probably  an  avoidance 
reaction,  as  McManus  (1972)  reported 
for  the  chinchilla  {CJunchilla  Icmiger) . 
This  initial  increase  in  fluid  consiunption 
probably  meant  that  as  the  kidneys  came 
closer  and  closer  to  reaching  maximum 
limits  of  concentrating  capacity,  more 
and  more  saline  water  was  required  from 
which  to  extract  the  same  volume  of  physi- 
ologically useful  water.  Possibly  the  de- 
crease in  fluid  consumption  at  concentra- 
tions gi-eater  than  0.3  M  is  attributable  to 
the  unpalatability  of  concentrated  salt 
solutions,  as  McManus  (1972)  suggested. 

These  data  indicate  that  O.  torridus  is 
an  effective  osmoregulator  when  exposed 
to  NaCl  concentrations  up  to  and  includ- 
ing 0.3  M.  Beyond  that  point,  the  spec- 
ies is  not  able  to  maintain  water  balance, 
and  it  may  be  unable  to  survive  pro- 
longed periods  of  exposure  to  solutions 
greater  than  0.4  M.  These  results  are 
similar  to  those  reported  by  Schmidt-Niel- 
sen and  Haines  (1964).  On  a  diet  of  la- 
boratory chow  and  with  increasing  salin- 
ity of  the  drinking  water,  all  of  their  O. 
torridus  maintained  weight  on  0.2  M  Na- 
Cl. Four  of  the  six  mice  maintained 
weight  on  0.3  M  NaCl,  and  all  mice  lost 
weight  rapidly  on  0.4  M  NaCl  solution. 

Initial  rates  of  consumption  of  tap  water 
in  O.  torridus  allow  a  comparison  with 
the  data  of  Boice  (1972)   for  O.  leucogas- 


ter.  Five  mice  in  this  study  averaged  0.23 
cc/g/day  after  20  days.  O.  torridus 
averaged  0.11  cc/g/Zday  after  three 
weeks.  Although  possible  differences  in 
humidity  in  the  two  laboratories  were 
not  reported,  these  data  suggest  that  O. 
torridus  is  capable  of  weight  maintenance 
on  smaller  daily  water  rations  than  is  O. 
leucogaster.  The  possibilit}'  that  differ- 
ences in  the  water  balance  of  O.  torridus 
and  O.  leucogaster  influence  their  local 
distributions  in  areas  of  sympatry  lends 
itself  to  further  investigation. 

The  efficiency  of  O.  torridus  in  main- 
taining body  weight  on  concentrations  of 
NaCl  solutions  is  similar  to  those  of  some 
other  myomorph  rodents  that  have  been 
investigated,  including  Neotoma  micropus 
and  Neotoma  floridajia  (Birney  and  Two- 
mey,  1970),  Microtus  ochrogaster  and  Mi- 
crotus  pentisylvanicus  (Getz,  1963,  1966), 
Peroniyscus  floridanus  (Fertig  and  l^oryne, 
1963),  and  Rattus  norvegicus  (Adolph, 
1943).  All  these  species  lost  weight  or 
died  at  concentrations  of  0.3  M  NaCl  or 
greater. 

The  data  in  our  study  support  the  con- 
clusion of  Schmidt-Nielsen  and  Haines 
(1964)  that  O.  torridus  is  physiologically 
unspecialized  for  maintaining  water  bal- 
ance in  a  xeric  environment.  We  con- 
cur that  the  southern  grasshopper  mouse 
is  adapted  to  its  xeric  environment  by  its 
carnivorous  diet,  which  provides  sufficient 
moisture  for  the  species  to  maintain  its 
water  balance. 

We  thank  S.  Bailey  and  L.  Terzenbach 
for  assistance  in  the  field.  J.  P.  Kinney 
kindly  allowed  the  use  of  field  facilities 
at  his  disposal  and  gave  us  permission  to 
trap  on  private  property.  C.  T.  Collins 
provided  the  impetus  for  this  study,  and 
we  thank  him  and  R.  B.  Loomis,  California 
State  University,  Long  Beach,  for  their 
helpful  comments. 

References 

Adolph,  E.  F.  1943.  Do  rats  thrive  when  drink- 
ing sea  water?    Am.  J.  Physiol.  140:25-32. 

Birney,  E.  C,  and  S.  L.  Twomey.  1970.  Ef- 
fects of  sodium  chloride  on  water  consump- 
tion, weight,  and  survival  in  the  woodrats, 
Neotoma  micropus  and  Neotoma  floridana.  J. 
Mammal.  51:372-375. 

Boice,  R.  1972.  Water  addiction  in  captive 
desert    rodents.     J.    Mammal.    53:395-398. 

Fertig.  D.  S.,  and  J.  N.  Layne.  1963.  Water 
relationships  in  the  Florida  mouse.  J.  Mam- 
mal.  44:322-334. 


64                                                                          GREAT   BASIN  NATURALIST                                              Vol.  35,  No.  1 

Getz,   L.   L.     1963.     A  comparison  of  the   water  chinchilla     Chinchilla    laniger.      Comp.     Bio- 
balance    of    the    prairie    and    meadow    voles.  chem.  Physiol.  41A:445-450. 
Ecology   44:202-207.  Schmidt-Nielsen,  K.,  and  H.  B.  Haines.     1964. 

1966      Salt     tolerances     of     salt     marsh  Water  balance  in  a  carnivorous  desert  rodent 

meadow   voles.     J.    Mammal.    47:201-207.  the  grasshopper  mouse.   Physiol.  Zool.  37:259- 

McManus,  J.   J.     1972.     Water  relations  of  the  265. 


A  SYSTEMATIC  STUDY  OF  COENIA  AND  PARACOENIA 
(Diptera:  Ephydridae) 

Wayne  N.  Mathis^ 

iAbstract. —  Shore  flies  of  the  genera  Coenia  Robineau-Desvoidy  and  Paracoenia  Cresson  are  re- 
vised, resulting  in  the  description  of  two  new  subgenera  of  Paracoenia,  Calocoenia  and  Leptocoenia, 
and  of  four  new  species,  Paracoenia  ampla,  P.  calida,  and  P.  wirthi  from  California,  and  Coenia  al- 
pina  from  Labrador,  Canada.  Biological  information  on  P.  turhida  and  P.  calida  is  given  and  tlie 
known  distribution  for  each  species  is  presented.  P.  paurosoma  is  reported  from  the  Palearctic  Region 
for  the  first  time  based  on  specimens  from  Sweden;  several  new  distribution  records  from  North  Amer- 
ica are  also  included.  Characters  of  the  male  postabdomen  are  used,  and  the  male  genitalia  of  each 
species  are  illustrated.    Keys  or  references  to  all  known  species  of  these  genera  are  included. 


Introduction  and  Review 

Shore  flies  of  the  ephydrid  genera  Co- 
enia Robineau-Desvoidy  and  Paracoenia 
Cresson  are  common  and  widely  distribu- 
ted in  the  Holartic  region.  Typically, 
flies  of  both  genera  are  associated  with 
semiaquatic  or  aquatic  environments  and 
many  tolerate  a  diverse  range  of  seem- 
ingly inhospitable  habitats.  They  are  of- 
ten abundant  around  mineral  or  hot 
springs,  alkaline  lakes,  and  marginal  a- 
quatic  areas  where  the  water  is  highly 
saline.  This  study  was  initiated  to  fur- 
ther the  systematic  knowledge  of  these 
tmique  flies  and  to  provide  a  basis  for  de- 
tailed biological  investigations. 

A  synoptic  series  on  the  North  Amer- 
ican Ephydridae  was  started  by  Ezra  T. 
Cresson,  >.  (1942,  1944,  1946,  1949), 
whose  papers  reviewed  most  Nearctic  gen- 
era of  the  subfamilies  Psilopinae,  Noti- 
philinae,  and  Parydrinae.  His  mitimely 
death  precluded  the  completion  of  this 
series  and  left  the  synoptic  study  of  the 
subfamily  Ephydrinae  largely  unfinished 
imtil  Sturtevant  and  Wheeler's  review  in 
1954.  This  review  was  the  last  compre- 
hensive treatment  of  the  genera  considered 
here. 

Prior  to  Sturtevant  and  Wheeler's  pa- 
per, the  American  species  of  Coenia  or 
Paracoenia  had  not  been  treated  together. 
Coquillet  (1902)  and  Curran  (1927)  de- 
scribed the  first  Nearctic  species,  Coenia 
bisetosa  and  C.  turhida  respectively,  and 
Johnson  (1925)  included  C  palustris  (Fal- 
len) in  his  list  of  Ephydridae  from  Massa- 
chusetts. Johnson's  identification  of  the 
latter  species  as  C.  palustris  is  question- 
able, and  Sturtevant  and  Wheeler  include 
this  citation  under  C.  curvicauda  Meigen. 
After    studying    the    Ephydrinae    in    the 


Naturhistorisches  Museum,  Wien,  Cres- 
son (1930)  reviewed  the  European  species 
of  Coenia.  He  distinguished  C.  curvicauda 
from  C.  palustris  and  designated  a  lecto- 
type  for  C.  curvicauda.  Cresson  (1935) 
described  a  new  genus,  Paracoenia^  that 
included  two  new  species,  platypelta  and 
fumosalis^  in  addition  to  Coenia  bisetosa 
and  C.  turbida,  described  previously  from 
North  America,  and  two  European  species, 
C.  futnosa  (Stenhammar)  and  C.  beckeri 
(Kuntze).  C.  curvicauda  and  C.  palustris 
were  left  in  the  geiuts  Coenia.  Sturtevant 
and  Wheeler  described  one  additional 
species,  C.  paurosoma.  in  their  review  of 
1954. 

Cresson  delimited  Paracoenia  from 
other  genera  based  on  comparative  dif- 
ferences he  noted  in  the  dimensions  of 
the  head,  in  the  number  of  dorsocentral 
bristles  (three  in  Coenia,  four  in  Para- 
coenia), and  in  the  presence  {Paracoenia) 
or  absence  {Coenia)  of  well-defined  hum- 
eral bristles.  Most  specialists  have  con- 
tinued to  recognize  both  Coenia  and  Para- 
coenia c[S  waWA  genera.  Dahl  (1959)  stud- 
ied the  male  genitalia  of  both  genera  and 
suggested  that  the  marked  differences  be- 
tween them  supported  Cresson's  view.  The 
two  genera  were  also  recognized  by  Wirth 
(1965)  in  the  catalog  of  North  American 
Diptera.  Sturtevant  and  Wheeler,  how- 
ever, recognized  only  Coenia,  relegating 
Paracoenia  to  subgeneric  status. 

This  revision  is  based  primarily  upon 
a  comparative  study  of  the  male  postabdo- 
men and  a  reevaluation  of  previously  rec- 
ognized characters.  From  these  studies,  I 
generally  concur  with  Cresson's  concept 
of  Coenia  and  Paracoenia;  however,  I 
now  recognize  three  subgenera  in  the  lat- 
ter   genus,    Paracoenia,    Calocoenia,    and 


'Department  of  Entomology,  Oregon  State  University,  Con-allis,  Oregon  97331 


65 


66 


GREAT   BASIN    NATURALIST 


Vol.  35,  No.  1 


Leptocoenia.  The  basis  for  these  proposals 
will  be  treated  in  greater  detail  in  the 
sections  on  ])hylogcny  and  under  the  ap- 
propriate generic  discussions.  Four  new 
s])ecies  are  described,  three  in  Parncoenia 
and  one  in  Cocnia. 

Acknowledgments 

This  study  was  initially  suggested  b}- 
Dr.  Stephen  L.  Wood  while  I  was  an  un- 
dergraduate at  Brighani  Young  University. 
Many  of  the  preliminary  observations 
were"  completed  then,  although  the  stud}' 
has  since  been  expanded  as  new  species 
were  discovered  and  specimens  were  ex- 
amined from  a  wider  geographic  area.  I 
thank  Dr.  Wood  for  his  encouragement 
and  for  sponsoring  two  summers  of  field 
work. 

Special  thanks  are  extended  to  Dr.  Paul 
H.  Arnaud.  California  Academy  of  Scien- 
ces, to  Dr.  Willis  W.  Wirth,  Systematic 
Entomology  Laboratory,  ARS,  USDA-Na- 
tional  Museum  of  Natural  History,  Wash- 
ington, D.C.,  and  to  Mr.  Guy  E.  Shewell, 
Canadian  National  Collection  of  Insects, 
for  their  extensive  cooperation  and  assist- 
ance. 

T^'pe  specimens  were  borrowed  from 
the  Academy  of  Natural  Sciences  of  Phil- 
adelphia (Drs.  David  C.  Rentz  and  Selw>n 
S.  Roback);  the  National  Museum  of  Na- 
tural History  (Dr.  W.  W.  Wirth);  and 
the  Museum  of  Comparative  Zoology 
(Mrs.  Janice  C.  Scott  and  Dr.  John  F. 
Lawrence) . 

In  addition  to  the  above,  the  following 
institutions  and  curators  kindly  loaned 
specimens,  wdthout  which  this  study  could 
not  have  been  completed:  LTniversity  of 
Minnesota  (Dr.  Philip  J.  Clausen);  Kent 
State  University  (Dr.  B.  A.  Foote);  Uni- 
versity of  California,  Riverside  (Dr.  Saul 
I.  Frommer) ;  Los  Angeles  County  Mu- 
seum of  Natural  History  (Dr.  Charles  L. 
Hogue);  Iowa  State  University  (Dr.  Rob- 
ert E.  Lewis);  Cornell  University  (Dr. 
L.  L.  Pechuman) ;  University  of  Califor- 
nia, Davis  (Dr.  R.  0.  Schuster);  Wash- 
ington State  University  (Dr.  William  J. 
Turner) ;  Florida  State  Collection  of  Ar- 
thropods (Dr.  Howard  V.  Weems,  Jr.); 
and  Brigham  Young  UniAorsity  (Dr.  Ste- 
phen L.  Wood).  Dr.  Marshall  R.  Wheeler 
generously  loaned  specimens  from  his  pri- 
vate collection. 

I  also  wish  to  gratefully   acknowledge 


Drs.  John  D.  Lattin,  Carol  A.  Musgrave, 
Paul  Oman,  and  Paul  O.  Ritcher,  Oregon 
State  University,  for  their  technical  and 
editorial  assistance  and  for  many  stimu- 
lating discussions  on  systematics.  The 
stereoscan  electron  micrograph  was  taken 
by  Mr.  Alfred  Soeldner,  Oregon  State  Uni- 
versity. Dr.  Willis  W.  Wirth,  USDA- 
USNM  was  also  kind  enough  to  review 
this  study. 

Methods  and  Discussion  of  Characters 

All  observations  were  made  using  a 
stereomicroscope;  a  filar  micrometer  was 
used  for  the  measurements.  Morpholog- 
ical characters,  especially  those  of  the 
male  postabdomen,  w^ere  illustrated  using 
an  ocular  grid.  All  illustrations  were 
drawn  to  the  same  scale  on  mylar  draft- 
ing film.  Preparation  of  the  male  or  fe- 
male postabdomen  for  study  involved  its 
removal  and  heating  in  a  10  percent  so- 
dium hydroxide  solution  to  remove  ex- 
traneous tissue.  The  abdomen  was  then 
washed,  further  dissected,  and  compared. 
For  permanent  storage  the  abdomens 
were  preserved  in  plastic  microvials  filled 
with  glycerin  and  attached  to  the  appro- 
priate specimen. 

During  the  course  of  the  study,  I  ex- 
amined approximately  4,000  specimens, 
including  the  type  specimens  of  all  but 
Cocnia  curvicauda.  and  I  did  examine 
European  specimens  of  this  species.  The 
type  specimens  examined  formed  the  basis 
for  the  species  descriptions. 

Species  descriptions  are  purj)osefully 
brief;  for  the  most  part  they  summarize 
specific  differences  or  additions  not  found 
in  the  more  detailed  generic  descriptions. 
The  diagnoses  will  differentiate  the  spec- 
ies from  similar  taxa.  For  previously  rec- 
ognized species,  the  descriptions  also  con- 
lain  any  newly  acquired  information  for 
comparative  purposes.  Polymorphic  and 
polytypic  variations  are  included  under  re- 
marks. 

Characters  considered  in  this  study  are 
ffom  all  bod}^  tagma  and  have  been  quan- 
lifi(>d  where  appropriate.  Ratio  values  are 
based  on  an  average  of  ten  specimens  se- 
lected because  of  obvious  size  differences. 

Head. —  Eye-to-cheek  ratio.  This  is  the 
ratio  of  genal  height  to^  eye  height.  Meas- 
urements are  taken  from  the  head  in  pro- 
file. This  ratio  is  a  convenient  character 
for  some  species  groups. 


March  1975 


MATHIS:    AMERICAN    EPHYDRIDAE 


67 


Width-to-height  ratio.  This  ratio  is  cal- 
culated as  head  height  to  head  width; 
measurements  are  made  from  a  cephalic 
orientation. 

Eye-width-to-foce-length  ratio.  This 
ratio  is  hased  on  measurements  from  the 
head  in  profile  and  is  calculated  as  face 
length  to  eye  width. 

Height-to-length  ratio.  This  is  the  ra- 
tio of  the  height  of  the  head  in  profile  to 
its  length,  measured  from  the  most  ante- 
rior surface  of  the  face  to  the  posterior 
margin  of  the  eye.  Cresson  first  used 
this  character  when  describing  Paracoenia. 

Aristal  -pectinations.  The  length  of  the 
pectinate  branches  on  the  dorsum  of  the 
arista  is  compared  with  the  base  width 
of  the  arista.  This  character  is  sometimes 
difficult  to  use  and  is  best  seen  in  well- 
preserved  specimens. 

Interfoveal  hump  and  marginal  bristles. 
This  character  is  correlated  with  the  eye- 
to-cheek  ratio.  It  is  the  comparison  of  the 
hump  height  to  the  length  of  the  bristles 
along  the  oral  margin.  Hump  height  is 
related  to  genal  height. 

Facial  color.  This  character  is  subject 
to  considerable  variation  in  many  species, 
especially  species  of  Paracoenia,  but  it  is 
of  some  diagnostic  value  in  others.  The 
species  of  Coenia  and  Calocoenia  ha^e  rel- 
atively constant  facial  color. 

Postocular  bristles.  The  development 
of  the  dorsalmost  postocular  bristles  is 
useful  in  distinguishing  Paracoenia  from 
Coenia. 

Thorax. —  Acrostichal  hairs.  The  ar- 
rangement and  degree  of  development  of 
the  acrostichal  hairs  have  been  overlooked 
as  a  diagnostic  character  other  than  at 
the  species  level.  These  characters  are 
important  in  distinguishing  the  subgenera 
of  Paracoenia. 

Dorsocentral  bristles.  The  number  of 
dorsocentral  bristles  has  been  extensively 
used  as  a  major  character  and  was  ac- 
corded significance  at  the  generic  level  by 
Cresson. 

Humeral  bristles.  The  presence  or  ab- 
sence of  well-developed  humeral  bristles 
also  was  used  by  Cresson  to  delimit  these 
genera.  I  have  followed  Cresson  in  at- 
tributing generic  importance  to  this  char- 
acter as  well  as  to  the  number  of  dorso- 
central bristles. 

Halters.  The  color  of  the  halters  can  be 
used  to  distinguish  some  groups  of  spec- 


ies. Cresson  (1930)  mentioned  that  this 
character  is  usually  variable  and  is  not 
important  as  a  key  character. 

Costal  vein  ratio.  This  is  the  ratio  of 
distance  along  the  coastal  margin  between 
Ri  and  R:,+,;  to  the  distance  between  R2+3 
and  Ro+i.  All  measurements  are  the  maxi- 
imum  straight-line   distances. 

Mi+2  ratio.  This  is  the  ratio  of  the  dis- 
tance of  the  Mi+o  anterior  to  the  posterior 
cross  vein  to  the  distance  posterior  to  the 
posterior  cross  vein. 

Costal  bristles.  The  presence  or  absence 
of  costal  bristles  along  the  dorsal  and/or 
ventral  surface  is  diagnostic  of  some  gen- 
era. Calocoenia  is  the  only  taxon  of  Scatel- 
lini  with  prominent  bristles  on  both  sur- 
faces, a  character  found  in  many  species 
of  Eph3drini. 

Femoral  comb.  The  femoral  comb  is  a 
sexually  dimorphic  character  restricted  to 
the  males  of  Paracoenia  s.  str. 

Abdomen. —  Male  postabdomen.  The 
male  genitalia  previously  have  not  been 
used  as  characters  at  the  species  or  generic 
level;  I  have  found  them  extremely  use- 
ful at  both  levels.  These  characters  are 
discussed  more  fully  in  the  generic  and 
specific  descriptions. 

Female  ventral  receptacle.  The  shape 
of  this  structure  seems  to  be  of  consider- 
able diagnostic  value,  especially  at  the 
generic  level. 

Phylogeny  and  Classification 

Both  Coenia  and  Paracoenia  belong  to 
Scatellini  as  it  is  presently  characterized. 
The  tarsal  claws  are  curved  and  short, 
and  the  pulvilli  are  developed  normally. 
The  tribal  concepts,  however,  have  not 
been  reassessed  since  Wirth  (1948,  1970, 
1971),  Oliveira  (1954a,  1954b,  1957),  and 
others  (Collin,  1963;  Steyskal,  1970)  be- 
gan incorporating  characters  of  the  male 
postabdomen  in  their  treatments  of  vari- 
ous Ephydrinae  genera.  This  is  especially 
evident  with  the  annectant  genus  Austro- 
coenia  Wirth  from  South  America  as  well 
as  several  und escribed  genera  from  the 
neotropics,  which  will  require  further 
evaluation  before  a  reliable  classification 
of  the  higher  categories  can  be  achieved. 
Biological  information  and  systematic 
studies  of  the  immature  stages  would  al- 
so be  most  useful. 

Because  biological  and  morphological 
data  of  the  larvae  are  lacking  in  many  re- 


68 


GREAT   BASIN    NATURALIST 


Vol.  35,  No.  1 


lated  genera  and  in  some  of  the  taxa  here- 
in considered,  I  have  elected  to  recognize 
a  conservative  classification  in  this  study. 
Taxa  above  the  species  level  but  within 
the  generic  limits  of  Paracoenia  as  Cres- 
son  described  it  are  given  subgeneric  stat- 
us. But  this  status  is  provisional,  await- 
ing the  accumulation  of  additional  in- 
formation as  outlined  above  and  further 
assessment. 

The  subgenera  of  Paracoenia  are  pri- 
marily based  on  characters  of  the  male 
postabdomen  and  correlated  external  fea- 
tures. The  resulting  concepts  are  suffi- 
ciently distinct  to  be  easily  recognizable 
as  delimited  in  the  diagnoses  and  as  seen 
by  reviewing  the  figures.  Paracoenia 
{Paracoenia),  for  example,  is  the  largest 
subgenus  with  eight  species,  yet  each 
known  taxon  belonging  to  this  category 
can  be  readily  placed  without  difficulty. 
Further,   most   of  the   diagnostic   charac- 


ters are  apomorphus  and  define  mono- 
phyletic  groups. 

On  the  other  hand,  the  relationships  be- 
tween subgenera  are  somewhat  obscure 
and  the  generic  concept  is  not  as  neatly 
circumscribed.  More  reliance  is  ])laced  on 
chaetotaxy  characters  of  doubtful  signifi- 
cance. However,  coupled  with  our  mea- 
ger knowledge  regarding  biology,  habitat, 
etc.,  I  feel  that  the  genus  is  convenient 
and  does  reflect  a  cohesive  unit. 

Coenia  has  only  three  species,  all  of 
which  are  evidently  closely  related.  Ex- 
cept by  association  with  males  and  in 
some  instances  with  locality,  the  females 
of  one  species  are  generally  indistinguish- 
able from  those  of  another.  Similarly,  the 
males  closely  resemble  each  other,  al- 
though their  genitalic  characters  are  con- 
sistent and  constant,  a  fact  that  facilitates 
identification  and  classification.  This 
group  in  particular  needs  biological  study. 


Taxonomy 
Key  to  Coenia  and  Paracoenia 

Four  pairs  of  dorsocentral  bristles;  humeral  bristles  well  developed;  dorsal- 
most  postocular  bristles  subequal  to  verticals ..  Paracoenia  Cresson 

Three  pairs  of  dorsocentral  bristles;   no  developed  humeral  bristles;  dorsal 

postocular  bristles  much  smaller  than  verticals Coenia  Robineau-Desvoidy 


Genus  Paracoenia  Cresson 

Paracoenia  Q-esson,  1935,  Trans.  Amer.  Ent.  Soc. 
61:356.  Type-species,  Coenia  bisetosa  Coquil- 
lett.  by  original  designation.  Sturtevant  and 
Wheeler,  1954.  Trans.  Amer.  Ent.  Soc.  79:164- 
166  (review  of  Nearctic  species  as  subgenus  of 
Coenia).  Wirth.  1965,  USDA  Agricultural 
Handbook  No.  276,  pp.  755-756   (catalog). 

Diagnosis. —  Members  of  this  genus 
are  similar  to  those  of  Coenia  but  can  be 
distinguished  from  the  latter  as  follows: 
Postocular  bristles  immediately  posterior 
to  the  vertical  bristles  subequal  to  verti- 
cals; at  least  one  humeral  bristle  well  de- 
veloped, much  larger  than  the  surround- 
ing setae;  foiu-  pairs  of  dorsocentral  ])rist- 
les. 

Description. —  Small  to  large,  length 
2.1  to  4.4  mm,  females  usually  larger 
than  males;  dark  species,  often  with  sub- 
shining  metallic  reflections;  head  with 
characteristic  arched  prefrons. 

Head.  Front  (postfrons)  rectangular, 
wider  than  long;  margins  of  mesofrons 
directed  inward  anteriorly;  mesofrons 
subshining  with  metallic  reflections,  setu- 
lose;  ocellar  triangle  equilateral,  concolor- 


ous  with  fronto-orbital  areas,  dull,  micro- 
sculptured.  One  large  pair  of  proclinate 
diverging  ocellar  bristles;  postvertical 
bristles  various;  two  pairs  of  fronto-orbi- 
tal bristles;  both  inner  and  outer  vertical 
bristles  well  developed;  two  pairs  of 
strong  postocular  bristles  immediately  pos- 
terior to  vertical  bristles.  Antennae  dark 
brown  to  black,  pollinose;  second  segment 
setulose,  especially  on  median  and  ventral 
surfaces;  dorsum  of  arista  pectinate,  pec- 
tinate branches  up  to  three  times  the  width 
of  aristal  base,  sometimes  equaling  third 
antennal  segment  width.  Pruinose  face 
(])refrons)  protruding,  arched  with  inter- 
foveal  hump,  setulose;  setae  descending 
from  hump  and  along  ventral  margin 
strongest.  Eye  suboval;  gena  variously  do 
veloped  with  genal  bristle  toward  ventral 
margin.  Mouthparts  dark;  prementum 
large,  bulbous. 

Thorax.  Dorsum  of  mesonotum  sub- 
shining  to  dull,  generally  concolorous, 
sometimes  with  discernible  median  and 
lateral  stripes.  Acrostichal  hairs  in  ap- 
proximately six  irregidar  rows  to  two  dis- 
tinct rows;  four  pairs  of  dorsocentral  brist- 


March  1975 


MATHIS:    AMERICAN    EPHYDRIDAE 


69 


les;  one  pair  of  intra-alars;  two  to  three 
pairs  of  humeral  bristles;  two  pairs  of 
notopleurals;  one  pair  of  presuturals;  one 
pair  of  supra-alars;  two  pairs  of  post-alars; 
dorsum  of  scutellum  convex  to  flat,  setu- 
lose;  at  least  two  well-developed  pairs  of 
lateral  scutellar  bristles;  one  ])air  each 
of  mesopleural  and  sternopleural  bristles. 
Wings  transparent  to  infuscated;  costal 
setae  weak  to  strong,  on  dorsal  and/or 
ventral  margins.  Legs  dark,  pollinose  to 
subshining. 

Abdomen.  Abdomen  of  males  with  five 
visible  tergites,  females  with  six  to  seven, 
subshining  to  pollinose;  most  species  with 
pollinose  band  near  the  posterior  margin 
of  each  tergite;  fifth  abdominal  tergite  of 
males  with  anteroventral  ])rojection  in 
some  species  of  Paracoenia  s.  str.  Female 
postabdomen  with  three  complete  seg- 
ments, six,  seven,  eight;  ninth  segment 
with  tergite  not  fused  dorsally,  with  one 
pair  of  long  spines  on  ventral  margin; 
sternite  of  ninth  segment  also  with  one 
pair  of  spines;  female  abdomen  terminat- 
ing with  cerci.  Ventral  receptacle  vari- 
ously shaped.  Male  postabdomen  sym- 
metrical, reduced;  sixth  segment  with 
spiracles  only  although  European  species 
of  Paracoenia  s.  str.  have  a  transverse 
sclerotized  band  posterior  to  fifth  sternite 
which  could  represent  sixth  sternite; 
seventh  and  eigth  segments  absent.  Spir- 
acles one  through  six  present,  sixth  spir- 


acle in  membrane  between  fifth  abdomin- 
al tergite  and  epandrium,  all  other  spira- 
cles in  ventral  margin  of  respective  terg- 
ites. Epandrium  suboval  with  scattered 
setae;  dorsum  of  epandrial  plate  bearing 
two  setulose  cerci;  ventrally  epandrium 
terminates  at  juncture  with  prominent, 
projecting  surstyli  of  various  shapes.  Hy- 
pandrium  extending  dorsally  and  ven- 
trally, attaching  to  epandrium,  lateral 
hypandrial  process  sheathing  aedeagus; 
aecteagus  well  sclerotized,  pointed  apically. 

Discussion. —  For  purposes  of  classifi- 
cation the  species  of  Paracoenia  are  ar- 
ranged in  three  subgenera,  Paracoenia, 
Calocoenia,  and  Lepiocoenia.  The  latter 
two  subgenera  might  be  accorded  generic 
status  by  future  revisers  since  the  male 
postabdomens  and  other  characters  are 
fairly  distinctive.  But  before  the  generic 
classification  of  these  taxa  is  assessed,  the 
higher  classification  of  the  subfamily 
Ephydrinae  should  be  revised  on  a  world- 
wide basis  to  insure  consistency  in  generic 
concepts,  especially  the  distinguishing  gap. 
Such  a  review  will  necessarily  entail  a 
great  deal  of  descriptive  work  because  of 
our  spotty  knowledge  of  many  faunal 
areas,  i.e.,  the  Neotropical  Region,  which 
are  replete  with  undescribed   species. 

The  following  key  works  best  for  male 
specimens  and  includes  both  European 
species.  Illustrations  of  the  male  genitalia 
will   facilitate   accurate  identification. 


Key  to  Paracoenia  subgenera  and  species 

Acrostichal  hairs  in  two  rows;  dorsum  of  scutellum  slightly  convex  to  flat; 
posteroventral  margin  of  mesofemora  without  row  of  comblike  bristles  in 
males 2 

Acrostichal  hairs  in  four  to  six  irregular  rows;  dorsum  of  scutellum  con- 
vex; posteroventral  margin  of  mesofemora  with  dense  row  of  bristles  in 
males    subgenus   Paracoenia  Cresson   3 

Length  over  3.25  mm;  eye-to-cheek  ratio  1:0.25  or  larger;  well-developed 
costal  bristles  projecting  anteriorly  from  ventral  and  dorsal  surfaces 
subgenus  Calocoenia,  Caloceonia  platypelta  (Cresson) 

Length  under  3.00  mm;  eye-to-cheek  ratio  1:0.25  or  less;  bristles  along  cos- 
tal margin  not  developed 

subgenus  Leptocoenia^  Lepiocoenia  paurosoma  (Sturtevant  and  Wheeler) 

Fifth  abdominal   sternite  of  male  deeply  U-shaped,  Figs.  11,  12;  Palearctic  ....     4 
Fifth  abdominal  sternite  broadly  U-shaped,  Figs.  1-6;  Nearctic 5 

Length  approximately  5  mm;  dorsum  of  thorax  and  abdomen  grayish-blue 

to  light  brown,  dull;  pleura  gray P.   beckeri    (Kuntze) 

Length  usually  less  than  4  mm;  thorax    and   abdomen   darker,   bluish-olive 

green  to  greenish-gray;  subshining  dorsally P.  fumosa  (Stenhammar) 


70  GREAT   BASIN    NATURALIST  Vol.  35,  No.  1 

5.  Postocellars  small,  no  longer  than  their  distance  apart  at  base;  facial  pru- 

inosity  yellowish-gold  with  greenish-blue  metallic  reflection  showing 
through  around  interfoveal  hump;  ventral  projection  of  fifth  abdominal 
tergite  in  males  blunt,  parallel  to  remainder  of  ventral  margin  ....  P.  fumosalis 

Cresson 
Postocellars  larger,  longer  than  their  distance  apart   at  base,   facial   pruin- 
osity  various;  ventral  projection  of  fifth  abdominal  tergite  absent  or  not 
as  above  6 

6.  Process  of  fifth  abdominal  tergite  in  males  not  developed 7 

Process  of  fifth  abdominal  tergite  developed  into  a  projection  of  various  shapes    8 

7.  Subshining    with    metallic    blue    reflections;   eye-to-cheek  ratio   1:0.45   or 

larger;  associated  with  hot  sulfur  springs  P.  calida  n.sp. 

Subshining  with  metallic  green  reflections;    eye-to-cheek   ratio   1:0.3   or 

less;   associated  with  thermal  or  cool  water  P.  turbida  (Curran) 

8.  Process   of   fifth   abdominal  tergite   of  male  broadly  produced;   acrostichal 

hairs  few;  male  genitalia  as  in  Fig.  1 ;  presently  known  only  from  vicin- 
ity of  Los  Angeles,  California P.  ampla  n.   sp. 

Process  of  fifth  abdominal  tergite  not  as  broadly  produced;  acrostichal  hairs 

stronger;  surstyli  of  male  genitalia  not  pointed  distally  or  bare 9 

9.  Surstyli  thickened  basally,  at  least  one-half  total  length;   sheathing  projec- 

tion of  hypandrium  broadly  rounded  apically;  smaller,  length  3.1-3.6 
mm;  setation  less  well  developed,  especially    on    face    and    mesonotum; 

male  genitalia  as  in  Fig.  6  P.  ivirthi  n.  sp. 

Basal  expansion  of  surstyli  less  than  one-third  total  length;  lateral  process 
of  hypandrium  pointed;  length  3.7-4.4  mm;  facial  and  mesonotal  hairs 
well  developed  P.  bisetosa  (Coquillett) 

Subgenus  Paracoenia  Cresson  hump,  strongly  arched;  eye-to-cheek  ra- 

n            ■    n            ,n:>r-   rr          .          t7  .  c  tio   usually    1:0.22-0.45;    width-to-height 

Paracoenia  Cresson,   19i5,   Irans.  Amer.  Ent.  boc.  ^-      a    r\  rn                  •  ^^^    \      r          i         ..i 

61:356.  Type-species  Coenia  bisetosa  Coquil-  ratio  1 : 0.67;  eye-width-to-face-length  ra- 
lett,  by  original  designation.  Sturtevant  and  tio  1:0.45  (1:0.85  in  P.  callda) ;  height- 
Wheeler,  1954,  Trans.  Amer.  Ent.  Soc.  79:164-  to-length  ratio   1:0.93. 

Thorax.  Dorsum  subshining  although 
Diagnosis. —  This  subgenus  is  similar  subdued  in  aged  specimens,  generally 
to  Calocoenia  but  may  be  distinguished  as  shining  more  posteriorly.  Acrostichal 
follows:  acrostichal  hairs  in  several  ir-  hairs  in  four  to  six  irregular  rows;  other- 
regular  rows,  no  prominent  hairs;  dor-  wise  chaetotaxy  as  in  generic  description; 
sum  of  scutellum  convex;  costal  setae  scutellum  convex.  Costal  vein  ratio  1:0.2; 
weak,  in  a  single  row;  postero ventral  mar-  Mi+..  vein  ratio  1:0.90;  costal  setae  gen- 
gin  of  mesofemora  with  distinct  row  of  erally  weak,  developed  setae  on  dorsal 
bristles  in  males;  many  species  with  an  margin  only.  Mesofemora  of  males  with 
anteroventral  projection  of  the  fifth  ab-  j)osteroventral  row  of  comblike  bristles, 
dominal  tergite;  epandrium  of  male  with  Abdomen.  Subshining  to  pollinose,  if 
an  anteromedian  triangular  projection;  shining,  reflection  somewhat  less  than 
aedeagal  apodeme  subquadrate.  dorsum  of  scutellum.  Female  ventral  re- 
Description. —  Moderately  large,  ceptacle  with  operculum,  wider  than  high, 
length  3.1-5.0  mm;  dark  species,  often  extending  process  not  longer  than  oper- 
with  subshining  metallic  reflections.  cukmi  length.  Surstyli  of  male  postab- 
Head.  Front  rectangular;  mesofrons  domen  projecting  from  lateral  margins 
subshining  to  shining  with  metallic  re-  <^f  epandrium,  long,  variously  shaped;  a 
flections;  pectinate  branches  of  arista  up  triangular  process  lies  between  surstyli, 
to  throe  times  the  width  of  aristal  base,  ^"me  species,  with  a  median  groove  (see 
often  equaling  third  antennal  segment  figures  of  included  species), 
width.    Face  with  prominent  interfoveal  Discussion. —  The  species  included  in 


March  1975 


MATHIS:    AMERICAN    EPHYDRIDAE 


71 


this  taxon  form  a  fairly  homogeneous 
group  based  mostly  on  apomorphous  char- 
acters (see  diagnosis).  The  joint  posses- 
sion of  these  characters  dehmits  the  mono- 
phyletic  grouping  here  understood  as  Para- 
coenia  s.  str.  and  determines  the  sub- 
generic  concept.  Two  species  previously 
included  here,  P.  paurosoma  (Sturtevant 
and  Wheeler)  and  P.  platypelta  Cresson, 
are  sufficient!}'  distinct  to  fomi  the  basis 
for  new  subgenera. 

Paracoenia  s.  str.  is  Holarctic,  but  no 
one  species  is  presently  known  to  occur 
in  both  Eurasia  and  North  America.  How- 
ever, many  species  of  the  subgenus  are 
widely  distributed,  and  it  is  not  uncom- 
mon to  collect  two  or  more  species  from 
the  same  general  locality-.  Other  species 
such  as  the  Nearctic  P.  calida  and  P.  am- 
pla  and  the  Palearctic  P.  beckeri  are 
known  only  from  very  localized  geo- 
graphic areas. 

The  subgenus  contains  eight  species; 
six  are  Nearctic  and  two  are  Palearctic. 
All  of  the  Nearctic  species  except  P.  fum 
osalis  are  found  principally  in  western 
North  Ainerica.    A  more  detailed  zoogeo- 


graphic  account,  aside  from  the  general 
distributions  indicated  under  the  appro- 
priate species,  will  not  be  possible  until 
more  collection  data  become  available. 

The  paucity  of  biological  studies  pre- 
cludes a  detailed  accounting  for  the  en- 
tire subgenus.  For  the  Nearctic  region, 
only  P.  turhida  has  been  studied  in  any 
detail  (Brock  et  al,  1968,  1969).  How- 
ever, Dr.  B.  A.  Foote  and  associates  at 
Kent  State  University  and  Dr.  Karl  W. 
Simpson  of  Cornell  University  are  cur- 
rently engaged  in  studies  of  various  eph- 
ydrid  species  that  will  greatly  enhance 
our   biological   knowledge. 

Most  species  can  tolerate  harsh  environ- 
ments, especially  aquatic  habitats  with 
high  concentrations  of  various  salts.  These 
shore  flies  are  often  abundant,  for  exam- 
ple, along  the  margin  of  Great  Salt  Lake 
or  associated  with  hot  sulfur  springs  in 
Yellowstone  National  Park,  Wyoming. 
Scheiring  and  Foote  (1973)  further  re- 
port finding  larvae  in  the  shoreline  mud 
of  alkaline  lakes  and  in  sewage-impreg- 
nated mud.  The  larva  and  pupa  of  P. 
fumosa.  a  Palearctic  species,  were  de- 
scribed by  Beyer  (1939). 


Map  1. —  Distribution  of  Paracoenia  wirihi, 
filled  stars;  Paracoenia  platypelta,  filled  circles; 
Paracoenia  ampla,  open  circle;  and  Paracoenia 
calida,  open  star. 


Paracoenia   (Paracoenia)   ampla,  n.  sp. 
Fig.  1,  Map  1 

Diagnosis. —  Although  this  species  is 
quite  similar  to  P.  hisetosa,  it  can  be 
readily  distinguished  from  the  latter  by 
comparing  male  postabdomens.  The  sur- 
styli  of  P.  ampla  are  sinuate,  bare,  and 
strongly  narrowed  apically.  Further,  the 
shape  of  the  lateral  hypandrial  process 
is  distinctive,  especially  the  clavate  ex- 
tension. This  species  is  larger  than  most 
P.  bisetosa  and  in  general  is  less  setulose. 
The  acrostichal  hairs,  in  particular,  are 
weak  and  widely  scattered. 

Description. —  Length  approximately 
4.0  mm  (the  abdomen  was  removed  for 
dissection  before  measurements  were  tak- 
en); generally  subshining  with  bluish- 
green  metallic   reflections. 

Head.  Width-to-height  ratio  1:0.7; 
height-to-length  ratio  1:1;  eye-width-to- 
face-length  ratio  1 : 0.44;  facial  pruinosity 
brownish-gold. 

Thorax.  Setae  generally  weak,  scattered; 
costal  vein  ratio  1:0.17;  Mi+o  ratio  1:1. 

Abdomen.  Fifth  tergite  ventrally  pro- 
duced into  broadly  based  processes  that 
extend  anteriorly  to  basal  margin  of  fused 


72 


GREAT   BASIN    NATURALIST 


Vol.  35,  No.  1 


U-shaped  fourth  and  fifth  abdominal 
sternites,  processes  pointed  apically;  fifth 
abdominal  sternite  thin,  weak,  deeply  U- 
shaped  with  parallel  arms;  epandrium 
subquadrate;  medial  triangular  projection 
proportionately  small  to  epandrial  size; 
surstyli  directed  inward,  apically  nar- 
rowed, sinuate,  bare;  hypandrium  in  pro- 
file almost  rectangular,  with  extending 
process  slightly  clavate;  aedeagus  broad 
basally,  tapering  rapidly,  curved  and 
pointed  apically.  Male  genitalia  as  in 
Fig.    1. 

Distribution. —  Los  Angeles,  Cali- 
fornia. 

Types. —  Male  holotype  with  the  fol- 
lowing label  data:  Los  Angeles,  Cal.,  Apr 
29th,  1915,  M.  VanDuzee;  a  determina- 
tion label,  Coenia  hisetosa  Coq.,  1919, 
Cresson;  a  blue  M  C  VanDuzee  collection 
label.  The  type  will  be  deposited  with 
the  California  Academy  of  Sciences,  type 
number  12032. 

Remarks. —  This  species  is  known  only 
from  the  unique  male  holotype.  Recogni- 
tion of  the  specimen  as  representing  a 
new  species  is  justified  in  view  of  the 
very  distinctive  male  postabdomen.  Un- 
fortunately, P.  ampla  inay  already  be  ex- 
tinct due  to  the  tremendous  and  rapid 
urban  growth  in  the  Los  Angeles  area 
since  1915.  I  have  examined  several  Para- 
coenia  specimens  from  Los  Angeles 
County  but  none  were  ampla. 

Paracoenia  {Paracoenia)  hisetosa 
(Coquillett) 
Fig.  4,  Map  3 

Coenia   hisetosa   Coquilllett.    1902.    J.    N.    Y.    Ent. 

Soc.  10:183 
Caenia    [sic]    hisetosa:    Aldrich,    1905,    Smithson. 

Misc.  Coll.  66(1444):  631 
Paracoenia  hisetosa:   Cresson,   1935,  Trans.  Amer. 

Ent.  Soc.  61:356 
Coenia     {Paracoenia )    hisetosa:     Sturtevant     and 

Wheeler,  1954,  Trans.  Amer.  Ent.  Soc.  79:164 

Types. —  Male  holotype,  Salt  Lake, 
Utah,  25  June,  E.  A.  Schwarz  collector. 
The  type  is  deposited  with  the  National 
Museum  of  Natural  History,  Washington, 
D.C.,  type  number  6644.  This  specimen 
is  in  relatively  good  condition,  although 
the  wings  are  ragged  and  torn. 

Diagnosis. —  P.  hisetosa  is  similar  to  P. 
turhida.  but  the  former  is  larger,  more 
setulose,  and  more  brownish.    The  poste- 


rior margin  of  the  fifth  abdominal  stern-  | 
ite  is  sclerotized  and  of  uniform  thickness 
throughout.  The  Aentral  process  of  the 
fifth  abdominal  tergite  is  well  developed 
and  pointed.  Basally,  the  surstyli  are  en- 
larged but  narrow  quickly,  making  the 
lateral  margin  sinuate.  The  hypandrial 
process  usually  tapers  evenly  to  a  rounded 
point.  Outwardly,  this  species  might  be 
confused  with  P.  ampla,  but  the  male 
genitalia  of  P.  hisetosa  differ  sufficiently 
from  the  latter  that  recognition  of  either 
species  should  not  be  difficult. 

Description. —  Length  3.7-4.4  mm; 
dark  greenish-brown  dorsally;  laterally 
quite  pollinose. 

Head.  Mesofrons  with  bronze  metallic 
reflections.  Eye-to-cheek  ratio  1:0.34; 
width-to-height  ratio  1:0.69;  height-to- 
length  ratio  1:0.9;  eye-width-to-face- 
length  ratio  1:0.5. 

Thorax.  Dorsum  with  pollinose  an- 
terior, becoming  subshining  posteriorly; 
pleural  areas  largely  pollinose.  Wings 
infuscated  with  light  brown. 

Abdomen.    As  in  diagnosis  and  Fig.  4. 

Specimens    examined. —    1881. 

Distribution. —  Like  P.  turhida,  this 
species  is  primarily  a  western  North 
American  taxon,  although  collecting  data 
indicate  eastern  extensions  to  New  York 
(5  miles  W  Cardiff),  Pennsylvania  (Phil- 
adelphia), Delaware  (Bombay  Hook),  and 
Virginia  (Saltville).  The  Northwest  Ter- 
ritories (Nyarling  River),  Canada,  is  the 
northernmost  collection  site,  and  the  spec- 
ies ranges  from  there  southward  through 
most  of  the  Midwest  to  Texas  (Buffalo 
Spring  Lake)  and  into  Mexico  (Guada- 
lupe Can.,  B.  Calif.).  Westward,  speci- 
mens have  been  collected  in  every  state 
and  Canadian  pro\ince  west  of  the  100th 
])arallel. 

Remarks.^ —  This  is  the  most  common 
species  of  the  genus  and  among  the  most 
widespread.  It  is  also  one  of  the  most 
variable.  Facial  pruinosity  color  runs 
from  bright  brownish-orange  to  silver, 
and  the  general  body  color  varies  from 
shining  greenish-brown  to  a  subdued, 
grayed  green.  Age  polymorphism  is  also 
apparent;  older  specimens  are  often  more 
brownish  and  are  worn. 

Dr.  Willis  W.  Wirth  has  made  several 
collections  of  P.  hisetosa  from  aquatic 
habitats  with  varying  concentrations  of 
both  alkaline  and  saline  salts. 


March  1975 


MATHIS:    AMERICAN    EPHYDRIDAE 


73 


Map  2. —   Distribution   of   Paracoenia  fumosalis,  filled  circles;  Paracoenia  turbida,  filled  stars;  and 
Coenia  alpina,  enclosed  stai-s. 


Paracoenia  {Paracoenia)  calida,  n.  sp. 
Fig.  3,  Map  1 
Diagnosis. —  This  is  the  most  distinc- 
tive Nearctic  species  and  is  easily  separa- 
ted from  all  others  of  the  genus.  Exter- 
nally, the  blue  metallic  reflections  from 
the  dorsum,  the  protruding  prefrons,  and 
the  eye-to-cheek  ratio  are  diagnostic.  The 
male  postabdomen  resembles  that  of  P. 
bisetosa  but  differs  in  the  shape  of  the 
hypandrial  process,  which  is  more  or  less 


of  uniform  thickness  and  has  a  noticeable 
taper  just  before  the  apices.  Additionally, 
the  fifth  abdominal  tergite  does  not  have 
a  ventral  extension,  although  the  margin 
is  pointed.  P.  calida  and  P.  bisetosa  are 
approximately  the  same  size. 

Description. —  Length  3.4-4.3  mm, 
holotype  male  3.9  mm,  generally  dark, 
gray  pollinose  on  head  and  thoracic  plurae 
with  subshining  blue  metallic  reflections 
dorsally. 


74 


GREAT   BASIN    NATURALIST 


Vol.  35,  No.  1 


Head.  Eye-to-cheek  ratio  1:0.45;  height- 
to-length  ratio  1:1;  width-to-height  ratio 
1 : 0.71 ;  eye- width-to-face-length  ratio 
1:0.85.  Fronto-orbital  and  ocellar  triangle 
areas  blackish-gray,  concolorous  with  mar- 
gins of  frons;  postocellar  bristles  weak  in 
some  specimens;  pruniose  face  grayish- 
tan. 

Thorax.  Pleural  areas  dull,  pollinose; 
dorsmn  subshining  to  shining,  brownish- 
blue;  halters  dark,  reddish-bro\^^l  to  black; 
wings  completely  infuscated,  brown  to 
smoky.  Tarsal  claws  well  developed,  as 
long  as  third  or  fourth  tarsomere;  pulvil- 
lar  pads  proportionately  small  to  claw  size. 

Abdomen.  Dorsum  of  all  segments  with 
distinct  blue  to  purplish-blue  reflections, 
reflections  stronger  in  general  than  else- 
where on  body;  ventral  margin  of  fifth 
abdominal  tergite  broadly  pointed,  with- 
out lobelike  projection;  fifth  abdominal 
sternite  broadly  U-shaped,  narrow  pro- 
jecting arms  forming  obtuse  angle;  setae 
along  posterior  margins  of  tergites  much 
larger  than  rest  of  setae,  at  least  twice 
as  long,  in  some  female  specimens  three 
to  four  times  as  long.  Surstyli  of  male 
genitalia  with  more  or  less  gradual  taper, 
not  sinuate  or  pedunculate;  aedeagus 
short;  hypandrial  process  as  described  in 
diagnosis. 

Distribution. —  Wilbur  Hot  Springs, 
Colusa  Co.,  California. 

Types. —  Male  holotype,  allotype,  and 
all  paratypes  are  from  the  type  locality. 
Two  male  and  9  female  paratypes,  H.  J. 
Jacob;  1  male  and  1  female  para  type,  27 
June  1950,  L  .W.  Quate;  holotype,  allo- 
type, and  205  male  and  96  female  para- 
types, 25  June  1974,  W.  N.  Ma  this.  Pri- 
mary types  will  be  deposited  with  the  U. 
S.  National  Museum  of  Natural  History, 
type  number  72975.  Male  and  female 
paratypes  will  be  deposited  with  the  Cali- 
fornia Academy  of  Sciences,  Canadian 
National  Collection,  Academy  of  Natural 
Sciences  of  Philadelphia,  Kent  State  Uni- 
versity, Washington  State  University,  and 
Oregon  State  University.  The  remaining 
paratypes  are  in  my  collection. 

Remarks. —  Of  all  the  Nearctic  species 
of  Paracoenia  s.  str.,  P.  calida  is  perhaps 
the  most  remarkable.  Its  known  distribu- 
tion is  limited  to  a  hot  sulfur  spring  in  the 
foothills  just  east  of  Clear  Lake,  California. 
The  larvae  develop  and  mature  in  all  but 
the  hottest  water  where  they  can  easily 


be  collected  in  great  numbers.  The  adults 
are  also  abundant  and  were  often  ob- 
served to  congregate  in  large  clumps  near 
the  spring  source  where  shaded  or  pro- 
tected areas  could  be  found.  Empty  pu- 
paria  that  are  scattered  on  the  surface  of 
the  effluent  and  along  its  margins  are 
often  utilized  as  oviposition  sites.  Figure 
13  is  a  stereoscan  electron  micrograph  of 
the  egg  of  P.  calida. 

The  effluent  of  the  spring  emptied  into 
a  small  creek  around  which  swarming 
numbers  of  other  ephydrids  were  encoun- 
tered on  emergent  grasses  and  in  quieter 
eddies  on  the  water's  surface.  P.  calida, 
however,  was  not  common  there,  and  only 
an  occasional  collection  was  made  away 
from  the  hot  springs.  It  is  also  of  interest 
that  a  new  saldid  species  was  recently 
described  from  the  same  locality  (J.  T. 
Polhemus,  1967). 

The  specific  name,  calida,  is  descriptive 
of  the  habitat. 

Paracoenia  {Paracoenia)  fumoscdis 

Cresson 

Fig.  2,  Map  2 

Paracoenia  fumosalis  Cresson,  1935,  Trans.  Amer. 

Ent.  Soc.  61:356 
Coenia    {Paracoenia)    fumosalis:    Sturtevant    and 

Wheeler,  1954,  Trans.  Amer.  Ent.  Soc.  79:164 

Types. —  Male  holotype,  Rockport, 
Massachusetts,  28  August  1913,  C.  W. 
Johnson  collector.  The  type  is  deposited 
with  the  Boston  Natural  History  Society 
(Museum  of  Comparative  Zoology,  Har- 
vard University,  Cambridge,  Massachu- 
setts), type  number  31759.  One  male  and 
two  female  paratypes  are  topotypical. 
Four  female  paratypes  were  collected  on 
Nantucket  Island,  13  July  1926  by  C.  W. 
Johnson.  All  paratyj)es  are  deposited  with 
the  Academy  of  Natural  Sciences  of  Phil- 
adelphia. 

Diagnosis. —  Cresson  (1935)  stated  that 
this  species  is  similar  to  P.  hisetosa  and  P. 
funiosa,  which  is  Palearctic.  My  observa- 
tions agree  with  Cresson's,  although  in 
many  respects  P.  fumosalis  is  unique 
among  Paracoenia  species.  The  males 
are  most  easily  distinguished  from  similar 
taxa  by  the  rounded,  fingerliko  projection 
of  the  fifth  abdominal  tergite.  This  pro- 
cess is  parallel  to  the  ventral  margin  of 
the  tergite  that  is  deeply  incised  and  of 
uniform  thickness  before  the  slightly  en- 
larged, rounded  apex.    The  fifth  abdomin- 


March  1975 


MATHIS:    AMERICAN    EPHYDRIDAE 


75 


Map  3. —  Distribution  of  Paracoenia  bisetosa, 
enia  curvicauda,  filled  circles. 

al  sternite  is  subrectangular  without  ex- 
tending arms  from  the  posterolateral  mar- 
gins. The  surstyli  are  narrowly  S-shaped, 
and  the  median  triangular  process  often 
has  a  median  groove  that  is  cleft  apically. 
The  hypandrial  process  is  much  longer 
than  the  aedeagus,  and  apically  it  is  trun- 
cate. Externally,  P.  fumosalis  differs 
from  all  others  in  the  length  of  the  post- 
ocellar  bristles,  which  are  usually  shorter 
than    their    distance    apart    at    the    base. 


filled  stars;   Coenia  paurosoma,  open  stars;  and  Co- 

Further,  the  tannish-bronze  color  of  the 
pruinose  face  seems  to  be  constant. 

Description. —  Length  3.5-4.0  mm; 
dark  brown,  lightly  pollinose  to  subshin- 
ing  dorsally;  some  greenish-blue  metallic 
reflections. 

Head.  Mesofrons  shining  with  bluish 
reflections;  fronto-orbital  areas  subshining 
brown;  face  concolorous  with  mesofrons, 
pruinose,  tannish-bronze.  Eye-to-cheek 
ratio  1:0.3;  width-to-height  ratio   1:0.64; 


76 


GREAT   BASIN    NATURALIST 


Vol.  35,  No.  1 


height-to-length  ratio  1:0.87;  eye-width- 
to-face-length  ratio  1:0.41. 

Thorax.  Dorsally  subshining,  purplish- 
blue  reflections,  dark  brown.  Pleural  areas 
pollinose  except  dorsal  margin.  Wings  in- 
fuscated  \\dth  brown. 

Abdomen.  Concolorous  with  dorsum  of 
thorax.  Male  postabdomen  as  in  diagno- 
sis and  Fig.  2. 


Specimens  examined. 


368. 


Distribution. —  P.  fumosalis  is  pre- 
dominately a  nothern  and  eastern  North 
American  species.  I  have  examined  ma- 
terial from  Alaska  (Matanuska  and  Eagle 
River  flats)  and  from  all  of  the  Canadian 
provinces  except  British  Columbia  and 
the  Yukon  Territory.  In  the  continental 
United  States,  P.  fumosalis  ranges  west- 
ward to  Montana  (Libby),  eastward 
through  the  Great  Lake  states,  some  mid- 
western  states  (Nebraska,  Iowa),  and  in- 
to the  Northeast.  It  has  also  been  col- 
lected as  far  south  as  Florida  (Archbold 
Biological  Station,  Lake  Placid),  and  it 
presumably  occurs  between  Florida  and 
the  Northeast.  Wirth  (1965)  lists  Cali- 
fornia as  the  westernmost  extension  of  P. 
fumosalis,  but  I  have  not  seen  specimens 
from  California. 

Remarks. —  Examination  of  over  350 
specimens  of  this  species  from  a  wide 
selection  of  localities  within  its  distribu- 
tion revealed  very  little  morphological 
variation.  This  species  is  very  uniform 
except  for  slight  artificial  size  differences 
reflecting  the  mode  of  preservation.  The 
facial  coloration  seems  to  be  a  consistent 
diagnostic  character  unlike  other  wide- 
spread species  of  Paracoenia.  I  suspect 
that  the  species  is  quite  vagile  and  that 
the  apparent  uniformity  is  due  to  exten- 
sive genetic  exchange. 

Although  I  do  not  know  of  any  locali- 
ties in  which  this  species  is  associated  with 
hot  springs,  it  has  been  collected  around 
alkaline  and  saline  habitats.  Scheiring 
and  Foote  (1973)  reared  specimens  and 
report  finding  larvae  in  mud  impregnated 
with  organic  sewage. 


Paracoenia  {Paracoenia)  turhida 

(Curran) 

Fig.  5,  Map  2 

Caenia  [sic]  turbida  Curran,  1927,  Can.  Ent.  59:91 

Coenia     {Paracoenia)     turbida:     Sturtevant     and 

Wheeler,  1954,  Trans.  Amer.  Ent.  Soc.  79:165 


Paracoenia    turbida:    Wirth,    1965,    USDA    Agri. 
Handbk.    No.   276:756 

Types. —  Male  holotype  and  allotype, 
Old  Faithful,  Yellowstone  National  Park, 
Wyoming,  30  September  1924,  N.  Crid- 
dle.  Two  male  and  one  female  paratype 
have  the  saine  label  data  as  the  type.  All 
types  are  deposited  with  the  Canadian  Na- 
tional Collection,  type  number  2370. 

Diagnosis. —  Externally,  this  species  re- 
sembles P.  ivirthi  and  is  sometimes  con- 
fused with  P.  hisetosa.  However,  it  differs 
from  both  in  the  shape  of  the  hypandrial 
process,  which  apically  narrows  more 
abruptly  although  the  apex  is  rounded. 
Also,  the  lateral  margins  of  the  surstyli 
do  not  taper  apically  as  rapidly  as  P.  hi- 
setosa nor  are  they  enlarged  basally  as  in 
P.  wirthi.  Outwardly,  P.  turbida  is  smal- 
ler than  P.  bisetosa  but  of  approximately 
the  same  length  as  P.  ivirthi.  However, 
the  acrostichal  hairs  are  weaker  in  turbi- 
da. The  fifth  abdominal  sternite  is  more 
similar  to  that  of  bisetosa,  although  the 
more  sclerotized  posterior  margin  is  not 
as  wide  nor  as  uniform  in  thickness  as  it 
is  in  bisetosa. 

Description. —  Length  3.25-3.75  mm; 
dark  greenish-brown,  subshining  dorsally. 

Head.  Fronto-orbital  areas  dark  brown, 
pollinose  to  subshining.  Eye-to-cheek  ratio 
1:0.3;  width-to-height  ratio  1:0.65;  height- 
to-length  ratio  1:0.93;  eye-width-to-face- 
length  ratio  1:0.44. 

Thorax.  Anterior  area  of  dorsum  slight- 
ly pollinose,  becoming  subshining  to  shin- 
ing posteriorly;  pleural  areas  in  general 
more  subdued  than  dorsum,  especially 
along  margins.  Wings  nearly  transparent 
to   light  brown. 

Abdomen.    As  in  diagnosis  and  Fig.  5. 

Specimens    examined. —   674. 

Distribution. —  The  majority  of  col- 
lection localities  are  west  of  the  Rocky 
Mountains,  although  they  do  extend  east 
to  Nebraska  (Cherry  Co.,  Big  Alkali 
Lake),  Iowa  (Ames),  and  Ohio  (Kent, 
5.6  miles  SE).  Specimens  have  been  col- 
lected as  far  north  as  Alaska  (Circle  Hot 
Springs)  and  southward  through  Canada 
(British  Columbia  to  Manitoba)  and  the 
western  United  States  into  Mexico  (60 
km  S  Tijuana). 

Remarks. —  Strength  of  setation,  gen- 
oral  body  color,  especially  the  facial  pru- 
inosity,  and  overall  size  show  considerable 
variability.    Polymorphism  within  a  sin- 


March  1975 


MATHIS:    AMERICAN   EPHYDRIDAE 


n 


Figs.  1-5. —  Male  and  female  genitalia.  1  a,b  Paracoenia  ampla,  2  a,b,c  Paracoenia  funiosalis; 
3  a,b,c,d  Paracoenia  calida;  4  a,b,c  Paracoenia  bisetosa;  5  a,b,c  Paracoenia  turbida.  Fig.  a,  ventral  view 
of  cerci,  epandrium,  and  surstyli;  Fig.  b,  lateral  view  of  cerci,  epandrium  (epn),  surstyli  (sur),  aede- 
agal  apodeme  (aeg  ap),  hypandrial  process  (hyp  pr),  and  aedeagus  (aeg);  Fig.  c,  ventral  view  of  male 
fifth   abdominal   steniite;   Fig.   d,  lateral  view   of  female  ventral  receptacle. 


gle  population  of  P.  turbida  seems  to  var}^ 
as  greatly  as  the  total  species  variance. 
Character  displacement  was  not  apparent 
in  areas  where  turbida  occurs  sympatri- 
cally  with  other  Paracoenia  species. 

Brock,  et  al.  (1968,  1969)  have  studied 
the  biology  of  turbida  from  near  the  type 
locality  in  Yellowstone  National  Park, 
Wyoming.    They  found  that  both  larvae 


and  adults  feed  on  blue-green  algae  and 
the  filamentous  bacteria  of  mucilaginous 
mats,  which  develop  under  a  variety  of 
hot  spring  flow  conditions.  Fly  activity 
is  generally  restricted  to  cooler  mats 
(from  30-35C),  where  most  egg  laying 
occurs.  Above  40C  the  eggs  fail  to  hatch 
and  first  instars  die.  At  35C  the  life  cycle 
takes  approximately  14  days  (egg  to  egg), 


78 


GREAT   BASIN    NATURALIST 


Vol.  35,  No.  1 


and  mature  females  can  produce  100  or 
more  eggs  a  day.  Turbida  can  exploit 
transient  islands  of  available  resource 
quickly,  and  the  lar^'ae  soon  decimate  the 
optimum  habitat  (Weigert  and  Mitchell, 
1973).  Wiegert  and  Mitchell  also  an- 
alyzed the  interactions  between  the  algal 
mats  and  turbida  and  between  turbida 
and  a  mite  parasite  Partnuniella  thermalis 
Viets.  Mitchell  and  Redmond  (1974)  de- 
scribe the  egg  of  turbida  (several  stereo- 
scan  electron  micrographs)  and  suggest 
inechanisms  of  respiration  under  varying 
environmental  conditions.  Paracoenia 
turbida,  however,  is  not  endemic  to  hot 
springs,  and  the  details  of  their  feeding 
habits  and  habitat  preferences  under  dif- 
ferent conditions  could  vary  considerably. 
More  notes  on  the  biology  of  this  species 
may  be  found  in  Scheiring  and  Foote 
(1973). 

Paracoenia  {Paracoenia)   wirthi,  n.  sp. 
Fig.  6,  Map  1 

Diagnosis. —  This  species  resembles 
both  P.  bisetosa  and  P.  turbida  and  in 
many  respects  is  intermediate.  As  in  P. 
bisetosa,  the  ventral  margin  of  the  fifth 
abdominal  tergite  is  produced  into  a 
pointed  extension,  but  the  projection  is 
not  as  long  as  that  of  P.  bisetosa.  The 
general  coloration  and  setal  characters 
more  resemble  P.  turbida;  however,  males 
of  P.  wirthi  are  destinct  from  either  spec- 
ies in  the  shape  of  the  surstyli,  the  hy- 
pandrial  process,  and  the  fifth  abdominal 
sternite.  Basally,  the  surstyli  are  wide,  a 
condition  that  extends  to  about  one-half 
their  total  length.  The  hypandrial  pro- 
cess is  broadly  produced  apically  with  a 
bluntly  rounded  apex.  The  fifth  abdomin- 
al sternite  is  broad,  and  the  projecting 
arms  are  subparallel  and  short. 

Description. —  Length  3.1-3.6  mm; 
subshinning  with  bluish-green  to  green 
metallic  reflections. 

Head.  Mesofrons  greenish-blue,  shin- 
ing; pruinose  face  grayish-tan;  eye-to- 
cheek  ratio  1:0.22;  width-to-height  ratio 
1:0.6;  height-to-length  ratio  1:0.87;  eye- 
width-to-face-length  ratio  1:0.44. 

Thorax.  Acrostichal  hairs  in  three  to 
four  rows  anteriorly,  becoming  irregular 
with  five  to  six  rows  posteriorly.  Pleural 
areas  although  subdued  not  grayed.  Wings 
in  many  specimens  almost  transparent,  in 
others  smoky  infuscate. 


Abdomen.  Metallic  reflections  green  to 
olive  green.  Male  genitalia  as  in  diagnosis 
and  Fig.  6. 

Distribution.^ —  This  species  is  found 
in  the  Sonoran  desert  from  southern  Cali- 
fornia and  northern  Baja  California  east- 
ward into  Arizona.  It  extends  south  into 
Mexico  (Distrito  Federal,  Mixquic). 

Types. —  Male  holotype,  allotype  and 
21  paratypes  (7  males,  14  females),  Cali- 
fornia, Inyo  Co.,  1  mile  N  Tecopa  Hot 
Springs,  24  June  1974,  Wayne  N.  Mathis. 
Thirty-nine  paratypes  as  follows:  Cali- 
fornia, Inyo  Co.,  Tecopa  Hot  Springs,  16 
May  1965,  W.  F.  Barr,  1  male;  Inyo  Co., 
Shoshone,  24  June  1974,  W.  N.  Mathis,  1 
male,  5  females;  Inyo  Co.,  Shoshone,  1 
October  1935,  A.  J.  Basinger,  1  male,  3 
females;  San  Diego  Co.,  Mtn.  Palm 
Springs,  Anza  Desert,  2  March  1964,  1 
male.  Arizona,  Pima  Co.,  Lowell  Ranger 
Station,  6-20  June  1916,  32°  18.5'  N,  110° 
49'  W,  ca.  2,700',  1  male,  1  female;  Bill 
Williams  Forest,  August,  F.  H.  Snow,  1 
male,  1  female.  Mexico,  Baja  California, 
Guadalupe  Canyon,  19  May  1957,  F.  X. 
Williams,  1  male,  3  females;  Distrito  Fed- 
eral, Mixquic,  9  km  SW  Chalco,  4  August 
1965,  K.  R.  Valley,  4  males,  16  females. 
The  type,  allotype,  and  8  paratypes  will 
be  deposited  in  the  California  Academy 
of  Sciences,  type  number  12033.  The  re- 
maining paratypes  will  be  deposited  with 
the  U.  S.  National  Museum  of  Natural 
History,  Cornell  University,  the  Canadian 
National  Collection,  The  Academy  of  Na- 
tural Sciences  of  Philadelphia,  and  my 
collection. 

Remarks. —  Although  P.  wirthi  ap- 
pears to  be  intermediate  in  many  charac- 
ters, their  consistency,  especially  the  male 
genitalia,  justifies  recognition  as  a  new 
species.  As  with  other  species  of  the 
genus,  P.  wirthi  is  sometimes  associated 
with  hot  springs,  usually  along  the  mar- 
gins  of   the   effluent. 

P.  wirthi  is  named  to  honor  Dr.  Willis 
W.  Wirth  for  his  contribution  to  the 
systematics  of  the  Ephydridae  and  for  his 
helpful,  encouraging  responses  to  my 
many  questions. 

Calocoenia,  n.  subgen. 

Type-species:  Paracoenia  plalypelta  Cresson,  1935, 
monobasic 

Diagnosis. —  Although  similar  to  Para- 
coenia s.  str.  and  Leptocoenia,  Calocoenia 


March  1975 


MATHIS:    AMERICAN    EPHYDRIDAE 


79 


Figs.  6-10.—  Male  and  female  genitalia.  6  a,b,c,d  Paracoenia  ivirthi;  7  a,b,c,d,e  Paracoema  plaly- 
pelta;  8  a,d,e  Coenia  curvicauda;  9  a,h,c4,e  Paracoenia  paurosoma;  10  a,b,c,d,e  Coenia  alpina.  Fig.  e, 
lateral  view  of  internal  genitalia;  otliers  as  in  caption  of  1-5. 


may  be  distinguished  from  either  as  fol- 
lows: Externally,  Calocoenia  differs  from 
Paracoenia  s.  str.  in  the  arrangement  of 
acrostichal  hairs,  which  are  in  two  dis- 
tinct rows,  by  the  absence  of  a  postero- 
ventral  comb  of  bristles  along  the  male 
mesofemora,  by  the  flattened  scutellum, 
by  the  prominent  costal  bristles  project- 
ing anteriorly  from  both  dorsal  and  ven- 
tral margins,  and  by  the  generally  uni- 
form, concolorous  abdomen  which  in  male 


specimens  is  more  noticeably  narrowed 
apically.  The  most  apparent  difference 
between  Calocoenia  and  Lcptocoenia  is 
size;  Calocoenia  is  nearly  twice  the  length 
of  most  Leptocoenia.  Further,  the  eye-to- 
cheek  ratio  of  Calocoenia  is  at  least  1:0.2 
and  the  M1+2  vein  ratio  is  under  1:0.8. 
The  male  postabdomen  is  symmetrical; 
the  epandrium  is  subeUiptical  with  closely 
fused  surstyli  ventrally  and  with  a  median 
groove.   The  hypandrial  process  and  aede- 


80 


GREAT    BASIN    NATURALIST 


Vol.  35,  No.  1 


agus  are  tusklike,  long,  and  well  sclero- 
tized.  The  aedeagal  apodeme  is  crescent 
shaped. 

Description. —  Length  3.4-4  mm;  sub- 
shining  to  shining,  metallic  brown  to 
greenish-brown;  pollinose,  gra^'  ventrally. 

Head.  Mesofrons  shining,  bronze-gold 
metallic  reflections;  pectinate  branches  of 
arista  not  more  than  twice  aristal  width 
at  base;  pruinose  face  tan;  interfoveal 
hump  not  as  prominent  as  Paracoenia  s. 
str.,  dorsally  sloping;  eye  large,  subcircu- 
lar,  width  in  profile  double  the  length  of 
projecting  face  in  profile;  eye-to-cheek 
ratio  1:0.25;  width-to-height  ratio  1:0.66; 
height-to-length  ratio  1:0.9.  Chaetotaxy 
of  head  and  thorax  like  Paracoenia  s.  str. 
except  acrostichal  hairs. 

Thorax.  Acrostichal  hairs  in  two  rows; 
dorsum  pollinose  to  subshining;  pleural 
areas  concolorous  with  mesonotum  cen- 
trally, becoming  pollinose,  grayed  mar- 
ginally; halters  yellow.  Male  mesofemora 
without    comb    of    bristles.     Wings    with 


costal  bristles  on  dorsal  and  ventral  mar- 
gins; costal  vein  ratio  1:0.2. 

Abdomen.  Subshining  to  shining,  brown 
metallic  reflections;  fifth  abdominal  ter- 
gite  of  male  more  or  less  truncate,  with- 
out anteroventral  process;  fifth  abdominal 
sternite  with  three  posteriorily  oriented 
prongs.  Female  postabdomen  similar  to 
Paracoenia  s.  str.  Male  postabdomen  as  in 
diagnosis.  Fig.  7.  Ventral  receptacle  with 
operculum  wider  than  high,  extending 
process  considerably  longer  than  opercu- 
lum. 

Discussion. —  Calocoenia  is  a  mono- 
typic  subgenus  known  only  from  the  No- 
arctic  Region.  Nothing  is  known  about 
the  biology  of  the  included  species. 

Although  the  type-species  of  Calocoejiia 
was  originally  described  in  Paracoenia  s. 
str.,  its  inclusion  with  the  latter  subgenus 
would  form  a  paraphyletic  grouping  since 
the  sister  group  of  Calocoenia  is  Lepto- 
coenia.  The  sister-group  relationship  with 
Leptocoenia  is  deduced  from  the  joint  pos- 


Figs.   11-13. —  Male  genitalia  and  egg.  11  a,b,c  Paracoenia  beckeri;  12c  Paracoenia  fumosa;  13  stereo- 
scan  electron  micrograph  of  Paracoenia  calida  egg,  100  X.    Figures  as  in  caption  of  1-5. 


March  1975 


MATHIS:    AMERICAN    EPHYDRIDAE 


81 


session  of  the  following  apomorphous  fea- 
tures not  found  in  Paracoenia  s.  str.:  eye- 
to-cheek  ratio  less  than  1:0.3;  interfoveal 
hump  not  as  pronounced  as  in  Paracoenia 
s.  str.;  and  the  fifth  abdominal  tergite 
lacking  an  anteroventral  process. 

Paracoenia  (Calocoenia)  platypelta 

(Cresson) 

Fig.  7,  Map  1 

Paracoenia  platypelta  Cresson,  1935,  Trans.  Amer. 

Ent.  Soc.  61:356 
Coenia    {Paracoenia)    platypelta:    Sturtevant    and 

Wheeler,  1954,  Trans.  Amer.  Ent.  Soc.  79:165 

Types. —  Male  holotype,  Pine  Lake,  So. 
Cal.,  Johnson.  The  type  specimen  also 
has  a  small  label  with  the  male  sex  sym- 
bol, Cresson' s  pink  type  label,  and  a  red 
USNM  type  label,  number  51110.  The 
type  is  deposited  with  the  U.  S.  National 
Museum  of  Natural  History.  Cresson's 
original  description  also  lists  a  topotypi- 
cal  female  paratype.  I  have  examined 
this  latter  specimen,  presently  with  the 
Academy  of  Natural  Sciences  of  Philadel- 
phia, and  determined  it  as  Paracoenia 
turbida. 

Diagnosis  and  Description. —  See 
generic  description. 

Specimens  examined. —  516 

Distribution. —  This  species  is  known 
only  from  the  West  (US).  I  have  ex- 
amined specimens  from  Washington  to 
Alberta  (Laggan)  and  southward  to  New 
Mexico,  Arizona,  and  California.  I  have 
not  seen  specimens  from  Montana  or 
Wyoming,  although  collection  attempts 
have  been  made. 

Remarks. —  C.  platypelta  is  a  very 
homogeneous  species  exhibiting  little 
morphological  variation.  There  is  some 
color  polymorphism,  but  this  could  repre- 
sent age  polymorphism.  As  mentioned 
previously,  nothing  is  known  regarding 
the  biology  or  larvae  of  this  species. 

Leptocoenia.  n.  subgen. 

Type-species. —  Coenia  paurasoma  Sturtevant 
and   Wheeler,   monobasic 

Diagnosis. —  Leptocoenia  resembles 
Calocoenia  but  the  body  size  is  much 
smaller,  length  2.1-2.6  mm,  and  the  male 
postabdomen  of  Leptocoenia  differs  con- 
siderably. The  surstyli  are  well  separated 
apically  with  a  small  median  triangular 


process  between  them,  which  is  very  simi- 
lar to  a  comparable  structure  in  Para- 
coenia s.  str.  rhe  triangular  process  in 
Paracoenia  s.  str.,  however,  is  better  de- 
veloped in  comparison  with  the  lateral 
surstyli.  Externally,  Leptocoenia  is  simi- 
lar to  Calocoenia.  and  both  share  the  fol- 
lowing character  states:  The  acrostichal 
hairs  are  in  two  distinct  rows;  the  scutel- 
Imn  is  slightly  flat;  the  ventral  margin 
of  the  fifth  abdominal  tergite  is  not  pro- 
duced into  a  lobe;  and  the  posteroventral 
surface  of  the  mesofemora  does  not  bear 
a  row  of  comblike  bristles. 

Description. —  Length  2.1-2.6  mm; 
dark  brown,  pollinose. 

Head.  Fronto-orbital  areas,  mesofrons 
nearly  concolorous,  the  later  subshining; 
pectinate  aristal  branches  at  most  two  and 
one-half  times  aristal  width  at  base;  in- 
terfoveal hump  not  prominent,  without 
pronounced  dorsal  indentation;  pruinose 
face  light  tan;  longest  bristles  along  ven- 
tral margin  of  face  approximately  three- 
fourths  length  of  interfoveal  himip  height; 
genal  bristle  weak,  subequal  to  humeral 
bristles.  Eye-to-cheek  ratio  1:0.175; 
width-to-height  ratio  1:0.65;  height-to- 
length  ratio  1:0.93;  eye-width-to-face- 
length  ratio  1:0.3. 

Thorax.  Lightly  pollinose  dorsally. 
Acrostichal  setae  in  two  rows;  four  pairs 
of  dorsocentral  bristles;  humeral  bristles 
present;  halters  yellowish-brown  to 
brown.  Costal  setae  weak,  developed  only 
on  dorsal  margin. 

Abdomen.  Male  postabdomen  as  in  Fig. 
9;  ventral  receptacle  as  in  Fig.  9d. 

Discussion. —  In  many  respects,  this 
subgenus  is  pivotal,  linking  Paracoenia 
with  Coenia.  This  is  evident  in  characters 
associated  with  size  and  dimension,  but 
the  annectant  role  of  Leptocoenia  is  best 
evidenced  by  the  shape  of  the  female  ven- 
tral receptacle.  The  ventral  receptacle 
closely  resembles  those  of  Coenia  species 
and  is  probabl}^  less  likely  to  be  affected 
by  selective  pressure,  which  would  bring 
about  convergence  in  dimension  and  size 
of  external  characters  except  by  pleio- 
trophy.  I  attribute  considerable  import- 
ance to  this  feature  and  the  relationship 
with  Coenia  that  it  demonstrates. 

The  internal  male  genitalia  of  Lepto- 
coenia also  reflect  the  intermediate  posi- 
tion of  this  subgenus  with  Coenia.  This 
is  best  seen  by  comparing  Figs.  9e  and  lOe. 


82 


GREAT    BASIN    NATURALIST 


Vol.  35,  No.  1 


Paracoenia  {Leptococnia)  pawosoma 

(Sturtevant  and  Wheeler) 

Fig.  9,  Map  3 

Coenia  paurosoma  Sturtevant  and  Wheeler,  1954, 

Trans.  Amer.  Ent.  Soc.  79:165 
Paracoenia  paurosoma:  Wirth,  1965,  USDA  Agri. 

Handbook  No.  276,  p.  567 

Types. —  Female  holotype,  three  para- 
types  (one  male,  two  females),  Lander, 
Wyoming,  16  August  1950.  The  holo- 
type also  bears  a  collector  label,  M.  R. 
Wheeler,  and  type  number  6696.  A  fourth 
paratype  (female).  Rainbow  Lake,  Colo- 
rado, ^elev.  10,200',  50  hi.  The  original 
description  indicates  that  K.  W.  Cooper 
was  the  collector  of  the  fourth  paratype, 
but  no  collector  label  accompanied  the 
specimen.  The  holotype  is  deposited  with 
the  Academy  of  Natural  Sciences  of  Phil- 
adelphia, two  paratypes  with  the  U.S. 
National  Museum  of  Natural  History, 
and  two  paratypes  with  M.  R.  Wheeler. 

Di.\GNOsis  AND  Description. —  See 
generic  description. 

Specimens  examined. —  41 

Distribution. —  Paurosoma  has  been 
collected  from  Colorado  north  through 
Wyoming,  Alberta  (Laggan),  and  into 
Alaska  (Matanuska  Flats).  I  have  also 
examined  five  specimens  from  Sweden 
(Norrbotten,  3  km  N  Messaure). 

Although  paurosoma  is  Holarctic,  it  was 
the  most  recently  discovered  species. 
Moreover,  nothing  is  known  about  its  bi- 
ology. The  habitat  of  this  species  is  mon- 
tane; at  lower  latitudes  it  is  found  at  ele- 
vations up  to  10,500  feet  (Rainbow  Lake, 
Colorado) . 

Genus  Coenia  Robineau-Desvoidy 

Coenia  Robineau-Desvoidy,  1830,  Essai  sur  les 
Myodaires  2:800.  Type-species  Coenia  caricicola 
Robineau-Desvoidy  i  =  Ephydra  palusiris  Fal- 
len), by  monotypy;  Sturtevant  and  Wheeler, 
1954,  Trans.  Amer.  Ent.  Soc.  79:164-166  (re- 
view of  Nearctic  species  as  subgenus  of  Coenia) ; 
Wirth.  1965,  USDA  Agri.  Handbook  No.  276, 
pp.  755-756   (catalog) 

Caenia  emendation:  Walker,  1853,  Insecta  Britan- 
nica  Diptera  ?:259  (preoccupied-Newman, 
1853,  Entomological  notes.  Art  X.  Ent.  mag., 
pp.  372-402,  Coleoptera) 

Diagnosis. —  Coenia  species  resemble 
those  of  Paracoenia^  especially  the  sub- 
genus Leptocoenia.  but  they  are  differen- 
tiated as  follows:  Dorsalmost  postocular 
bristles  not  subcqual  to  the  verticals;  no 


well-developed  humeral  bristles;  two  pairs 
of  dorsocentral  bristles;  halters  brownish- 
yellow  to  dark  brown;  and  fifth  abdomin- 
al sternite  of  males  longer  than  wide.  The 
size  of  Leptocoenia  is  within  the  dimen- 
sions of  Coenia^  and  superficially  they  ap- 
pear very  similar.  However,  the  chaeto- 
taxy  characters  readily  distinguish  either 
and  set  the  Coenia  species  apart  from  any 
Paracoenia  subgenus. 

Description. —  Length  2.2-2.8  mm; 
dark  brown,  pollinose  to  subshining. 

Head.  Mesofrons  subshining  with  me- 
tallic reflections;  postocular  bristles  weak, 
at  most  slightly  larger  than  their  distance 
apart  as  base;  postorbital  bristles  usually 
weak;  pectinate  aristal  branches  various; 
bristles  of  face  comparatively  large, 
bristles  along  ventral  margin  subequal  in 
length  to  interfoveal  hump  height;  facial 
pruinosity  various;  chaetotaxy  of  head 
similar  to  Paracoenia  except  as  noted.  Eye 
relatively  large,  subcircular  although 
higher  than  wide;  width-to-height  ratio 
1:0.62;  height-to-length  ratio  1:0.95;  eye- 
width-to-face-length  ratio  1:0.3. 

Thoj-ax.  Dark  brown,  generally  con- 
colorous;  strength  of  pollinose  covering 
various.  Acrostichal  hairs  in  two  rows,  in- 
distinct in  some  species.  Three  or  four 
pairs  of  dorsocentral  bristles;  humeral 
bristles  various;  otherwise  chaetotaxy  as 
in  Paracoenia.  Wings  without  prominent 
costal  bristles,  at  most  weakly  developed 
on  dorsal  margin;  costal  vein  ratio  1:0.3; 
Mi+_.  vein  ratio  1:0.7.  Legs  uniformly 
dark  brown;  male  mesofemora  without 
posteroventral  comb.  Halters  as  in  diag- 
nosis. 

Abdomen.  Subshining  to  shining  dark 
brown;  male  with  five  visible  tergites; 
female  with  six  to  seven;  male  with  i\\e 
sternites;  fifth  sternite  narrowed  to  small 
strip;  spiracle  arrangement  as  in  Para- 
coenia. Male  postabdomen  with  surstvli 
closely  apposed  basally,  becoming  well 
se])arated  on  apical  2/3;  aedeagal  apo- 
deme  broad  to  narrowly  crescent-shaped; 
aedeagus  curved,  pointed  apically.  Ven- 
tral receptacle  with  small  operculum,  ex- 
tending process  C-shaped. 

Discussion.- —  Of  the  three  known  Co- 
enia species,  one  is  aj)parently  endemic 
to  the  Nearctic  Region,  a  second  is  limited 
to  the  Palearctic,  and  a  third  species  is 
Holarctic.  Cresson's  comments  regarding 
the  European  species  and  their  distinguish- 


March  1975 


MATHIS:    AMERICAN    EPHYDRIDAE 


83 


ing  characteristics  should  bo  consulted  for  tions  of  various  salts.    Dahl    (1959)    and 

species  separation  from  that  fauna.  Scheiring  and  Foote  (1973)  report  on  as- 

Species  of  Coenia  are  not  usually  asso-  sociations  of  Coenia  curvicauda  (Meigen) 

elated  with  aquatic  environments  that  are  with  mud  shore  habitats  and  to  a  lesser 

contaminated  or  contain  high  concentra-  extent  with  the  limnic  wrack. 

Key  to  Coenia  species 

1.    Male  genitalia  large,  exposed  ventrally;  epandrium  over  twice  as  long  as 

wide,  with  median  suture  - C.  curvicauda  (Meigen) 

Male  genitalia  more  compact,  not  exposed;   epandrium   less  than  twice   as 

long  as  wide,  without  median  suture  --..  C.  alpina  n.  sp. 


Coenia  alpina,  n.  sp. 

Fig.  10,  Map  2 

Di.'\GNosis. —  C.  alpina  most  closely  re- 
sembles C.  palustris,  a  Palearctic  species. 
It  is  distinguished  from  the  latter  by  dif- 
ferences in  male  genitalic  structures.  The 
surstyli  are  proportionately  shorter  to 
the  epandrial  length,  the  aedeagus  is 
deeper  and  Aentrall}'  rounded,  the  aede- 
agal  apodeme  is  longer,  and  the  hypandri- 
al  processes  are  thinner  and  project  mesad 
to  the  hypandrial  connection  with  the 
aedeagus.  The  females  of  both  species  are 
very  similar. 

Description. —  Length  2.1-2.6  mm; 
dark  brown  with  some  purplish  metallic 
reflections  dorsally. 

Head.  Interfoveal  hump  more  or  less 
prominent;  pruinose  face  brown;  pectin- 
ate branches  of  arista  long,  approximately 
three  times  aristal  width  at  base;  post- 
orbital  setae  not  developed  dorsally.  Eye- 
to-cheek  ratio  1:0.12;  width-to-height  ra- 
tio 1:0.6;  height-to-length  ratio  1:0.96; 
eye-width-to-face-length   ratio    1:0.33. 

Thorax.  Acrostichal  setae  weak,  in  two 
rows;  three  pairs  of  dorsocentral  bristles; 
humeral  bristles  absent;  halters  dark 
brown. 

Abdomen.  Subshining  to  shining  with 
some  purplish  reflections;  ventral  recep- 
tacle as  in  Fig.  lOd;  male  genitaha  as  in 
Fig.  10. 

Distribution. —  C.  alpina  has  been 
collected  in  Colorado  (Rabbit  Ears  Pass), 
in  the  Northwest  Territories  (Aklavik), 
and  in  Labrador   (Cartwright). 

Types. —  Male  holotype,  allotype,  and 
21  para  types  (8  males,  13  females).  Cart- 
wright,  Labrador,  29  June  1955,  E.  F. 
Cashman;  17  paratypes  with  same  data 
as  type  except  as  follows:  5  males,  3  fe- 
males, 3  July  1955;  4  males,  2  females,  2 
July  1955,  E.  E.  Sterns;  1  male,  6  August 


1955,  E.  E.  Sterns;  1  male,  12  August 
1955.  The  type,  allotype,  and  32  para- 
types will  be  deposited  wdth  the  Canadian 
National  Collection,  type  number  13435. 
A  male  and  female  paratype  also  will  be 
deposited  with  the  California  Academy  of 
Sciences,  the  U.S.  National  Museum  of 
Natural  History,  and  in  my  collection. 

In  addition  to  the  type  series,  I  have 
examined  31  specimens  of  this  species 
from  the  following  localities:  9  males,  17 
females,  Aklavik,  Northwest  Territories, 
May-August  1930  and  1931;  2  males,  3 
females,  Rabbit  Ears  Pass,  Colorado,  11 
June  1968,  S.  L.  W. 

Remarks. —  This  species  is  alpine, 
which  accounts  for  the  specific  name.  Al- 
though the  known  distribution  is  based 
on  minimal  data  that  is  rather  disjunct,  I 
feel  that  C.  alpina  is  distributed  through- 
out the  Rocky  Mountains  at  higher  ele- 
vations  and   across   northern   Canada. 

From  my  study,  I  liaAe  found  very 
httle  variation  except  for  minor  size  dif- 
ferences as  indicated  in  the  description. 
Otherwise,  C.  alpina  seems  to  be  a  very 
uniform  species. 

Coenia  curvicauda   (Meigen) 
Fig.  8,  Map  3 

Ephydra  curvicauda  Meigen,    1830,   Syst.   Beschr. 

6:116 
Coenia    curvicauda:    Macquart,    1835,    Hist.    Nat. 

Ins.  Dipt.  2:530 

Types. —  Cresson  (1930)  designated  a 
lectotype  for  this  species  from  material 
in  the  Naturhistorisches  Museum,  Wien. 
According  to  Cresson,  the  male  lectotype 
has  the  following  label  data:  "curvicauda 
Coll.  Winth,"  "curvicauda."  A  second 
male  specimen  with  similar  data  was  de- 
signated a  parat3^pe  b}'  Cresson. 

Diagnosis. —  C.  curvicauda  is  similar  to 
C.  alpina  and  to  C.  palustris  but  differs 


84 


GREAT    BASIN    NATURALIST 


Vol.  35,  No.  1 


from  either  by  the  well-developed  male 
postabdomen  which  })rotrudes  from  the 
venter  of  the  abdomen.  The  epandrium 
plus  surstjdi  are  over  twice  as  long  as 
the  epandrial  width,  and  the  epandrium 
is  divided  by  a  median  groove.  The  sur- 
styli  arms  are  longer  than  their  base,  the 
aedeagal  apodeme  is  slender  and  C-shaped 
in  profile,  and  the  aedeagus  is  broadly 
developed  basally  and  curves  forming  a 
J-shaped  structure. 

Description. —  Length  2.3-2.8  mm; 
dark  brown,  subshining  to  shining. 

Head.  Mesofrons  shining,  dark  brown; 
fronto-orbital  areas  subshining;  pectinate 
aristal  branches  subequal  to  third  anten- 
nal  segment  width;  face  lightly  pollinose, 
mostly  dark  brown;  chaetotaxy  as  in  C. 
alpina;  eye-to-cheek  ratio  1:0.11;  width- 
to-height  ratio  1:0.6;  height-to-length  ra- 
tio 1:1;  eye-W'idth-to-f ace-length  1:0.3. 

Thorax  and  abdomen  as  in  C.  alpina 
except  as  given  in  diagnosis. 

Specimens  examined. —   150 

Distribution. —  Coenia  cwvicauda  is 
a  Holarctic,  boreal  species.  In  the  Nearc- 
tic  region  it  is  found  in  Montana  (Big- 
fork)  and  east  through  the  Midwest  to 
the  Northeast.  It  extends  north  to  Alaska 
(Tonsina)  and  east  to  Quebec  (Cross 
Point) . 

Remarks. —  Although  I  did  not  ex- 
amine the  lectotype,  I  have  studied  Euro- 
pean specimens  of  this  species.  The 
aedeagus  in  some  specimens  does  not 
curve  apically  to  the  extent  found  in  Ne- 
arctic  specimens,  but  I  did  not  find  any 
other  major  differences.  I  consider  all 
specimens   I  examined   to   be  conspecific. 


Literature    Cited 

Aldrich,  J.  M.  1905.  A  catalogue  of  Nortli 
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Arten   Derselben.    Neue   Beitrage   zur   Kennt- 

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Berlin. 
M.\CQU.\RT,  J.     1835.     Histoire  Naturelle  des  In- 

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bung  der  bekannten  europaischen  zweiflugel- 

igen  Insekten.  vol.  5.  Hamm. 
Mitchell,    R.    M.    ,\nd    B.    L.    Redmond.     1974. 

Fine  structure  and  respiration  of  the  eggs  of 

two    ephydrid    flies     (Diptera:     Ephvdridae). 

Trans.  Amer.  Micros.  Soc.  93:113-118. 
Newm.\n,    E.     1838.     Entomological    notes.    Art. 

XL.  Ent.  Mag.,  pp.  372-402. 
Oliveir.\,  S.  J.      1954a.     Contribuicao  para  o  con- 

hecimento   do   genero   ''Dimecoenia"   Cresson. 

1916.  I.  ''Dimecoenia  lenii"  sp.  n.  encontrada 

no  Chile   (Diptera,  Ephydridae).  Rev.  Brasil. 

Biol.   14:187-194. 
.     1954b.     Contribuigao     para     o     conheci- 

mento  do  genero  ''Dimecoenia'"  Cresson,  1916, 

II.  Sobre  3  especies  novas  do  Brasil  (Diptera. 

Ephydridae).   Rev.   Brasil.   Biol.   14:269-278. 
.      1957.     Contribuigao  para  o  conhecimen- 

to  do  genero  ''Dimecoenia'  Cresson,  1916.  III. 

Sobre    uma    especie    nova    do    Estado    de    Sao 

Paulo.    Brasil     (Diptera.    Ephydridae).     Rev. 

Brasil.  Biol.   17:305-308. 
PoLHEMUs,    J.     T.     1967.     A    new    saldid     from 

California    (Hemiptera:    Saldidae).  Proc.  Ent. 

Soc.  Wash.   69:346-348. 
Rorineau-De-svoidy.    a.    J.  "B.     1830.     Essai    sur 

les     Myodaires.      Memoires     da      I'Academie 

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March  1975 


MATHIS:    AMERICAN    EPHYDRIDAF. 


85 


ScHEiRiNG.  J.  F.  AND  B.  A.  FooTE.  1973.  Hab- 
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eastern Ohio  (Diptear:  Ephydridao).  Ohio 
Jour.   Sri.   7^:152-166. 

Stenh.\mm.\r.  C.  18-H,  Forsok  till  gruppering 
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Vetensk.  Akad.  Handl.   1843:75-272. 

Steyskal,  George  C.  1970.  The  species  of  the 
genus  Dimecoenia  (Diptera:  Ephydiidae)  in 
America  north  of  Panama,  with  the  descrip- 
tion of  a  new  species.  Ann.  Ent.  Soc.  Amer. 
63:462-465. 

SxURTEyANT,    A.    H.    AND    M.    R.    WlIEEI.ER.        1954. 

Synopses  of  nearctic  Ephydridae  (Diptera). 
Trans.  Amer.  Ent.  Soc.  79:151-261. 

Walker,  F.  1849.  List  of  the  specimens  of  dip- 
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Museum.   Vol.   4.   pp.   689-1172. 

.     1853.     Insecta   Britannica.   Diptera.   Vol. 

II,  297  pp. 


WiEGERT,  R.  G.  and  R.  Mitciiei.e.  1973.  Ecol- 
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intersects  of  blue-green  algae,  grazing  flies 
(Paiaroenia,  Ephydridae)  and  water  mites 
{Part/iuniclla,  Hydrachnellae).  Hydrobiol- 
ogia  M: 25 1-271. 

WiRTir,  W.  W.  1948.  A  taxonomic  study  of 
Hawaiian  Ephydridae  (Diptera)  related  to 
Scalclla  Robineau-Desvoidy.  Proc.  Hawaiian 
Ent.   Soc.    13:277-304. 

—  -.  1965.  Ephydridae.  Pages  734-759  in 
Alan  Stone,  C.  W.  Sabrosky,  W.  W.  Wirth, 
R.  H.  Foote,  and  J.  R.  Coulson,  eds.  A  catalog 
of  the  diptera  of  America  north  of  Mexico. 
U.   S.   Dep.  Agr.  Handb.  No.  276. 

.      1970.     A  new  genus  and  species  of  shore 

fly  (Diptera.  Ephydridao)  from  southern  Pata- 
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.      1971.     The     brine     flies     of     the     genus 

Ephydra  in  NoiUi  America  (Diptera:  Eph- 
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ENVIRONMENTAL  FACTORS  IN  RELATION  TO  THE  SALT  CONTENT 
OF  SALICORNIA  PACIFIC  A  VAR.  UTAHENSIS' 

D.  J.  Hansen-  and  D.  J.  Weber 

ABSTII.A.CT. —  The  stability  of  the  salt  content  in  Salicornia  pacifica  Standi,  var.  utahensis  (Tide- 
strom)  Munz  in  relation  to  environmental  changes  was  investigated.  Salicornia  pacifica  communities 
have  a  characteristic  soil  pH  of  7.5  to  8.0  ±  0.2  and  a  constant  subsurface  soil  moisture  level  of  25  to 
35  percent.  The  ion  content  in  the  tissue  of  S.  pacifica  remained  constant  despite  increased  moisture 
stress  throughout  the  growing  season.  The  concentrations  of  the  salts  were  significantly  higher  in  the 
surface  soil  layers  than  in  the  subsurface  layers  around  the  roots.  Normal  metabolic  processes  in  the 
tissues  of  S.  pacifica  appear  to  occur  even  though  some  fluctuations  in  the  ionic  balance  and  concen- 
tration of  ions  in  the  plant  occur. 


Introduction 

Halophytic  plants  are  among  the  few 
species  of  higher  plants  that  can  with- 
stand high  sahne  soil  conditions  without 
detrimental  effects  (Waisel,  1972).  The 
is  mainly  associated  with  an  increase  in 
the  chloride  content  of  the  tissues  (Adri- 
ani,  1958;  Bernstein,  1961).  Steiner 
(1935)  found  that  chloride  ions  accounted 
for  67  to  88  percent  of  the  increase  of 
osmotic  potential  of  different  species  of 
salt  marsh  plants,  whereas  other  osmotic- 
ally  active  substances  had  only  a  negligi- 
ble effect.  Chloride  accounted  for  80  per- 
cent of  the  total  osmotic  potential  in  5^//- 
cornia  ambigua,  91  percent  in  S.  stricta, 
and  93  percent  in  S.  mucronata  (Arnold, 
1955).  Harward  and  McNulty  (1965), 
on  the  other  hand,  found  that  chloride 
accounted  for  less  than  49  percent  of  the 
osmotic  potential  in  S.  ?ubra. 

Osmotic  values  for  leaf  saps  of  a  num- 
ber of  herbaceous  halophytes  including  S. 
herbacea,  ranged  from  25  to  75  atm  when 
grown  in  salt  marshes  (Yabe  et  al.,  1956). 
Seasonal  changes  in  osmotic  potentials  in 
tissues  of  5".  rubra  ranged  from  40  atm  to 
over  100  atm  over  a  two-month  period 
(Harward  and  McNulty,  1965).  Higher 
osmotic  potential  values  have  been  record- 
ed for  a  number  of  halophytes;  for  ex- 
ample, Waisel  (1972)  reported  that  os- 
motic potentials  of  Rhizophora  and  Ain- 
cennia  leaves  reached  values  of  148  and 
163  atm,  respectively. 

According  to  Bowen  and  Rovira  (1966), 
salt  ions  can  cause  toxicity  in  the  follow- 
ing ways:  (1)  acting  as  antimetabolites, 
(2)  binding  or  precipitating  various  me- 
tabolites, (3)  catalyzing  rapid  decomposi- 
tion of  essential  elements,    (4)  combining 


with  cell  membranes  and  affecting  their 
permeability,  and  (5)  displacing  essential 
elements  but  failing  to  fulfill  their  func- 
tions. Bowen  and  Rovira  (1966)  sug- 
gested that  salt  injury  is  not  due  to  a 
direct  effect  of  the  salts  but  to  the  indirect 
effects  of  one  or  more  of  the  above  men- 
tioned metabolic  disturbances.  Waisel 
(1972)  singled  out  nitrogen  metabolism  as 
an  important  area  affected  by  high  salts. 
Salt-induced  growth  retardation  leads  to 
an  accumulation  of  unused  substances 
that  may  be  toxic  (Gauch  and  Eaton, 
1942). 

The  degree  of  salt  injury  or  tolerance 
of  plants  may  be  affected  by  a  nmnber 
of  environmental  factors  such  as  water- 
logged soils.  Some  plants,  however,  have 
adapted  to  waterlogged  conditions.  Sali- 
cornia foliosa  not  only  tolerates  water- 
logged conditions  but  appears  to  benefit 
from  them,  because  of  increased  capacity 
to  obtain  iron  under  these  conditions 
thereby  avoiding  chlorosis  (Adams,  1963). 
In  habitats  with  marked  fluctuations  in 
salt  concentration,  only  the  species  with 
high  osmotic  shock  resistance  {Salicornia 
sp.)  can  survive  (Levitt,  1972).  Some  of 
the  facultative  halophytes,  such  as  Sali- 
cornia rubra,  are  found  at  the  highest 
salinities  3^et  are  capable  of  growing  nor- 
mally in  low  to  nonsaline  environments 
(Ungar  et  al.,  1969). 

Since  species  of  Salicornia  are  among 
the  most  salt  tolerant  forms  of  higher 
plants  (Chapman,  1960),  S.  pacifica 
Standi,  var.  utahensis  (Tidestrom)  Munz., 
a  halophyte  conunon  to  inland  salt  playas 
of  northern  Utah,  was  selected  for  this 
investigation.  Due  to  the  lack  of  basic  in- 
formation   about    environmental    fluctua- 


'This  research   was  supported  in  part  by  NSF  Grant  No.   GB3I0G7    and    a 
-Department    of   Botany,    Brighani    Young   University,    Prove,    Utah  84602. 


It    from    Brigham    Young    University. 


86 


March  1975 


HANSEN,  WEBER:    SALICORNIA 


87 


tions  in  S.  pacifica  habitats  and  difficulties 
in  providing  an  artificial  environment 
that  parallels  the  natural  environment, 
investigations  were  undertaken  to  corre- 
late fluctuations  of  the  natural  environ- 
ment with  physiological  and  morphologi- 
cal characteristics  in  relation  to  salt  con- 
tent of  5'.  pacifica. 

Methods  and  Materials 

This  investigation  was  conducted  dur- 
mg  a  typical  growing  season  for  Salicor- 
nia  pacifica  (April  to  August  1972) .  Three 
sites  were  selected  to  compare  soil  and 
moisture  factors  in  relation  to  changes  in 
salt  content  of  S.  pacifica.  Sites  1  and  2 
were  six  miles  (9  km)  north  and  Site  3 
was  about  one-half  mile  (1  km)  east  of 
Goshen,  Utah.  Site  1  was  especially  se- 
lected because  it  was  an  ecotone  between 
a  stand  of  S.  pacifica  and  a  stand  of  Dis- 
tichlis  stricta.  It  was  hoped  that  the  data 
from  this  site  could  be  used  to  explain 
some  of  the  environmental  factors  respon- 
sible for  separating  the  Salicornia  com- 
munities from  the  Distichlis  communities. 

The  ground  at  Site  1  was  covered  by  a 
thin  layer  of  dried  algae  of  the  genus, 
Oscillatoria.    This  covering  formed  a  sur- 


face mulch  which  increased  the  moisture 
of  the  soil  surface.  Site  2  was  about  100 
m  west  of  Site  1  on  the  opposite  side  of  a 
large  drainage  basin.  Site  3  (Fig.  1 )  was 
6  miles  (9  km)  from  Sites  1  and  2  adja- 
cent to  a  natural  drainage  system.  Site 
3  was  selected  because  of  reduced  fluctua- 
tions in  soil  moisture  throughout  the  grow- 
ing season.  Soil  and  plant  samples  were 
taken  every  two  weeks  at  all  three  sites. 

A  standardized  hygrothermograph  unit 
was  used  to  continuously  monitor  temper- 
ature and  relative  humidity.  The  housing 
unit  for  the  hygrothermograph  was  loca- 
ted three  inches  above  the  ground  about 
100  m  from  Sites  1  and  2.  Measurements 
were  recorded  from  1  May  through  20 
September  1972. 

The  percentage  of  plant  cover  for  Sites 
1,  2,  and  3  was  determined  using  eight 
randomly  distributed  rectangular  1/4  m- 
quadrats.  Dry  weight  production  in  each 
site  was  determined  by  clipping  four  ran- 
domly distributed  rectangular  %.  m-  quad- 
rats. 

Soil  samples  were  taken  with  a  soil  core 
borer.  Each  core  measured  1  inch  (2:54 
cm)  in  diameter  and  was  extended  to  a 
depth  of  10  inches  (25.4  cm).  The  cores 
were  separated  to  provide  surface  (upper 


Fig.    1.     Site    3.    located   one   mile   east   of   Goshen,    Utah,    sliowi 
pacifica   (Photograph  courtesy  of  W.  M.  Hess.) 


(!    (if    Salicornia 


88 


GREAT   BASIN    NATURALIST 


Vol.  35,  No.  1 


2  inches  or  5  cm  of  the  core)  and  subsur- 
face (lower  6-10  inches  or  15-25  cm  of 
the  core)  soil  samples.  Five  to  eight  core 
samples  were  taken  at  each  site  and  pooled 
to  obtain  the  soil  sample.  The  samples 
were  placed  in  plastic  bags,  sealed,  and 
immediately  taken  to  the  laboratory  for 
analysis.  Samples  were  weighed  to  the 
nearest  one-hundredth  of  a  gram  and 
dried  in  an  oven  at  HOC  for  48  hours. 
The  samples  were  weighed  again,  and  the 
percentage  of  moisture  was  calculated. 

Measurements  of  the  soil  pH  were  de- 
termined from  saturated  soil  paste  sam- 
ples using  the  Sargent- Welch  pH  Meter, 
Model  PAX,  with  a  combination  electrode. 

Osmotic  potentials  of  soil  samples  were 
determined  from  saturated  soil  pastes  by 
freezing-point  depression  methods  accord- 
ing to  the  procedure  outlined  by  Hansen 
and  Weber  (1974). 

Soil  samples  (5  g)  were  leached  of  ex- 
changeable cations  and  anions  by  flush- 
ing with  four  25  ml  volumes  of  IN  CH3- 

C-NHo,  pH  7.0.  Each  volume  was  allowed 
to  drain  before  the  next  was  applied.  The 
cations  (sodium,  potassium,  magnesium, 
and  calcium)  were  detected  in  the  filtrate 
according  to  procedures  outlined  bv  Per- 
kin-Elmer  (1971)  on  the  Modef  290B 
Atomic  Absorption  Spectrophotometer. 
The  anion,  chloride,  was  detected  in  the 
filtrate  according  to  the  procedure  outlined 
by  Marius/Fiske  (1972)  using  a  Mari- 
us/Fiske  Chlor-o-counter. 

Osmotic  potential  measurements  for 
Salicornia  were  determined  by  freezing- 
point  depression  techniques  as  described 
by  Gary  and  Fisher  (1969,  1971)  and 
Fisher  (1972).  The  circuitry  was  modi- 
fied by  replacing  the  two  1.35  vdc  Hg  bat- 
tery cells  with  an  alkaline  9  vdc  battery. 
A  lOK  1-tum  potentiometer  was  installed 
to   compensate   for  voltage   drop   that  oc- 

T.ABLE  1.  Percentage  moisture,  dry  weight 
production,  and  percentage  cover  of  Salicornia 
pacifica  in  three  salt  desert  playas. 


Dry  weight 

Percentage 

production 

Percentage 

Site 

moisture 

g/m' 

cover 

1 

79.0 

73.1 

56.0  S.  pacifica 
2.5  S.  rubra 

80.6 

26.7 

7.6  S.  pacifica 
1.0/1.  occidentalis 

6 

80.6 

141.2 

65.0  S.  pacifica 
2.5  S.  rubra 

curred  with  time  and  usage.  Measure- 
ments were  made  at  each  internode  for 
several  plants.  Measurements  were  occa- 
sionally made  using  a  vapor  pressure  os- 
mometer. Model  301  Mechrolab  Inc.,  to 
verify  freezing-point  measurement  values. 
The  percentage  of  crude  protein  in  each 
plant  sample  was  determined  by  the  micro- 
Kjeldahl  method    (Horwitz,   1970). 

Results 
Climatic  and  Growth  Factors 

Continuous  hygrothermograph  monitor- 
ing of  climatic  factors  indicated  that  tem- 
peratures fluctuated  consistently  through- 
out the  growing  season.  The  average 
temperatures  gradually  increased  through 
May  and  June.  A  peak  was  reached  dur- 
ing July,  followed  by  a  gradual  decrease 
from  August  to  September.  The  average 
of  the  daily  highs  during  July  was  34C. 
The  hottest  temperature  in  July  was  38G 
on  12  July.  The  coolest  temperature  re- 
corded in  July  was  6G  on  25  July. 

Lowest  daily  means  (calculated  on  an 
hourly  basis)  for  relative  humidity  oc- 
curred from  15  July  through  20  August. 
A  series  of  rain  storms  increased  the  rela- 
tive humidity  during  late  August  and 
early  September.  Light  rain  was  also 
common  diu-ing  late  May  and  early  June. 
No  quantitative  data  concerning  the 
amount  of  rain  was  taken.  Relative  hu- 
midity reached  100  percent  every  night 
except  for  about  six  days  during  the  grow- 
ing season. 

The  dry  weight  of  the  standing  crop  of 
S.  pacifica  plant  tissue  was  the  highest  on 
Site  3  (141.2  g/m-)  and  the  lowest  on 
Site  2  (26.7  g/m-)  as  shown  in  Table  1. 
The  percentage  cover  was  also  highest  on 
Site  3  (65  percent)  and  lowest  on  Site  2 
(7.5  percent).  The  percentage  moisture 
of  the  plants  in  all  three  sites  was  about 
80  percent. 

Soil  Moisture 

The  subsurface  and  surface  soil  mois- 
ture readings  for  Site  2  were  highest  dur- 
ing June,  gradually  decreasing  through- 
out the  growing  season  (Fig.  2).  The  sub- 
surface soil  moisture  for  this  site  was  less 
than  the  surface  soil  moisture  from  1 
April  through  15  July.  This  may  have 
been  due  to  the  heavy  mulch  of  algae  on 
the  surface.  However,  from  mid-July 
through  the  rest  of  the  season,  this  trend 


March  1975 


HANSEN,  WEBER:    SALICORNIA 


89 


Fig.  2.  Soil  moisture  for  three  salt  desert 
playas  (Sites  1,  2,  and  3).  Solid  lines  indicate 
the  plot  of  percentage  values  for  surface  soil  sam- 
ples (upper  2").  Dashed  lines  indicate  the  plot 
of  percentage  values  for  subsurface  soil  samples 
(6-10"). 

was  reversed.  In  Sites  2  and  3  the  subsur- 
face soil  moisture  was  consistently  higher 
than  the  surface  soil  moisture  throughout 
the  growing  season,  except  for  one  week  in 
June  when  Site  2  had  an  increase  in  the 
surface  soil  moisture  due  to  rainfall.  Sur- 
face and  subsurface  soil  moisture  percent- 
ages at  Site  3  were  well  above  moisture 
percentages  of  Sites  1  and  2.  From  June 
through  July  soil  moisture  in  Site  3  was 
above  30  percent,  whereas  soil  moisture 
of  Sites  1  and  2  was  well  below  30  per- 
cent. This  was  probably  due  to  under- 
ground seepage  from  the  nearby  drainage 
system  adjacent  to  Site  3.  Soil  moisture 
was  lowest  in  all  three  sites  during  May. 

Soil  pH 

The  surface  soil  pH  of  the  three  sites 
was  generally  higher  and  fluctuated  more 
than  the  corresponding  subsurface  pH 
(Fig.  3).  The  subsurface  pH  of  these 
sites  changed  very  little  during  the  grow- 
ing season.  The  decrease  in  the  pH  of 
the  soil  surface  appeared  to  be  correlated 
to  the  amount  of  rainfall.  This  effect  was 
probably  due  to  the  transporting  of  solu- 
ble salts  into  the  subsurface  layers  by  the 
percolating  rain.  A  decrease  in  the  pH 
of  the  surface  generally  was  inversely 
proportional  to  the  increase  in  the  subsur- 
face pH. 

Ion  Content  of  the  Soil 
Site  1 

Osmotic  potential  measurements  of  the 
surface  soil  were  high  during  Jul}'  and 
September.   Osmotic  potential  values  were 


Fig.  3.  Soil  pH  as  recorded  for  three  salt 
desert  playas  (Sites  1,  2,  and  3).  The  solid  lines 
indicate  the  plot  of  tlie  pH  for  surface  samples 
(upper  2").  The  dashed  lines  indicate  the  plot 
of  the  pH  for  subsurface  samples   (6-10"). 


as  high  as  135  atm  (Fig.  4).  Osmotic 
potential  measurements  of  subsurface  soil 
samples  were  considerably  lower  than  os- 
motic potential  measurements  of  the  siu-- 
face.  Osmotic  potential  values  for  the  sub- 
surface soil  samples  gradually  increased. 
The  highest  value  reached  was  48  atm. 
This  value  was  recorded  on  20  September, 
when  the  study  terminated.  Values  dur- 
ing the  hottest  month  did  not  exceed  23 
atm. 

Individual  ion  analysis  of  soil  samples 
from  the  surface  (Fig.  5)  and  the  subsur- 
face layers  (Fig.  6)  showed  that  sodium 
and  chloride  were  the  two  ions  responsi- 
ble for  most  of  the  osmotic  potential.  The 
increasing  ion  accumulation  of  the  soil 
closely  paralleled  the  increase  in  osmotic 
potential  previously  described.    The  con- 


's— ik — to — it  «»"• 

LIST       I     UPTiytl* 


Fig.  4.  Osmotic  potential  of  saturated  soil 
paste  for  a  salt  desert  playa  (Site  1)  expressed  as 
atmospheres  pressure.  The  solid  line  indicates 
the  plot  of  osmotic  potential  values  for  the  sur- 
face soil  samples  (upper  2").  The  dashed  line 
indicates  the  plot  of  osmotic  potential  values  for 
tlie  subsurface  samples  (6-10"). 


90 


GREAT   BASIN    NATURALIST 


Vol.  35,  No.  1 


Fig.  5.  Ion  content  of  the  soil  for  the  surface 
layer  (upper  2")  of  a  desert  playa  (Site  1)  ex- 
pressed as  a  percentage  of  the  dry  soil.  The  solid 
line  indicates  the  percentage  of  chloride.  The 
dashed  line  indicates  the  percentage  of  sodium. 
The  dash-dotted  line  indicates  the  percentage  of 
calcium.  The  solid  vertical-barred  line  indicates 
the  percentage  of  potassium. 

centratioiis  of  sodimn  and  chloride  were 
highest  during  July  and  September.  The 
concentration  levels  of  calcium  and  potas- 
sium ions  were  fairly  constant  through- 
out the  season.  Concentration  values  sel- 
dom exceeded  0.5  percent  of  the  dry  soil 
weight  in  either  the  surface  or  the  subsur- 
face layers. 

Site  2 

Osmotic  potential  values  for  the  surface 
layer  samples  were  uniform  and  high 
(over  130  atm)  throughout  the  season 
with  the  exception  of  a  substantial  de- 
crease on  29  August,  when  values  dropped 
to  80  atm  (Fig.  7).  The  subsurface  values 
steadily  increased  throughout  the  growing 
season  to  a  high  of  about  96  atm  on  20 
September.  In  most  cases  osmotic  poten- 
tial values  of  the  subsurface  soil  samples 
were  50  to  100  atm  lower  than  osmotic 
potential  values  of  the  surface  soil  sam- 
ples.   Thus,  roots  near  the  surface  would 


SUBSURFACe     LAYEn 


Fig.  6.  Ion  content  of  the  soil  for  the  sub- 
surface layer  (6-10")  of  a  salt  desert  playa  (Site 
1)  expressed  as  a  percentage  of  the  dry  soil.  The 
solid  line  indicates  the  percentage  of  chloride.  The 
dashed  line  indicates  the  percentage  of  sodium. 
The  dash-dotted  line  indicates  the  percentage  of 
calcium.  The  solid  vertical-barred  line  indicates 
the  percentage  of  potassium. 


V) 


o 

0 


VZ       4T"6       k   I  lb      12     114     16 
APRIL*     MAY     '    JUNE    '     JULY     '      AUO 

OSMOTIC   POTENTIAL  OF  SOIL  PASTE 

Fig.  7.  Osmotic  potential  of  satui-ated  soil 
paste  for  a  salt  desert  playa  (Site  2)  expressed  as 
atmospheres  pressure.  The  solid  line  indicates  the 
plot  of  osmotic  potential  values  for  the  surface 
soil  samples  (upper  2").  The  dashed  line  indi- 
cates the  plot  of  osmotic  potential  values  for  the 
subsurface  soil  samples  (6-10"). 

be  in  an  environment  of  higher  moisture 
stress  than  deeper  roots. 

Ion  analysis  of  the  surface  layer  soil 
samples  showed  that  sodium  and  chloride 
ions  were  responsible  for  most  of  the  os- 
motic potential  of  the  soil  samples  (Fig. 
8).  The  calcium  ion  concentration  of  the 
surface  layer  for  this  site  was  consider- 
ably higher  than  for  Site  1.  The  subsur- 
face calcium  and  potassium  ion  concen- 
trations were  comparable  with  other  sites 
and  rarel}^  exceeded  0.5  percent  of  the 
soil  dry  weight  (Fig.  9).  The  combined 
ion  concentration  in  the  subsurface  layer 
increased  gradually  over  the  growing  sea- 
son and  reached  a  peak  of  about  6  per- 
cent  soluble   salts  on  20   September. 

Site   3 

Osmotic  potential  measiu-ements  of  soil 
samples  on  this  site  gradually  increased 
in  both  the  surface  and  subsurface  layers. 
Measurements  were  highest  on  20  July 
and  20  September  (Fig.  10)  and  were 
comparable  with  measurements  from  Site 
2,  both  of  which  were  considerably  higher 
than  Site  1 . 

Chloride  ion  content  from  the  surface 
layer  fluctuated  considerably,  but  the  so- 
dium content  was  more  stable  (Fig.  11). 
The  major  increase  in  osmotic  potential 
of  the  subsurface  layer  from  20  July 
through  20  September  was  due  chiefly  to 
the  chloride  ion  content.  The  calcium  ion 
concentration  level  of  "this  site  was  con- 
siderably higher  than  Site  1  and  not  as 
high  as  Site  2.  The  potassiimi  ion  con- 
centration level  was  less  than  0.5  percent 


March  1975 


HANSEN,  WEBER:    SALICORNIA 


91 


~!2  WEEKS 
SEPTEHBEK 


Fig.  5.  Ion  content  of  the  soil  for  the  surface 
layer  (upper  2")  of  a  salt  desert  playa  (Site  2) 
expressed  as  a  percentage  of  the  dry  soil.  The 
solid  line  indicates  the  percentage  of  chloride. 
The  dashed  line  indicates  the  percentage  of  sodi- 
um. The  dash-dotted  line  indicates  the  percentage 
of  calcium.  The  solid  vertical-ban-ed  line  indicates 
the  percentage  of  potassium. 


APRILI  U*r  I       JUNE  I  JULY  I  AUGUST        I     SEP 


Fig.  9.  Ion  content  of  tlie  soil  for  the  sub- 
surface layer  (6-10")  of  a  salt  desert  playa  (Site 
2)  expressed  as  a  percentage  of  the  dry  soil.  The 
solid  line  indicates  the  percentage  of  chloride. 
Th  dashed  line  indicates  the  percentage  of  sodium. 
The  dash-dotted  line  indicates  the  percentage  of 
calcium.  The  solid  vertical-barred  line  indicates 
the  percentage  of  potassium. 


and  did  not  fluctuate  significantly.  The 
concentration  level  gradually  increased 
to  a  peak  of  0.5  percent  on  20  July  and 
then  gradually  decreased  to  0.3  percent 
on  20  September.  Concentrations  of  so- 
dium and  chloride  ions  in  the  subsurface 
soil  samples  gradually  increased  (Fig.  12). 
Calcium  and  potassium  ion  concentra- 
tions in  this  layer  remained  low  and  rela- 
tively constant  through  the  growing  sea- 
son; these  ion  concentrations  were  com- 
parable to  ion  concentrations  of  the  other 
sites. 

Ion  Content  in  Salicornia 
Site  1 

The  chloride  ion  content  in  Salicornia 
tissues  remained  constant  throughout  most 
of  the  growing  season  but  increased 
slightly  in  April  (Fig.  13).  This  increase 
was  followed  by  a  stabilization  of  the  con- 


centration at  about  12  percent  of  the  dry 
weight.  Sodimn  ion  concentration  gradu- 
ally increased  from  about  4.2  to  9  percent, 
while  potassium  ion  concentration  de- 
creased throughout  the  season  from  2.2 
to  1  percent. 

Site  2 

The  chloride  content  gradually  in- 
creased from  14.2  to  16.1  percent  at  the 
end  of  the  growing  season  (Fig.  14).  In- 
crease in  the  sodium  content  was  ]:)ropor- 
tional  to  the  increase  in  the  chloride  con- 
tent and  was  10.2  percent  at  the  end  of 
the  growing  season.  The  potassium  ion 
concentration  increased  from  4.5  to  7.5 
percent  and  was  closely  correlated  to  de- 
creases in  the  sodium  ion  concentration. 


n — •  I  'o  ,,«    |i — ^ — f 

I      JUNE  I  JUtT  I         AUOUST        I 


~iz  WEEKS 


Fig.  10.  Osmotic  potential  of  saturated  soil 
paste  for  a  salt  desert  playa  (Site  3)  expressed  as 
atmospheres  pressure.  The  solid  line  indicates  the 
plot  of  osmotic  potential  values  for  the  surface 
5oil  samples  (upper  2").  The  dashed  line  indi- 
cates the  plot  of  osmotic  potential  values  for  the 
subsurface  samples   (6-10"). 


SEPTEHSEI) 


Fig.  1 1 .  Ion  content  of  the  soil  for  the  sur- 
face layer  (upper  2")  of  a  salt  desert  playa  (Site 
3)  expressed  as  a  percentage  of  the  dry  soil.  The 
solid  line  indicates  the  percentage  of  chloride. 
The  dashed  line  indicates  the  percentage  of  so- 
dium. The  dash-dotted  line  indicates  the  percent- 
age of  calcium.  The  solid  vertical-barred  line 
indicates  the  percentage  of  potassium. 


92 


GREAT   BASIN    NATURALIST 


Vol.  35,  No.  1 


Fig.  12.  Ion  content  of  the  soil  for  the  sub- 
surface layer  of  a  salt  desert  playa  (Site  3)  ex- 
pressed as  a  percentage  of  the  diy  soil.  The  solid 
line  indicates  the  percentage  of  chloride.  The 
dashed  line  indicates  the  percentage  of  sodium. 
The  dash-dotted  line  indicates  the  percentage  of 
calcium.  The  solid  vertical-barred  line  indicates 
the  percentage  potassium. 


irr 


Fig.  13.  Ion  content  in  Salicornia  pacifica 
(Site  1)  expressed  as  a  percentage  of  the  dry 
weight.  The  solid  line  indicates  tlie  plot  of  the 
percentage  values  for  chloride  ions.  The  dash- 
dotted  line  indicates  the  plot  of  the  percentage 
values  for  sodium  ions.  The  dashed  line  indicates 
the  plot  of  the  percentage  values  for  potassium 


Site  3 

The  ion  content  in  Salicornia  tissues  in 
this  site  showed  a  decrease  in  chloride,  so- 
dium, and  potassium  throughout  the  sea- 
son (Fig.  15).  The  chloride  content 
ranged  from  16  percent  on  1  May  to  12.4 
percent  on  20  September.  The  sodium 
content  ranged  from  1 1  percent  on  1  May 
to  7.6  percent  or  20  September.  The  po- 
tassium content  ranged  from  4.8  percent 
on  1  May  to  2.1  percent  on  20  September. 

Osmotic  Potentials  in  Salicornia 

Early  in  the  investigation  it  was  dis- 
covered that  each  internode  of  the  plant 
was  osmotically  different  from  other  in- 
temodes  of  the  same  plant.  Freezing- 
point  depression  measurements  showed 
that  osmotic  potential  values  increased 
from  the  base  of  the  aerial  shoot  upward 
to  the  top  of  the  plant  (Table  2).  The 
lowest  intemodes  near  the  base  generally 
had  a  lower  osmotic  potential  than  inter- 
nodes  near  the  middle  of  the  plant.  In 
some  cases  there  was  as  much  as  15  atm 


difference  between  two  regions  (inner 
and  outer)  of  the  cortex,  although  usually 
the  difference  was  only  about  5  atm 
(Table  2).  Because  of  the  complexity  of 
such  differences  in  osmotic  potential  read- 
ings, only  periodic  measurements  were 
made  on  the  plants.  The  measurements 
ranged  from  a  low  of  about  80  atm  in 
May  to  about  130  atm  to  150  atm  in  mid- 
July  and  August.  However,  the  average 
was  about  90  to  100  atm. 

Crude  Protein  Analysis 

Crude  protein  analysis  showed  a  gradu- 
al decrease  in  the  total  crude  protein  con- 
tent in  the  plants  from  all  three  sites  from 
1  April  through  30  July.  Crude  protein 
decreased  from  20  to  8  percent  on  a  dry 
weight  basis.  From  30  July  through  20 
September  the  content  remained  about  8 
percent. 

Phenology  and  Morphology 

Growth  of  5".  pacifica  usually  begins 
with  development  of  the  subterranean  or 

ION  CONTENT     IN     SALICORNIA 


.... 


-iSf 


Fig.  14.  Ion  content  in  Salicornia  pacifica 
(Site  2)  expressed  as  a  percentage  of  the  dry 
weight.  The  solid  line  indicates  the  plot  of  the 
percentage  values  for  chloride  ions.  The  dash- 
dotted  line  indicates  the  plot  of  the  percentage 
values  for  sodium  ions.  The  dashed  line  indi- 
cates the  plot  of  the  percentage  values  for  po- 
tassium ions. 


IST 


Ho      n  wtEW 


Fig.  15.  Ion  content  in  Salicornia  pacifica 
(Site  3)  expressed  as  a  percentage  of  the  dry 
weight.  The  solid  line  indicates  the  plot  of  the 
percentage  values  for  chforide  ions.  The  dash- 
dotted  line  indicates  the  plot  of  the  percentage 
values  for  sodium  ions.  The  dashed  line  indicates 
the   plot   of   the   percentage  values   for  potassium 


March  1975 


HANSEN,  WEBER:    SALICORNIA 


93 


Table  2.  Osmotic  potentials  of  four  different  5.  pacifica  plants  showing  the  values  (atm)  of  the 
outer  and  inner  "cortex"  tissues  from  the  bottom  of  the  shoot  (Node  1)  to  the  top  of  the  shoot  (Nodle 
5). 


Node 


Plant  A 
Inner     Outer 


Plant  B 
Inner     Outer 


Plant  C 
Inner     Outer 


Plant  D 
Inner     Outer 


1 

76 

76 

81 

81 

73 

73 

84 

89 

2 

69 

89 

57 

75 

61 

80 

64 

74 

3 

71 

86 

93 

86 

83 

96 

71 

74 

4 

67 

67 

96 

106 

68 

83 

73 

79 

5 

106 

130 

96 

106 

103 

108 

84 

88 

6 

89 

92 

93 

108 

7 

106 

112 

near-subterranean  axillary  buds  of  older 
shoots  that  start  to  grow  in  March  and 
emerge  in  late  April  or  early  May.  Flow- 
ering occurs  in  mid-June  and  is  generally 
completed  in  July. 

By  late  August  lower  internodes  begin 
to  wither  and  die  sequentially  from  the 
base  of  the  shoot  to  the  tip  (Fig.  16).  Oc- 
casionally internodes  may  become  injured 
or  for  other  reasons  may  wither  and  die. 
The  central  stele  continues  to  function  in 
a  normal  manner.  Seed-producing  inter- 
nodes are  the  last  to  die  (Fig.  16).  It  is 
not  known  whether  the  internodes  die 
strictly  by  senescence,  whether  accumu- 
lation of  additional  salts  in  these  areas 
causes  death,  or  whether  death  is  related 
to  some  other  phenomenon. 

The  seeds  are  shed  in  October  and  No- 
vember. A  few  seeds  adhere  to  the  mother 
plant  until  rain  or  heavy  snows  separate 
them.  Seeds  are  protected  by  a  bulky, 
lightweight  seed  coat  that  aids  in  dis- 
persal and  absorption  of  moisture. 
Seasonal  Variations  of  the  Environment 

Temperatures  were  highest  during  July, 
while  relative  humidity  was  lowest  dur- 
ing July  and  August.  These  high  tem- 
peratures would  cause  increased  moisture 
stress. 

The  toxicity  of  salts  increases  with  the 
temperature,  according  to  Kaho  (1926) 
and  Waisel  (1972).  The  high  moisture 
content  of  the  soil  and  the  subsequent 
evaporation  may  have  had  a  cooling  ef- 
fect upon  the  temperature  of  the  soil. 
Both  lower  temperatures  and  high  rela- 
tive humidity  would  have  a  favorable 
effect  upon  plant  survival. 

At  all  three  sites  there  was  an  increase 
in  moisture  stress  throughout  the  season 
primarily  due  to  increases  in  salt  from 
underground  sources  and  slight  decreases 
in  soil  moisture.  Soil  moisture  of  25  to 
35  percent  is  perhaps  an  important  fac- 


tor in  maintaining  S.  pacifica  in  the  en- 
vironment. 

Sodium  and  chloride  were  the  two  prin- 
cipal ions  responsible  for  increases  in  os- 
motic potential  of  soil  samples.  They  were 
also  responsible  for  85  to  95  percent  of 
the  osmotic  potential  of  S.  pacifica.  Har- 
ris (1915)  considered  NaCl  the  most  toxic 
of  several  soluble  salts  but  concluded  that 
salt  mixtures  were  not  as  toxic  in  soils  as 
in  culture  solutions. 

The  salt  concentration  of  the  soil  sur- 
face layers  was  considerably  higher  than 
the  salt  concentration  in  the  rooting  lay- 
ers. Soil  surface  layers  were  often  en- 
crusted with  deposits  of  white  salt,  which 
gave  the  impression  that  the  plants  were 
surrounded  by  extremely  high  concentra- 
tions of  salt  when,  in  fact,  the  rooting 
zones  or  layers  were  only  moderately  sa- 
line. This  observation  was  also  reported 
by  Wiesel  (1972).  The  wicking  action 
caused  by  evaporation  of  water  resulted 
in  salt  crystallization  at  the  surface,  which 
removed  salts  from  the  rooting  layers  and 
provided  a  unique  environmental  niche 
for  S.  pacifica  to  occupy.  Succulents  such 
as  Salicornia  appear  to  lose  the  function 
of  portions  of  their  fleshy  cortex  and 
leaves  (Fig.  16),  yet  the  vascular  system 
continues  to  function  for  the  upper  fleshy 
tissue  (Fig.  16).  The  thick  cuticle  and 
waxy  layer  of  the  plants  and  the  active 
phellogen  of  the  central  stele  appear  to 
protect  the  upper  shoots  from  the  more 
severe  environmental  stresses. 

The  soil  pH  was  relatively  stable 
throughout  the  growing  season  in  all  three 
sites.  The  surface  soil  pH  of  each  site 
was  about  one-half  pH  unit  higher  than 
the  subsurface  pH.  Decreases  in  the  pH 
of  the  surface  layers  were  proportional 
to  increases  in  pH  of  the  subsurface  layer. 
Conversely,  increases  in  the  pH  on  the 
surface    layer    were    proportional    to    de- 


94 


GREAT   BASIN   NATURALIST 


Vol.  35,  No.  1 


Fig.  16.     Shoots  of  Salicornia  pacifica  showing  healthy  fleshy  portions  of  stem  above  dead  fleshy 
sections  of  cortex. 


creases  in  the  pH  of  the  subsurface  layer. 
Calcium  ion  concentrations  in  the  surface 
layers  were  considerably  higher  at  Sites 
2  and  3  than  at  Site  1.  Soil  pH  at  Sites 
2  and  3  was  significantly  lower  than  at 
Site  1.  The  pH  may  have  been  indirectly 
affected  by  the  calcium  concentration  or 
may  haAe  been  related  to  the  algal  cover- 
ing of  the  soil  surface  in  Site  1. 


Ecological  Interactions  and 
Plant  Distribution 

There  appear  to  be  two  groups  of 
thought  explaining  the  restriction  of  Sali- 
cornia to  a  saline  environment.  New- 
wohner  (1938)  stated  that  halophytes  fail 
to  succeed  in  fresh-water  habitats  because 
of  competition  with  other  species.  This 
idea  was  supported  by  Stalter  and  Batson 
(1969),  who  stated  that  survival  and 
growth  rate  data  of  transplanted  salt 
marsh  vegetation  suggest  that  several 
species  of  halophytes  (S.  virginica  includ- 
ed) can  tolerate  conditions  not  found  in 
their  usual  zones.  Montfort  et  al.  (1927), 
on  the  other  hand,  claimed  that  this  ex- 
planation of  plant  distribution  must  be 
displaced  by  the  concept  of  "direct  ecolog- 


ical salt  action."  He  pointed  out  that  S. 
herbacea  has  an  optimum  growth  between 
1.5  and  3  percent  salt.  Webb  (1966)  al- 
so pointed  out  that  Salicornia  plants  grown 
without  salt  soon  die.  Waisel  (1972) 
stated  that  the  proportion  of  chlorides  to 
sulfates  and  the  total  salt  content  were 
important  in  determining  distribution.  For 
example,  S.  herbacea  was  more  sensitive 
to  a  high  proportion  of  sulfates  than  was 
Aster  tripolium. 

The  fact  that  Site  1  is  an  ecotone  be- 
tween S.  pacifica  and  Distichlis  stricta 
suggests  that  differences  in  this  site  com- 
pared with  the  other  two  sites  would 
provide  information  regarding  some  par- 
ameters of  S.  pacifica.  Distichlis  stricta 
in  this  site  appears  to  tolerate  a  higher 
pH  than  S.  pacifica  and  may  survive  in 
pH  values  from  8.0  to  9.0.  The  soil  pH 
apparently  affects  the  availability  of  nu- 
trients. At  pH  values  over  7  the  avail- 
ability of  manganese,  iron,  copper,  molyb- 
denum, and  zinc  declines  (Buckman  and 
Brady,  1969).  At  high  pH  values  phos- 
phorus forms  insoluble"  complexes  of  cal- 
cium that  are  unavailable  to  the  plants 
(Buckman  and  Brady,  1969).  Levitt 
(1972)    also  pointed  out  that  Allenrolfea 


March  1975 


HANSEN,  WEBER:    SALICORNIA 


95 


occidentalis  and  S.  suhtcrminalis  are 
highly  resistant  to  salt  but  sensitive  to  al- 
kalinity. A  change  from  saline  to  sodic 
soils  may  kill  these  plants.  Site  3,  where 
S.  pacifica  cover  and  production  were  the 
highest,  had  an  average  pH  of  7.7  and  a 
moisture  content  of  32  percent.  At  Site  1 
reduction  in  the  soil  moisture  and  the  in- 
crease in  the  total  concentration  of  salts 
of  the  subsurface  soil  layers  during  the 
season  apparently  also  favored  growth  of 
D.  stricta  over  S.  pacifica.  Distichlis  stric- 
ta  appears  to  be  a  better  competitor  for 
low  soil  moisture  than  S.  pacifica.  It 
seems  probable  that  during  the  initial 
adaptation  of  Salicornia  to  the  saline  en- 
vironment competition  played  a  major 
role  in  determining  plant  distribution.  As 
natural  selection  of  Salicornia  occurred, 
adaptations  allowing  the  plant  to  maintain 
itself  in  an  optimal  salt  concentration 
were  reinforced.  Plasticity  could  be  re- 
duced and  that  portion  of  the  gene  pool 
lost  that  allowed  the  plant  to  move  back 
into  non-saline  areas.  As  such  selective 
forces  would  be  localized,  it  would  be  im- 
proper to  generalize  and  say  that  all  spe- 
cies of  Salicornia  have  optimiun  growth  at 
high  salt  concentration.  The  variations 
in  salt  concentrations  that  yield  optimum 
growth  of  Salicornia  would  bear  this  out 
(Halket,  1915;  Webb,  1966;  Levitt,  1972; 
Waisel,  1972). 

The  Nature  of  the  Fleshy  Stem 

Anatomical  investigations  with  light 
microscopy  indicate  that  as  internodes 
mature  or  become  injured  a  phellogen 
that  produces  fibers  and  suberized  cells  is 
formed  in  the  central  stelar  region  (un- 
published results).  These  cells  seal  off 
the  conductive  tissues  from  water  and 
nutritive  loss  as  the  internodes  wither  and 
die.  This  development  would  protect  the 
plant  when  the  salt  and  moisture  stresses 
are  the  greatest  near  the  surface  in  July 
and  August. 

Osmotic  potential  measurements  of  the 
plants  indicated  that  each  internode  is 
probably  osmotically  independent  from 
other  internodes.  This  would  allow  the 
plant  to  survive  in  the  environment  if  a 
few  of  the  internodes  were  attacked  and 
destroyed  by  insects  or  disease.  Hill 
(1908)  showed  that  osmotic  adaptation 
differs  not  only  between  species  but  also 
between  organs  and  even  between  cells. 


In  his  investigation  root  hairs  varied 
greatly  in  this  respect,  even  in  the  same 
individual  plant.  Measurements  of  freez- 
ing-point depression  of  S.  pacifica  inter- 
nodes indicated  that  at  least  two  areas  of 
osmotically  different  tissues  were  present 
within  one  internode,  corresponding  to 
the  inner  "cortex"  and  outer  chloren- 
chyma  tissues.  Osmometer  measurements 
on  dissected  portions  of  these  tissues  indi- 
cated that  the  chlorenchyma  tissue  had  a 
higher  osmotic  potential.  Scholander  et 
al.  (1966)  suggested  that  xylem  sap  of 
S.  pacifica  may  contain  very  little  salt  as 
freezing-point  depression  of  S.  pacifica 
sap  was  found  to  approach  freezing-point 
depression  of  pure  water.  Since  chloride 
ions  were  equally  distributed  in  these  two 
tissues  (unpublished  data),  the  difference 
was  probably  due  to  sodium  ions  or  sug- 
ars and  other  organic  molecules  that  were 
produced  by  chloroplasts  in  the  immediate 
vicinity.  Steiner  (1935)  has  shown  that 
chloride  ions  account  for  67  to  88  percent 
of  the  increase  in  osmotic  potential  of  dif- 
ferent species  of  salt  marsh  plants,  where- 
as other  osmotically  active  substances  had 
only  negligible  effects.  This  increase  in 
chloride  was  shown  to  account  for  in- 
creases in  the  osmotic  potential  of  the  en- 
tire plant.  This  does  not  rule  out  the  pos- 
sibility that  differences  in  osmotic  poten- 
tials of  individual  tissues  might  be  due 
to  other  osmotically  active  substances. 
Increase  in  the  chloride  and  sodiimi  con- 
tent of  the  succulent  tissues  appears  to 
be  the  primary  means  of  osmotic  adapta- 
tion whereby  the  plant  can  survive  under 
increased  salt  and  moisture  stress.  Sodium 
and  chloride  ions  account  for  85  to  95 
percent  of  the  osmotic  potential  of  S. 
pacifica.  These  results  agree  with  the 
findings  of  Harward  and  McNult}'  (1965) 
and  Scholander  et  al.  (1966).  Ion  analysis 
of  S.  Pacifica  tissues  in  Site  1  indicated 
that  increases  in  osmotic  potential  were 
due  to  increases  in  Na+  but  not  CI".  In- 
creases in  osmotic  potential  at  Site  2  were 
due  to  both  Na+  and  C1-.  However,  at  Site 
3  there  was  a  decrease  in  both  Na"^  and 
CI".  As  the  concentration  of  the  soil  salts 
increased,  causing  an  increase  in  moisture 
stress,  osmotic  potential  of  the  plant  also 
increased.  This  increase  was  due  chiefly 
to  ions  other  than  Na+  and  CI"  or  sugars 
and  other  organic  molecules. 

Ion     analysis     of     S.     pacifica     tissues 
showed  that  the  chloride  ion  was  present 


96 


GREAT   BASIN    NATURALIST 


Vol.  35,  No.  1 


in  higher  concentrations  than  sodium  or 
potassiiim  ions.  Azizbekova  and  Babaeva 
(1970)  found  that  the  amount  of  absorbed 
Na+,  CI",  and  Mg"^""  in  Salicornia  increased 
with  increasing  salt  concentrations. 

During  the  period  of  greatest  moisture 
stress,  from  July  through  September,  the 
percentage  of  crude  protein  in  S.  pacifica 
for  all  three  sites  remained  constant  at 
about  8  percent  of  the  dry  weight.  The 
percentage  ion  content  of  sodium,  potas- 
sium, and  chloride  also  remained  relative- 
ly constant  throughout  the  growing  sea- 
son, even  when  environmental  stresses 
fluctuated  considerably. 

The  metabolism  of  S.  pacifica  func- 
tioned even  with  changes  in  the  ionic 
balance  and  concentration  of  ions  in  the 
plant.  The  changes,  however,  were  gradu- 
al. 

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Halket,  a.  C.  1915.  The  effect  of  salt  on  the 
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Hansen,  D.  J.,  and  D.  J.  Weber.  1974.  Ap- 
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Levitt,  J.  1972.  Responses  of  plants  to  environ- 
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1961.  Plant  communities  of  saline  soils  at 
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82:564-577. 

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demic Press,  New  York. 

Webb,  K.  L.  1966.  NaCl  effect  on  growth  and 
transpiration  in  Salicornia  bigelovii,  a  salt 
marsh  halophyte.  Plant  Soil  24:261-265. 

Yabe,  a.,  G.  Kuse,  T.  Mur.\ta,  and  H.  Takada. 
1965.  Physiology  of  halophytes  I.  The  os- 
motic value  of  leaves  and  the  osmotic  role  of 
each  ion  in  cell  sap.  Physiol.  Ecol.   13:25-33. 


NEW  RECORDS  OF  STONEFLIES  (PLECOPTERA)  FROM  NEW  MEXICO^ 


Bill  P.  Stark ,=  Theodore  A.  Wolff '  and  Arden  R.  Gaufin' 


Abstract. —  Distributional  data  are  presented  for  30  species  of  New  Mexico  Plecoptera  including  8 
species  new  to  the  state  list.  Previouslj',  many  of  the  included  species  were  known  from  a  single  lo- 
cation in  the  state.    A  revised  checklist  of  46  confirmed   species  is  presented  for  the  state. 


Distribtitional  data  on  the  New  Mexico 
stonefly  fauna  have  accumulated  slowly 
because  of  the  isolated  nature  of  much 
potential  stonefly  habitat.  Records  of  win- 
ter-emerging forms  are  particularly  mea- 
ger, with  three  sjiecies  of  Capniidae  listed 
for  New  Mexico  in  a  review  of  southwest- 
ern stoneflies  by  Stewart  et  al.  (1974). 
Two  species  were  recorded  from  a  single 
location  and  the  other  from  two  locations 
in  the  state. 

In  this  paper  we  report  eight  additional 
species,  Malenka  flexura  (Claassen),  Pod- 
mosta  delicatula  (Claassen),  Zapada  haysi 
(Ricker),  Capnia  confusa  Claassen,  Cap- 
nia  gracilaria  Claassen,  Paraleuctra  rick- 
eri  Nebeker  and  Gaufin,  Diura  knowltoni 
(Frison),  and  Isogenoides  zionensis  Han- 
son; new  distributional  data  for  species 
previously  reported  are  also  given.  Two 
of  us  (Stark  and  Wolff)  collected  all  ma- 
terial unless  otherwise  noted. 

The  authors  thank  M.  R.  Cather  for 
providing  records  from  her  personal  col- 
lection, and  R.  W.  Raumann  for  sending 
records  from  the  United  States  National 
Museum. 

Taenionema  nigripennis  (Ranks). — New 
Records:  Sandoval  Co.,  Santa  Clara  Can- 
yon, Wem  Povi  Pond,  elev.  6,900',  20-10- 
73,  nymphs.  Santa  Fe  Co.,  Rig  Tesuque 
Creek,  Rig  Tesuque  Campground,  elev. 
9,700',  21-IV-73,  nymphs;  Rio  Enmidio, 
Hyde  Park  Ski  Rasin,  lO-VI-74,  1    d^. 

Malenka  coloradensis  (Ranks). — New 
Records:  Catron  Co.,  Willow  Creek,  Wil- 
low Creek  Campground,  11-VT74,  2  d"  1 
?  .  Sandoval  Co.,  Tschicoma  Pond,  Santa 
Clara  Canyon,  13-VI-74,  3   d'   9    9  . 

Malenka  flexura  (Claassen). — Distribu- 
tion: Taos  Co.,  West  Fork  Red  River, 
A^heeler  Peak  Wilderness  Area,  10  miles 
iouth  of  Red  River,  9-VI-74,  25  c^   14   9  • 

Podmosta  delicatula  (Claassen). — Dis- 
tribution: Rio  Arriba  Co.,  Rio  Chama, 
Chama,  elev.  7,870',  8-VT74,  9  d"   19    9  . 


Taos  Co.,  Red  River,  2  miles  east  of  Red 
River,  9- VI- 74,  1    9  . 

Prostoia  hesametsa  (Ricker). — New 
Records:  Santa  Fe  Co.,  Rio  Santa  Cruz, 
near  Espanola,  elev.  5,600',  17-III-74,  1 
d" ;  Rig  Tesuque  Creek,  Rig  Tesuque 
Campground,  elev.  9,700',  17-III-74,  ex- 
uvium. 

Zapada  cinctipes  (Ranks). — New  Rec- 
ords: Sandoval  Co.,  Santa  Clara  Canyon, 
Wem  Povi  Pond,  elev.  6,900',  20-111-73, 
1  9 .  Santa  Fe  Co.,  Rig  Tesuque  Creek, 
Rig  Tesuque  Campground,  elev.  9,700', 
21-IV-73,  5   d'   10    9. 

Zapada  frigida  (Claassen). — New  Rec- 
ords: Santa  Fe  Co.,  Rig  Tesuque  Creek, 
Rig  Tesuque  Campground,  elev.  9,700', 
10-VT74,  1   d. 

Zapada  haysi  (Ricker). — Distribution: 
Lincoln  Co.,  North  Fork  Ruidoso  River, 
17-V-72,  S.  M.  Fiance,  1  9,  nymphs. 
Santa  Fe  Co.,  Rig  Testique  Creek,  Rig 
Tesuque  Campground,  elev.  9,700',  10- 
VT74,  2  d"  5  9  ;  same  location,  21-IV-73_, 
nymphs;  Rio  Enmidio,  Hyde  Park  Ski 
Rasin,  10-VT74,  elev.  10,560',  2  d'  7  9  . 
Taos  Co.,  Wheeler  Peak,  15  to  25-VT60, 
Rurks  and  Kinzer,  1  d"  1  9  ;  4  miles 
north  of  Arroyo  Seco,  22-VI-61,  S.  G. 
Jewett,  Jr.,  2  d"  1  9  ;  Rio  Hondo,  Taos 
Ski  Valley,  22-in-67,  R.  W.  Raumann, 
nym])hs;  Rio  Trampas,  above  El  Valle, 
lO-VI-74,  2  c^  8  9  ;  West  Fork  Red  River, 
Wheeler  Peak  Wilderness  Area,  10  miles 
south  of  Red  River,  9-VI-74,  3  d"  4   9  . 

Capnia  confusa  Claassen. — Distribtttion: 
Rio  Arriba  Co.,  Rio  Chama,  Chama,  elev. 
7,870',  19-IV-73,  13  cf  9  9  .  Sandoval  Co., 
Santa  Clara  Canyon,  Wem  Povi  Pond, 
elev.  6,900',  20-IIT73,  2  d-  San  Miguel 
Co.,  Pecos  River,  Tererro,  elev.  7,600',  21- 
IV-73,  26  d"  10  9  .  Taos  Co.,  Rio  Tram- 
pas,  1 1/2  miles  southeast  of  El  Valle,  elev. 
7,800',  20-IV-73,  1    9  . 

Capnia  gracilaria  Claassen. — Distribu- 
Tesuque  Creek, 


tion:    Santa  Fe  Co.,  Rig 

Study   supported   by   EPA   Grant    3053-3G4  and   NSF  Grant   G98G-600. 
■Department   of  Biology,   University  of  Utah,   Salt   Lake  City. 


97 


98 


GREAT    BASIN    NATURALIST 


Vol.  35,  No.  1 


Big  Tesuque  Campground,  elev.  9,700', 
21-IV-73,  3  d"  13  ?  ;  same  location,  17- 
III-74,  2  cf ;  Rio  Enmidio,  Hyde  Park  Ski 
Basin,  eler.  10,560',  lO-VI-74,  2  9  .  Taos 
Co.,  West  Fork  Red  River,  Wheeler  Peak 
Wilderness  Area,  9-VI-74,  1    cf    1    $  • 

Eucapnopsis  brevicauda  Claassen. — 
New  Records:  Santa  Fe  Co..  Big  Tesuque 
Creek,  Big  Tesuque  Campground,  elev. 
9,700',  21TV-73,  \   d- 

Paralcuctra  rickeri  Nebeker  &  Gaufin. 
— Distribution:  Taos  Co..  West  Fork  Red 
River,  Wheeler  Peak  Wilderness  Area,  10 
miles  south  of  Red  River,  9 -VI- 74,  12  d 
26   9. 

Paralcuctra  vershina  Gaufin  and  Rick- 
er. — This  species  has  been  previously  re- 
ported from  the  state  as  P.  sara  (Claas- 
sen). New  records:  Rio  Arriba  Co..  Rio 
Chama,  Chama,  elev.  7,870'  8-VI-74,  2 
cT  2  9  .  Taos  Co..  Rio  Trampas,  above  El 
Valle,  9-VI-74,  6  c?   H    ?• 

Pteronarcella  badia  (Hagen). — New 
Records:  Sandoval  Co..  Santa  Clara  Can- 
yon, Wem  Povi  Pond,  elev.  6,900',  20-III- 

73,  nymphs. 

Diura  knowltoni  ( Prison). — Distribu- 
tion: Rio  Arriba  Co..  Rio  Chama,  Chama, 
elev.  7,870',  8-VI-74,  2  d'  6  9  ;  same  loca- 
tion, 25-V-74,  M.  and  E.  Cather,  nymphs. 
Taos  Co..  Rio  Trampas,  IV2  miles  south- 
east of  El  Valle,  elev.  7,800',  20-IV-73, 
n}TTiphs. 

Isogenoides  elongatus  (Hagen). — New^ 
Records:  San  Miguel  Co..  Pecos  River, 
Pecos,  24-V-74,  M.  and  E.  Cather,  5c^  5 
9  .  _ 

Isogenoides  zionensis  Hanson. — Distri- 
bution: Catron  Co..  San  Francisco  River, 
Glen  wood,  19-11-70,  M.  Suavely,  nymph. 
McKinley  Co..  McGaffey  Lake,  lO-iV-70, 
n^niph.  Rio  Arriba  Co..  Rio  Chama, 
Chama,  elev.  7,870',  19-IV-73,  1  d 
(reared) ;  same  location,  8-VI-74,  6  cT  2 
9  .  Taos  Co..  Red  River,  Red  River,  5- 
III-61,  A.  R.  Gaufin,  nymph;  Red  River, 
3  miles  east  of  Cuesta, '9-VI-74,  nymphs. 

Kogotus  modestus  (Banks). — New  Rec- 
ords: Taos  Co.,  Red  River,  2  miles  east 
of  Red  River,  9-VI-74,  nymphs. 

Megarcys  si  gnat  a  (Hagen). — New  Rec- 
ords: Santa  Fe  Co..  Big  Tesuque  Creek, 
Big  Tesuque  Campground,  elev.  9,700', 
21-IV-73,  nymphs;  same  location,  10-VI- 

74,  46  cf  1  ?  ;  Rio  Enmidio,  Hyde  Park 
Ski  Basin,  elev.  10,560',  17-III-74, 
nymphs.  Taos  Co.,  Rio  Trampas,  11/2 
miles  southeast  of  El   Valle,  elev.   7,800', 


20-IV-73,  iiymphs;  West  Fork  Red  River. 
Wheeler  Peak  Wilderness  Area,  9-VI-74, 
nymphs. 

Skwala  parallel  a  ( Frison ) . — New  Rec- 
ords: 7V/O.S-  Co..  Rio  Hondo,  Hwy.  3,  5-III- 
61,  A.  R.  Ciaufin,  nymphs. 

Isoperla  ebria  (Hagen). — New  Records: 
Santa  Fe  Co.,  Big  Tesuque  Creek,  Big 
Tesuque  Campground,  elev.  9,700',  10- 
VI-74,  1  cf;  same  location,  21-IV-73, 
nymphs;  Rio  Enmidio,  Hyde  Park  Ski 
Basin,  elev.  10,560',  17-III-74,  nymphs. 

Isoperla  fulva  Claassen. — New  Records: 
Santa  Fe  Co..  Big  Tesuque  Creek,  Big 
Tesuque  Campground,  elev.  9,700',  21- 
IV- 73,  n}Tnphs. 

Isoperla  mormona  Banks. — New  Rec- 
ords: Catron  Co..  Whitewater  Creek,  Cat- 
walk Picnic  Area,  ll-VI-74,  1  d ;  West 
Fork  Gila  River,  17-V-74,  M.  and  E. 
Cather,  15  d'  6  9  .  Grant  Co.,  Gila  River, 
31  miles  north  of  Pinos  Altos,  Hwy  15, 
ll-VII-73,  1  (S  (reared);  same  location, 
17-V-74,  M.  and  E.  Cather,  26  d"  17   9  . 

Claassenia  sabulosa  (Banks). — New 
Records:  San  Miguel  Co..  Pecos  River, 
Tererro,  elev.   7,600',  21-IV-73,  nymphs. 

Hesperoperla  pacifica  (Banks). — New 
Records:  Catron  Co..  Whitewater  Creek, 
16-V-74,  M.  and  E.  Cather,  exuvium. 

Sweltsa  coloradensis  (Banks). — New 
Records:  Catron  Co.,  Willow  Creek,  Wil- 
low Creek  Campground,  ll-VI-74,  8  d 
7  9  .  Rio  Arriba  Co..  Rio  Chama,  Chama, 
elev.  7,870',  8-VI-74,  1  d'  3  9  .  Sandoval 
Co.,  Tschicoma  Pond.  Santa  Clara  Can- 
yon, 13-VI-74,  2    9. 

Sweltsa  lamba  (Needham  &  Claassen). 
— New  Records:  Taos  Co..  West  Fork  Red 
River,  Wheeler  Peak  Wilderness  Area, 
10  miles  south  of  Red  River,  9-VI-74,  1  d- 

Triznaka  diver sa  (Frison). — New  Rec- 
ords: Santa  Fe  Co..  Big  Tesuque  Creek, 
Big  Tesuque  Cami)ground,  elev.  9,700', 
10- VI- 74,  31    d^  24    9  . 

Triznaka  pintada  (Bicker). — New  Rec- 
ords: Taos  Co..  Rio  Pueblo,  Penasco,  9-VI- 
74,  2  d'   1  3    9  . 

Triznaka  signata  (Banks).— New  Rec- 
ords: Rio  Arriba  Co..  Rio  Chama,  Chama, 
elev.  7.870',  8-VI-74,  17   d   10   9  . 

Ni.w  Mi:xic:o  List 

Below  is  a  checklist  of  46  stonefly  spe- 
cies that  lune  been  confirmed  for  New 
Mexico.    Sjiecies  regarded  as  questionable 


March  1975 


STARK,   ET   AL.:    NEW   MEXICO   PLECOPTERA 


99 


or  unconfirmed  by  actual  specimens  and 
distributional  data  have  been  omitted. 

Taeniopterygidae 
Taenionema  nigripennis  (Banks) 
T.  pallida  (Banks) 
Taenioptcryx  sp. 

Nemouridae 
Amphinemura  mogollonica  Baumann  and  Gaufin 
Malenka  coloradensis  (Banks) 
M.  flexura  (Claassen) 
Podmosta  delicatiila  (Claassen) 
Prostoia  besametsa  (Ricker) 
Zapada  cinctipes  (Banks) 
Z.  frigida  (Claassen) 
Z.  haysi  (Ricker) 

Capniidae 
Capnia  confusa  Claassen 
C.  fibula  Claassen 
C.  gracilaria  Claassen 
Eucapnopsis  brevicauda  Claassen 
Mesocapnia  frisoni  (Baumann  and  Gaufin) 

Leuctridae 
Paraleuctra  rickeri  Nebeker  and  Gaufin 
P.  vershina  Gaufin  and  Ricker 
Perlomyia  utahensis  Needham  and  Claassen 

Pteronarcidae 
Pteronarcella  badia  (Hagen) 
Pterormrcys  califnrnica  Newport 

Perlodidae 
Cultus  aestivalis  (Needham  and  Claassen) 
Diura  knoivtoni  (Prison) 
Isogenoides  elongatus  (Hagen) 


/.  zionensis  Hanson 
Kogotus  modestus  (Banks) 
Megarcys  signata  (Hagen) 
Skwala  parallela   (Prison) 
Isoperla  ebria  (Hagen) 


.  fulva  Claassen 
/.  longiseta  Banks 
/.  mormona  Banks 
/.  patricia  Prison 
/.  phalerata  (Smith) 
/   quinquepunctata  (Banks) 

Perlidae 
Claassenia  sabulosa  (Banks) 
Hesperoperla  pacifica  (Banks) 
Neoperla  clymene  (Newman) 

Chloroperlidae 
Paraperla  frontalis  Banks 
Suwallia  pallidula  (Banks) 
Sweltsa  borealis  (Banks) 
S.  coloradensis  (Banks) 
S.  lamba  (Needham  and  Claassen) 
Triznaka  diver sa  (Prison) 
T.  pintada  (Ricker) 
T.  signata  (Banks) 


Literature    Cited 


Stewart,  K.  W..  R.  W.  Baumann,  and  B.  P. 
Stark.  1974.  The  distribution  and  past 
dispersal  of  southwestern  United  States  Ple- 
coptera.  Trans.  Amer.  Entomol.  Soc.  99:507- 
546. 


THE  AUTHORSHIP  AND  DATE  OF  PUBLICATION  OF 
SIREN  INTERMEDIA  (AMPHIBIA:  CAUDATA) 

Hobart  M.  Smith^,  Rozella  B.  Smith-,  and  H.  Lewis  Sawin^ 


Abstract. —  However  "just"  it  might  be  to  credit  LeConte,  1828.  with  Siren  intermedia,  both 
Harlan,  1826  (not  1827  as  often  cited),  and  Barnes,  1826,  antedate  LeConte's  proposal  of  the  name. 
As  the  earliest,  Barnes,  1826,  stands  credited  with  it.  In  analysis  of  precedent  for  these  conclusions, 
types  of  taxonomic  plagiarism  (calculated  vs.  innocent,  homoplagiarism  vs.  heteroplagiarism)  and  the 
distinctions  between  nomina  nuda  and  nomina  dubia  are  reviewed,  giving  examples  of  each  category. 

does  not  nullify  applicability  of  Art.  50  of 
the  International  Code  of  Zoological  No- 
menclature (ICZN,  1964:  49),  which 
states,  "The  author  ...  of  a  scientific 
name  is  .  .  .  the  person  .  .  .  who  first  pub- 
lishes it  in  a  way  that  satisfies  the  criteria 
of  availability,  unless  it  is  clear  from  the 
contents  of  the  publication  that  .  .  .  some 
other  person  is  alone  responsible  for  both 
the  name  and  the  conditions  that  make  it 
available"   (italics  ours). 

LeConte  obviously  was  responsible  for 
the  name  but  equally  clearly  was  not  re- 
sponsible for  the  "description"  that  "satis- 
fies the  criteria  of  availability."  Harlan 
obviously  wrote  the  description;  and  de- 
spite his  aj)parent  wish  to  the  contrary, 
the  present  rules  would  require  that  he 
be  regarded  as  author  of  the  name  in  zoo- 
logical nomenclature  if  indeed  his  ac- 
count were  the  earliest  to  have  appeared. 

There  is  ample  precedent  for  crediting 
the  immediate  source  of  an}"  given  name 
and  its  characterization,  however  ques- 
tionable may  be  the  derivation  of  either, 
for  that  name.  This  policy  unfortunately 
rewards  plagiarism  with  permanence  un- 
less the  International  Commission  on  Zo- 
ological Nomenclature  intercedes.  On  the 
other  hand,  plagiarism  seldom  occurs, 
either  inadvertently  or  deliberately.  Nev- 
ertheless, it  does  occur  on  occasion,  and 
the  Code  requires  that  the  perpetrator 
bear  responsibility  for  his  act,  whether  it 
be  innocent  or  calculated.  Examples  of  cal- 
culated plagiarism  are  provided  by 
Thom]:)son's  three  privately  printed  not- 
ices of  1912;  the  first  two  antedated  Van 
Denburgh's  competitive  advance  diagnosis 
of  1912,  and  although  Thompson's  de- 
scri])tions  are  sourced  directly  from  Van 
Denburgh's  manuscript,  insofar  as  they 
antedate     Van     Denburgh's     descriptions 


Martof  (1973:  1-3),  in  the  most  recent 
review  of  Siren  intermedia,  notes  that  the 
earliest  full  description  of  the  species  in 
LeConte  (1828:  133-134,  pi.  1)  actually 
was  antedated  by  a  brief  but  nominally 
occupying  characterization,  credited  to  Le- 
Conte, that  appeared  in  a  work  b}"  Har- 
lan (1826:  322),  dated  1827  by  Schmidt 
(1953:  14)  and  others. 

Two  points  merit  observation  in  this 
context:  (1)  the  particular  page  on 
which  the  "description"  of  Siren  inter- 
media appeared  in  Harlan's  work  was  ac- 
tually published  in  1826,  fide  the  1913  In- 
dex to  the  Scientific  Contents  of  the 
Journal  and  Proceedings  of  the  Academy 
of  Natural  Sciences  of  Philadelphia,  p.  viii; 
and  (2)  there  is  reason  to  accept  the 
author  of  this  "description"  as  Harlan, 
not  LeConte.  Harlan  "read"  his  paper  at 
the  12  Dec.  1826  meeting  of  the  Academy, 
and  accordingly-  the  pages  published  in 
1826  (pp.  317-324)  must  have  appeared 
sometime  after  12  Dec,  the  remainder 
(pp.  325-372)  in  February  1827,  accord- 
ing to  the  Index.  The  article  was  com- 
pleted in  no.  1  of  vol.  6  of  the  Journal 
(pp.  7-38)  appearing  in  March  1827  fide 
the  same  source. 

The  author  of  the  description  appearing 
in  Harlan  (1826:  322)  is  clearly  Harlan, 
not  LeConte,  despite  the  fact  that  Harlan 
attributed  the  name  to  LeConte  and  stat- 
ed (in  a  footnote)  that  the  material  on 
this  species  was  sourced  from  "manuscript 
notes."  The  characterization  obviously 
was  written  by  Harlan,  not  LeConte,  as 
becomes  evident  when  one  consults  Le- 
Conte's formal  description  that  appeared 
in  1828.  Harlan  seemingly  saw  the  Le- 
Conte ms.  and  published  in  his  own  words 
the  name  and  certain  characters  cited  in 
the  ms.    The  acknowledgment  of  source 

'Department    of    Environmental.    Population,    and    Organist 


Biology     and     Museum,     University     of     Colorado,     Boulder 


^Department   of  EPO    Biology   and   Center   for   Computer  Research    in    the    Iluma 
'Center   for   Computer   Research   in   the   Humanities,    University  of  Colorado. 


lies.    University    of   Colorado. 


100 


March  1975 


SMITH,   ET  AL.:    AMPHIBIAN   NOMENCLATURE 


101 


they  are  accepted  under  the  Code  as  valid 
(see  Barbour,  1917,  for  details). 

Examples  of  innocent  taxonomic  plagi- 
arism fall  into  two  categories:  self -plagi- 
arism (or  homoplagiarism)  and  hetero- 
plagiarism.  A  medium  for  frequent  homo- 
plagiarism  is  Dissertation  Abstracts, 
wherein  summaries  of  doctoral  disserta- 
tions occasionally  include  sufficient  in- 
formation with  a  new  name  or  a  new  com- 
bination to  occupy  them;  for  example 
Walker's  abstract  (1967)  includes  suffi- 
cient information  on  two  new  names 
{Cnernidophorus  gularis  rciuni,  C.  g.  semi- 
annulatus)  to  occupy  both,  whereas  it 
was  intended  that  these  names  not  be  en- 
tered into  nomenclature  before  full  docu- 
mentation could  be  provided  (full  descrip- 
tions have  not  even  yet  appeared).  A 
similar  case  occurred  in  another  journal 
(Harris,  1974),  wherein  a  photograph  and 
brief  notice  of  some  characteristics  of  a 
new  subspecies  of  rattlesnakes  was  ac- 
companied by  a  name  {Crotalus  ivillardi 
obscurus),  thus  occupying  the  name  in 
advance  of  the  intended  date  and  work 
which  was  then  in  press. 

An  example  of  heteroplagiarism  oc- 
curred with  inadvertent  mention  of  Palm- 
atotriton  by  Smith  (1945),  who  used  the 
name  under  the  impression  that  his  for- 
mer professor,  E.  H.  Taylor,  had  a  ms.  in 
press  establishing  the  genus,  and  that  the 
casual  mention  in  the  popular  journal 
would  be  meaningless.  Unfortunately 
Taylor  had  decided  against  erection  of 
the  genus,  and,  more  regrettably.  Smith's 
use  of  the  name  was  accompanied  by  a 
few  incidental  comments  inadvertently 
serving  to  occupy  the  name  nomenclatur- 
ally.  It  was  necessary  to  appeal  to  the 
International  Commission  on  Zoological 
Nomenclature  to  "deoccupy"  Palmatotri- 
ton  as  of  Smith,  1945,  making  the  name 
available  for  use  by  anyone  else,  in  any 
desired  sense  (ICZN,  1956). 

In  all  these  examples,  including  that 
of  Harlan,  it  is  clear  that  intent  has  noth- 
ing to  do  with  result;  only  the  briefest 
characterization,  in  but  a  few  words,  may 
serve  to  occupy  a  name  even  when  not  so 
intended,  and  the  person  responsible  is 
the  one  presenting  those  words,  even 
though  he  may  not  have  intended  to  re- 
ceive   that    responsibility. 

In  this  context  it  is  important  to  recog- 
nize that  a  name  may  be  occupied  even 


though  its  characterization  may  be  inade- 
quate for  definitive  allocation  to  its  prop- 
er taxon  in  nature;  such  names  are  nom- 
ina  diibia  despite  the  fact  that  they  are 
occupied  names.  There  is  a  rather  wide 
misapprehension  that  a  full  characteriza- 
tion is  required  in  order  to  occupy  a  new 
name,  but  this  is  not  so.  Nomina  dubia 
are  often  rendered  identifiable  (i.e.,  nom- 
ina  clara)  by  subsequent  provision  of 
further  details,  as  is  true  in  the  case  of 
Siren  intermedia.  Harlan's  description, 
although  adequate  to  occupy  the  name 
were  it  the  original  usage,  would  not  alone 
have  sufficed  for  allocation;  but  with  Le- 
Conte's  full  account,  no  doubt  remains. 
As  of  Harlan,  Siren  intermedia  is  a  no- 
men  dubiiim;  as  of  LeConte,  it  became  a 
nomen  clarum  although  occupied  at  an 
earlier  date  by  another  author.  Harlan's 
usage  was  not  of  a  nomen  nudum,  which 
is  nonexistent  nomenclaturally,  because  it 
did  provide  some  distinguishing  informa- 
tion. The  Code  makes  clear  (Art.  13)  that 
any  "statement  that  ]:)urports  to  give  char- 
acters differentiating  the  taxon"  (italics 
ours)  suffices  to  occupy  an  accompanying 
name,  and  practice  has  conformed  with 
this  liberal  rule. 

In  the  case  of  Siren  intermedia,  how- 
ever, the  comedy  of  errors  did  not  really 
begin  with  Harlan,  even  of  1826.  There 
is  a  still  earlier  usage  that  occupied  the 
name.  Barnes  (1826:  269,  footnote)  saw 
or  otherwise  knew  of  LeConte's  ms  and 
rendered  the  name  Sir-eii  intermedia  avail- 
able in  almost  precisely  the  same  way  that 
Harlan's  work  would  have  done  had  it 
been  the  earliest  usage.  The  Barnes  foot- 
note follows:  ''Additional  note  communi- 
cated by  the  author,  Aug.  15,  1826.  The 
delay  in  the  printing  of  this  paper  has 
given  the  author  an  opportunity  of  an- 
nouncing, in  this  place,  the  discoverv  of 
ANOITTER  NEW  SPECIES  OF  SIREN, 
by  Capt.  LECONTE.  It  belongs  to  this 
section,  and  is  called  by  its  discoverer  Siren 
intermedia.  In  its  color  it  resembles  the 
Lacertina,  and  in  its  gills,  the  Striata; 
but  it  has  peculiar  characters  of  its  own, 
which  will  be  explained  at  length  in  a 
paper  soon  to  be  published  in  the  Annals 
of  the  Lyceum.  Length  about  one  foot, 
inhabits  the  Southern  states  in  large  num- 
bers. Specimens  are  preserved  in  the 
Cabinet  of  the  Lyceum.  Fig.  Annals  of 
the  Lyceum,  Vol.  2,  fig.  1."  That  Barnes 
knew   of   LeConte's   description   long  be- 


102 


GREAT   BASIN    NATURALIST 


Vol.  35,  No.  1 


fore  its  publication  is  not  surprising,  in- 
asmuch as  he  was  the  "Recording  Secre- 
tary of  the  New  York  L}xeum,"  as  indi- 
cated (p.  268)  in  his  1826  paper.  Only 
by  the  close  familiarity  permitted  by  such 
an  association  could  he  have  known  some 
two  years  in  advance  of  publication  that 
LeConte's  account  would  appear  in  Vol- 
ume 2  and  incorporate  Figure  1  of  the 
Annals  of  the  Lyceuin! 

The  Harlan  and  Barnes  works  were  both 
dated  1 826,  but  the  Harlan  paper  appeared 
very  late  in  the  year — certainly  after 
December  12 — whereas  the  Barnes  paper, 
read  before  the  Lyceum  in  July  1825,  was 
surely  published  shortly  after  15  August 
1826^  when  Barnes  inserted  his  footnote 
on  S.  intermedia.  We  have  not  been  able 
to  pinpoint  the  exact  date  of  publication 
of  either  work,  but  the  evidence  that 
Barnes'  work  preceded  that  of  Harlan  is 
overwhelming. 

The  same  generalities  })ertinent  to  Har- 
lan's use  of  the  name  SireJi  intermedia 
are  equally  pertinent  to  the  earlier  Barnes 
usage.  Barnes  actually  must  be  regarded 
as  the  author  of  Siren  intermedia  (which 
accordingly  dates  from  1826),  unless  the 
case  is  appealed  to  the  ICZN,  asking  for 
rejection  of  the  contributions  of  both 
Barnes  and  Harlan  on  that  species,  giving 
LeConte  (1828)  priority.  The  effort  is  not 
warranted,  however,  since  the  significance 
of  the  case  is  grossly  inadequate  to  justify 
the  protracted,  laborious  protocol  involved 
in  ICZN  action.  Custom  dictates  that  sus- 
pension of  the  rules  be  requested  only  for 
names  of  relatively  broad  familiarity 
among  zoologists;  the  present  certainl}- 
does  not  fall  into  that  category. 

We  are  accordingly  left  with  the  con- 
clusion that  the  proper  citation  for  the 
species  under  consideration  is  Siren  inter- 
media Barnes,   1826. 


Literature  Cited 

B.^RBOUR,  T.  1917.  A  most  regrettable  tangle  of 
names.  Occ.  Pap.  Mus.  Zool.  Univ.  Michigan 
44:1-9. 
Barnes,  D.  H.  1826.  An  arrangement  of  the 
genera  of  batracian  [sic]  animals,  with  a  de- 
scription of  the  more  remarkable  species;  in- 
cluding a  monograph  of  the  doubtful  reptils 
[sic].  Am.  J.  Sci.  11:268-297. 
Harl.^n,  R.  1826-27.  Genera  of  North  American 
Reptilia  and  a  synopsis  of  the  species.  J. 
Acad.  Nat.  Sci.  Philadelphia  5:317-372;  ibid., 
6:7-38  (pp.  317-324.  Dec,  1826;  325-372,  Feb., 
1827;  7-38,  Mar.,  1827). 
Harris,  H.  S.,  Jr.  1974.  The  New  Mexican 
ridge-nosed  rattlesnake.  Nat.  Parks  Cons. 
Mag.  48(3): 22-24,  3  figs. 
International  Commission  on  Zoological  Nomen- 
clature. 1956.  Opinion  425:  Addition  to  the 
"Official  index  of  rejected  and  invalid  generic 
names  in  zoology"  of  the  name  ''Pabnatotri- 
ton'  Smith  (H.  M.).  1945  (Class  Amphibia). 
Opin.  Decl.  Int.  Comm.  Zool.  Nom.  14:243- 
256. 

.      1964.     International    code    of    zoological 

nomenclature  adopted  by  the  fifteenth  inter- 
national congress  of  zoology.  London,  Intern. 
Trust  Zool.  Nomencl.  xviii,  176  pp. 
LeConte,  J.  E.  1828.  Description  of  a  new 
species  of  Siren.  Ann.  Lyceum  Nat.  Hist. 
New  York  2:133-134,  pi.  1. 
Martof,  B.  S.     1973.     Siren  intermedia.   Cat.  Am. 

Amph.    Kept.    127:1-3.   map. 
Schmidt,    K.    P.     1953.     A    check    list   of    North 
American   amphibians   and   reptiles.    Chicago, 
Am.  Soc.  Ichth.  Herp.  vii.  280  pp. 
Smith,    H.    M.     1945.     Herpetological    collecting 
in  banana  fields  of  Mexico.    Ward's  Nat.  Sci. 
Bull.   19(l):3-7.  figs.  1-6. 
Thompson.  J.   C.     1912.     Prodrome  of  a  descrip- 
tion of  a  new  genus  of  Ranidae  from  the  Loo 
Choo    Islands.     Herpetological    Notices    1:1-3, 
1   pi. 

.      1912.     Prodrome   of  descriptions   of  new 

species    of    Reptilia    and    Batrachia    from    the 
Far    East.     Herpetological    Notices   2:1-4. 

.      1912.     On  reptiles  new  to  the  island  arcs 

of  Asia.  Herpetological  Notices  3:[i-ii],  1-5. 
Van  Denrurgh.  J.  1912.  Advance  diagnoses  of 
new  reptiles  and  amphibians  from  the  Loo 
Choo  Islands  and  Formosa.  Privately  Printed. 
5  pp. 
Walker.  J.  M.  1967.  Morphological  variation 
in  the  teid  lizard  Cnemidophorus  gularis. 
Diss.  Abst.  Int.,  B  28:1738-1739. 


NEW  MITES  FROM  THE  YAMPA  VALLEY^ 
(ACARINA:  CRYPTOSTIGMATA:  ORIBATULIDAE,  PASSALOZETIDAE) 

Harold  G.  Higgins-  and  Tyler  A.  Woolley^ 

Abstract. —  A  study  was  made  of  the  soil  mites  from  under  different  vegetative  types  near  a 
coal-burning  power  plant  in  the  Yampa  Valley  near  Hayden,  Colorado.  The  following  new  species  of 
oribatids  were  found:  Zygoribatula  apletosa  n.sp.,  Multoribates  haydeni  n.sp.,  Paraphauloppia  cordylin- 
osa  n.sp.,  Passalozetes  moniles  n.sp. 


Concentrated  collections  have  been 
made  near  a  coal-burning  power  plant  in 
the  Yampa  Valley  near  Hayden,  Colora- 
do. A  number  of  new  and  unrecorded 
species  of  soil  mites  for  Colorado  have 
been  found  in  the  project  area.  The  col- 
lections were  made  with  reference  to  the 
soil  forms  and  to  vegetative  types  with 
which  they  were  found.  As  might  be  ex- 
pected, many  species  appear  to  be  more 
abundant  at  one  season  of  the  year  than 
at  another  or  may  be  more  closely  as- 
sociated with  certain  vegetative  types  than 
with  others. 

This  concentrated  collecting  over  a  two- 
year  period  has  given  new  insight  as  to  the 
importance  of  the  microclimate  in  the 
biology  of  oribatids.  For  example,  depend- 
ing on  the  amount  of  moisture,  sunlight 
or  shade,  and  slope,  many  species  may  be 
more  abundant  under  one  side  of  a  bush 
than  under  the  other.  Preliminar}^  studies 
also  indicate  that  destruction  of  vegetation 
and  distiu-bances  of  the  soil  in  such 
changes  as  the  formation  of  spoil  banks 
or  strip-mining  and  pollution  from  coal- 
burning  power  plants  seriously  depletes 
the  numbers  and  kinds  of  soil  arthropods. 

Following  are  descriptions  of  four  new 
species  representing  two  families  of  orib- 
atids found  in  the  Hayden  area. 

Family  Oribatulidae 

Zygoribatula  apletosa,  n.sp. 

Figs.   1   and  2 

Diagnosis. —  Large  size,  larger  than 
any  known  Zygoribatula;  with  14  pairs  of 
large,  setose  notogastral  setae;  rostral  hairs 
further  apart  than  lamellar  hairs;  lamel- 
lae curved  inward  with  distinct  translam- 
ella;  areae  porosae  Aa  located  near  the 
small  shoulder  projections.  The  trivial 
name  apletosa  is  modified  from  the  Greek, 


and  implies  "immense,"  referring  to  the 
size  of  these  oribatids. 

Description. —  Color  reddish-brown; 
rostrimi  rounded;  rostral  hairs  heavy, 
reaching  beyond  tip  of  rostrum  by  about 
half  their  lengths,  hairs  farther  apart 
than  lamellar  setae;  lamellae  of  almost 
uniform  width  throughout,  length  curved 
inward  toward  anterior  tip,  slightly  less 
than  one-half  as  far  apart  at  tip  as  at 
base;  translamella  narrower  than  lamel- 
lae; lamellar  hairs  similar  to  rostral  hairs 
but  about  one-third  longer  inserted  in 
anterolateral  ends  of  lamellae;  interlamel- 
lar  hairs  situated  midway  between  inser- 
tions of  lamellar  hairs  and  pseudostig- 
mata,  closer  to  inner  margin  of  lamellae; 
pseudostigmata  cuplike  with  edge  erected 
above  surface  of  prodorsiun;  sensillum 
with  broad,  rounded  setose  head  and  short 
pedicel,  about  half  as  long  as  interlamel- 
lar  hair;  exobothridial  hair  rather  heavy 
and  stiff. 

Hysterosoma  longer  than  broad,  widest 
near  middle,  with  tapering  posterior  end; 
dorsal  surface  with  14  pairs  of  heavy, 
long,  spined  setae,  many  extending  beyond 
body  outline  as  shown  in  Figure  1;  areae 
porosae  all  large,  Aa  much  longer  than 
broad  and  located  near  small  humeral 
process. 

Camerostome  oval  in  outline;  ventral 
surface  with  apodemata  and  setae  as 
shown  in  Figure  2;  genital  and  anal  aper- 
tures far  apart,  smaller  genital  opening 
more  than  twice  its  length  anterior  to 
larger  anal  aperture;  each  genital  cover 
with  four  setae;  aggenital  setae  as  shown 
in  Figure  2;  each  anal  cover  with  two 
setae  (2  of  12  specimens  with  3  anal  se- 
tae) ;  two  adanal  setae  present. 

Legs  all  about  equal  in  size;  heterotri- 
dactylous,  median  claw  larger  than  later- 
als. 


^Yampa    Valley    Project,    Ecologj-    Consultants,    Inc. 

-Biology-    Department,    Granger    High    School,    Granger,    Utah. 

^Department   of  Zoologj-  and  Entomology,   Colorado  State  University,  Fort  Collins,  Colo.  80523. 


103 


104 


GREAT  BASIN  NATURALIST 


Vol.  35,  No.  1 


Fig.  1.  Zygoribatula  apletosa,  dorsal  aspect, 
legs  omitted. 

Measurements. —  length  .705  mm; 
width,  .495  mm.  (Range  .853-.705  mm  X 
.600-. 495  mm).  The  type  (a  male)  and  6 
paratypes  (6  females)  were  taken  under 
serviceberry  about  1/4  mile  NE  power 
plant,  Hayden,  Colorado,  7  Oct.  1971,  by 
H.  G.  Higgins;  2  specimens,  females,  were 
taken  under  rosebush,  I/2  mile  N  power 
plant,  Hayden,  Colorado,  21  June  1972; 
2  specimens  (males)  were  taken  i^  mile 
N  power  plant,  Hayden,  Colorado,  1  Aug. 
1971;  1  specimen  (male)  was  taken  under 
aspens,  4  miles  S  Seneca  Road,  Hayden, 
Colorado,  1  Aug.  1971;  all  by  H.  G."  Hig- 
gins. 

Discussion. —  This  species  stands  a- 
part  from  other  known  North  American 
Zygoribatula  by  its  large  size,  its  long, 
heavy,  setose  body  setae,  and  by  its  big, 
long,  areae  porosae  Aa  located  near  the 
shoulder.  In  general  appearance  Z.  aple- 
tosa n.sp.  resembles  Z.  lata  Hammer  but 
differs  in  the  much  larger  size  as  well  as 
in  the  size  and  locations  of  areae  porosae. 


Fig.  2.     Z.  apletosa,  ventral  aspect. 

To  date,  this  species  has  always  been 
found  associated  with  rather  heavy,  moist 
litter  under  dense  vegetation. 

This  species  shows  interesting  variation 
in  the  width  of  the  translamellae  and  lo- 
cation of  body  setae.  Also,  as  pointed  out 
earlier,  2  of  12  specimens  have  3  pairs, 
rather  than  2  pairs,  of  anal  setae. 

Multoribates  haydeni,  n.sp. 
Fig.    3 

Diagnosis. —  Similar  to  Multoribates 
chavinensis  Hammer,  1961,  but  larger, 
and  with  only  11  pairs  of  dorsal  setae; 
lacks  the  ventral  keel  on  femur  II.  The 
trivial  name  is  indicative  of  location. 

Description. —  Large  size;  color  yel- 
lowish to  light  brown;  body  egg-shaped 
with  the  pteromorphs  hardly  projecting 
beyond  lateral  outline  of  body;  prodorsum 
triangular  in  outline  with  rostrum  fairly 
pointed,  often  hyaline;  rostral  setae  in- 
serted posteriorly  on  lateral  margins  of 
propodosoma,  much  wider  apart  than  la- 
mellar hairs;  lamellae  narrow,  tapering 
slightly  anteriorly;  translamellae  absent; 
lamellar  hairs  stiff,  setose  about  same 
length    as    lamellae;    interlamellar    hairs 


March  1975 


HIGGINS,    WOOLLEY:    COLORADO    MITES 


105 


/f  ^^ 


Fig.  3.  Multoribates  haydeni,  dorsal  aspect, 
legs  omitted. 

heavy,  nearly  equal  in  length  to  lamellar 
hairs;  pseudostigmata  cuplike,  rim  project- 
ing beyond  the  body  level;  sensillum  with 
narrow  stalk  and  broad,  setose  head  that 
is  bent  backwards;  dorsosejugal  suture 
curved  anteriorly. 

Hysterosoma  longer  than  broad  with 
small  pteromorphs  that  project  only 
slightly  beyond  the  outline  of  body;  11 
pairs  of  fine,  simple  dorsal  hairs  visible  as 
shown  in  Figure  3;  areae  porosae  absent, 
but  replaced  with  chitinous  pores;  muscle 
scars  and  markings  visible  round  edge  of 
hysterosoma  as  indicated  in  Figure  3.  Var- 
iations occur  in  the  locations  of  body  se- 
tae. 

Ventral  surface  similar  to  M.  chavinen- 
sis  with  only  a  few  minor  exceptions;  gen- 
ital plates  separated  from  larger  anal 
plates  by  approximately  twice  their 
length,  each  plate  with  four  setae;  ana] 
plates  much  larger  than  genital  plates, 
situated  near  posterior  end  of  body,  each 
anal  plate  with  two  hairs;  adanal  and 
aggenital  setae  place  similarly  to  M.  chav- 
inensis;  fissure  iad  located  near  antero- 
mediad  margin  of  anal  plates. 

Legs  about  equal  in  size;  all  legs  hetero- 
tridactylous,  median  claw  larger  than  lat- 
erals; femur  II  without  visible  keel. 

Measurements. —  Length,  .45  mm; 
width,  .26  mm.    The  type,  a  gravid  fe- 


male was  collected  at  Seneca  #2,  Hayden, 
Colorado,  under  serviceberry,  10  April 
1971,  by  H.  G.  Higgins  and  T.  A.  Wool- 
ley.    Additional  specimens  are  as  follows: 

I  specimen  at  Seneca  :^2,  Hayden,  Colo- 
rado, 9  June  1971,  in  sagebrush,  by  T.  A. 
Woolley  and  H.  G.  Higgins;  3  specimens 
from  under  aspens  associated  with  bitter- 
brush  (Purshia),  8  June  1971,  4  miles  S 
Seneca  Road,  Hayden,  Colorado,  by  H.  G. 
Higgins  and  T.  A.  Woolley;  6  specimens 
from  under  bitterbrush,  5  miles  S  Seneca 
Road,  Hayden,  Colorado,  1  Aug.  1971,  by 
H.  G.  Higgins;  8  specimens  from  under 
bitterbrush,  5  miles  S  Seneca  Road,  Hay- 
den, Colorado,  8  Oct.  1971,  by  H.  G.  Hig- 
gins; 1  specimen  from  under  bitterbrush, 
4  miles  S  Seneca  Road,  Hayden,  Colorado, 
21  June  1972,  by  H.  G.  Higgins. 

Discussion. —  In  general  appearance 
M.  haydeni,  n.sp.  resembles  M.  chavinen- 
sis  Hammer  but  is  larger,  lacks  the  ventral 
keel  on  femur  II,  and  has  11  rather  than 
14  pairs  of  dorsal  setae.  Preliminary  study 
seems  to  indicate  that  although  this  new 
species  is  found  in  several  habitats,  it  pre- 
fers the  microhabitat  under  bitterbrush  in 
rather  arid  conditions. 

Paraphauloppia  cordylinosa,  n.sp. 

Fig.    4 

Diagnosis. — Similar  in  outline  to  Para- 
phauloppia novazealandica  Hammer, 
1967,  but  with  much  larger  lamellae  and 

II  pairs  of  notogastral  hairs.  The  name 
cordylinosa  refers  to  the  clublike  sensillum 
of  the  new  species. 

Description. —  Color  yellowish;  ros- 
tral setae  large,  rough,  situated  on  the  an- 
terolateral margins  of  the  propodosoma; 
lamellae  quite  large,  extended  more  than 
half  the  length  of  propodosoma,  about 
equal  in  width  throughout  theirs  lengths, 
with  a  small  spur  (prolamella)  located 
anteromediad;  lamellar  hairs  inserted  in 
anterior  tip  of  lamellae,  extending  to  tip 
of  rostrum,  heay>',  barbed,  and  about 
equal  in  length  to  rostral  hairs;  interla- 
mellar  hairs  three-fourths  as  long  as  la- 
mellar hairs,  inserted  mediad,  closer  to 
pseudostigmata  than  to  tip  of  lamellae  ex- 
tending to  the  sides  of  prodorsum;  pseudo- 
stigmata cuplike  with  a  short  stalk  and 
expanded,  rounded,  setose  head. 

Hysterosoma  oval,  tapering  anteriorly; 
dorsosejugal  suture  greatly  arched;  ptero- 


106 


GREAT  BASIN  NATURALIST 


Vol.  35,  No.  1 


Fig.  4.  Paraphauloppia  cordylinosa,  dorsal  as- 
pect, legs  omitted. 

morphs  absent;  11  pairs  of  simple  smooth, 
dorsal  hairs  as  shown  in  Figure  4.  Areae 
porosae  visible  but  not  noticeably  en- 
larged as  in  P.  novazealandica;  a  line  of 
light  marking,  probably  muscle  scars,  vis- 
ible mediad  of  lateral  margins. 

Anal  opening  much  larger  than  geni- 
tal opening  and  situated  near  posterior 
end  of  body,  each  cover  with  two  setae; 
genital  opening  smaller,  situated  about 
twice  its  length  in  front  of  anal  opening, 
three  pairs  of  visible  hairs  on  each  plate; 
aggenital  setae  posterior  to  genital  plate; 
aggenital  and  adanalsetae  located  similarly 
to  P.  novazealandica,  adi  ])osterior  to  anal 
plate,  ado  situated  near  the  middle  of  plate 
along  lateral  side,  and  ads  situated  anteri- 
or to  anal  plate. 

All  legs  about  equal  in  size,  hetero- 
tridactylous,  with  median  claw  larger  than 
laterals. 

Measurements. —  Length,  282^1,;  width, 
132/i.  The  type  (a  female)  and  a  para- 
type  from  sagebrush,  Yampa  Airport, 
Hayden,  Colorado,  9  June  1971;  6  speci- 
mens from  sagebrush,  2  miles  S  Yampa 


Airport,  Hayden,  Colorado,  9  June  1971; 
all  by  H.  G.  Higgins  and  T.  A.  Woolley. 

Discussion. ^ —  In  general  appearance 
this  species  resembles  P.  novazealandica 
Hammer  but  differs  in  having  much  lar- 
ger, heavier  lamellae  and  11  rather  than 
10  pairs  of  dorsal  setae.  Although  this 
taxon  differs  somewhat  from  Paraphaulop- 
pia in  the  number  of  dorsal  setae,  it  ap- 
pears to  be  nearer  this  genus  than  to 
Phauloppia.  We  hesitate,  at  this  time, 
to  describe  a  new  genus  based  on  these 
minor  differences  and  because  in  the  small 
sample  of  mites  collected  there  is  great 
variation  in  the  exact  location  of  the  dor- 
sal setae,  and  the  lengths  of  the  lamellar 
hair.  It  is  interesting  that  although  col- 
lections were  made  at  the  same  general 
site  several  times  a  year,  and  over  a  two- 
year  span,  specimens  of  this  species  were 
found  only  once,  in  June  1971,  and  those 
in  rather  dry  sagebrush  habitat.  Prelimin- 
ary postulations  attribute  this  to  the  pos- 
sible influence  of  pollutants  in  the  area. 

Family  Passalozetidae 

Passcdozetes  moniles,  n.sp. 
Figs.    5-6 

Diagnosis. —  This  species  is  readily 
separated  from  P.  linearis,  the  only  other 
known  North  American  species,  by  the 
banded,  beadlike  pattern  of  dorsal  and 
ventral  integuminal  and  by  the  smooth 
sensillum  with  a  pointed  tapered  head. 
The  trivial  name  moniles  is  modified  from 
the  Latin  meaning  "necklace"  and  refers 
to  the  beaded  appearance  of  the  integu- 
ment. 

Description. — Yellowish  in  color;  pro- 
dorsimi  slightly  wider  than  long;  rostrum 
blunt,  rounded;  rostral  hairs  simple,  in- 
serted near  tip  of  rostrum,  curved  medi- 
ally toward  tip  of  rostrimi;  lamellae  ab- 
sent; lamellar  hairs  small,  simple,  slightly 
longer  than  rostral  hairs,  curved  down  to- 
ward tip  of  rostrum;  interlamellar  hairs 
threadlike,  simple,  inserted  anteromediad 
of  pseudostigmata,  adjacent  to  coalesced 
median  section  of  dorsosejugal  suture; 
l)seudostigmata  cuplike,  separated  from 
each  other  by  a  little  more  than  length  of 
sensillum;  sensillum  with  a  narrow  curved 
base  and  tapering  into  a  narrow  pointed 
head. 

Hysterosoma  oval,  anterior  margin  ex- 
tended forward,  coalesced  medially  with 


March  1975 


HIGGINS,    WOOLLEY:    COLORADO   MITES 


107 


^rv/,l;M^»\ 


Fig.  5.     Passalozetes    moniles,     dorsal     aspect, 
legs  omitted. 


ted. 


Fig.  6.     P.  moniles,  ventral  aspect,  legs  omit- 


dorsum  of  propodosoma  beyond  level  of 
interlamellar  hairs;  dorsosejugal  suture  in- 
terrupted by  this  middle  projection;  lenti- 
culus  clear,  round,  surrounded  by  lines; 
dorsum  with  simple  hairs  as  shown  in 
Figure  5.  Integument  of  fine  lines  with 
darker  pigmented  areas  resembling  strings 
of  beads  (the  beadlike  cerotegument  may 
be  removed  by  soaking  in  lactophenol) ; 
two  pairs  of  area  porosae  and  a  glandular 
fissure  as  seen  in  Figure  5. 

Camerostome  with  rather  parallel  sides, 
longer  than  wide;  ventral  plate  as  seen  in 
Figure  6;  genital  covers  each  with  four 
pairs  of  genital  setae;  aggenital  setae  in- 
serted about  twice  their  lengths  postero- 
laterad  of  genital  aperture;  anal  aper- 
ture nearly  one-third  larger  than  anal 
opening;  anal  aperture  in  the  posterior 
end  of  ventral  plate,  each  cover  with  two 
setae;  adanal  setae  difficult  to  find  in  the 
cerotegument,  visible  setae  and  glands  as 
shown  in  Figure  6. 

All  tarsi  heterobidactylous,  heavier  of 
the  two  claws  toothed  (median  or  lateral) . 

Measurements. —  Length,  .36  mm; 
width,  .15  mm. 

The  type  (a  male)  and  3  para  types  were 
taken  from  under  bitterbrush  and  squaw- 
brush,  6  miles  E  Craig,  Colorado,  14  June 


1972;  4  specimens  from  burned-over  area 
at  Seneca  #3,  Hayden,  Colorado,  21  June 
1972;  all  by  H.  G.  Higgins. 

Discussion. —  As  is  to  be  expected, 
there  is  considerable  variation  in  the  ar- 
rangement of  setae  and  pigmentation  in 
specimens  examined.  Those  examples 
from  the  burned-over  area  were  more 
heavily  pigmented  and  have  a  wider  hys- 
terosoma  than  those  taken  near  Craig. 
The  only  previously  described  Passalozetes 
from  this  western  area  was  taken  from  a 
dry  woodrat  nest  in  Tooele  Co.,  Utah, 
several  hundred  miles  to  the  west  of  the 
location  of  this  new  species.  Both  North 
American  species  of  Passalozetes  have 
been  taken  from  dry  desert  sands,  which 
implies  that  they  are  found  in  xeric  habi- 
tats. 


References 

AoKi,  J.  1961.  On  six  new  oribatid  mites  from 
Japan.    Jap.  J.  Sanitary  Zool.  12(4):  233-238. 

.     1964.     Some    oribatid    mites     (Acarina) 

from  Laysan  Island.  Pacific  Insects  6(4): 
649-664. 

Balogh,  J.  1965.  A  synopsis  of  the  world  ori- 
batid (Acari)  genera.  Acta  Zool.  11(1-2):  5- 
99. 

CoETZER,  A.     1967-68.     New  Oribatulidae  THOR, 


108 


GREAT  BASIN  NATURALIST 


Vol.  35,  No.  1 


1929    (Oribatei,    Acari)    from    South    Africa, 

new   combinations   and   a   key   to   the   genera 

of    the    family.     Mems.    Inst.    Invest.    Ciest. 

Mocamb.  9,  Serie  A:    13-126. 
EwiNG,  H.  E.     1913.     New  Acarina.    Bull.  Amer. 

Mus.  Nat.  Hist.  32(5):   93-121. 
.     1917.     New  Acarina.    Bull.  Amer.  Mus. 

Nat.    Hist.    37(2):    149-172. 
Hammer,  M.     1961.     Investigations  on  the  oriba- 

tid  fauna  of  the  Andes  Mountains.  II.  Peru. 

Biol.   Skr.  Dan.  Vid.   Selsk.   13(1):    1-150   + 

plates. 
.     1967.         Investigations    on    the    oribatid 

fauna    of   New    Zealand,    Part   II.    Biol.    Skr. 

Dan.  Vid.  Selsk.  15(4):    1-64  +  plates. 
Higgins,  H.  G.,  .\nd  T.  a.  Woolley.     1962.     A 

new   species  of  Passalozetes  from  Utah  with 


notes  on  the  genus.  (Acarina:  Oribatei). 
Great  Basin  Nat.  22(4):  93-100. 

Jacot,  a.  p.  1961.  Journal  of  North  American 
moss-mites.  J.  N.  Y.  Ent.  Soc.  45(3-4):  353- 
375. 

WiLLMANN,  G.  1931.  Moosmilben  oder  Oribati- 
den  (Oribatei).  In:  Tierwelt  Deutschlands 
22(5):  79-200. 

Woolley,  T.  A.  1957.  Redescription  of  Ewing's 
oribatid  mites.  Ill — Family  Eremaeidae 
(Acarina:  Oribatei).  Ent.  News  68(4):  147- 
156. 

.  1961.  Redescriptions  of  Ewing's  oriba- 
tid mites.  Ill — Family  Oribatulidae  (Acarina: 
Oribatei).  Trans.  Amer.  Microscop.  Soc. 
80(1):     1-15. 

Higgins,  H.  G.,  and  T.  A.  Woolley 


THE  IDENTITY  OF  BOCOURT'S  LIZARD  EUMECES 
CAPITO    1879 

Hobai't  M.  Smith\  Rozella  B.  Smith\  and  Jean  Guibe" 

Abstract. —  Eumeces  capito  Boucourt,  1879,  is  a  senior  synonym  of  Eumeces  xanthi  Giinther, 
1889.  No  exception  to  application  of  the  Law  of  Priority  is  recommended  in  this  case.  The  type 
locality  of  E.  capito  as  originally  published  ("La  cote  oriental  des  Etats-Unis")  is  erroneous.  Undoubt- 
edly the  correct  locality  is  China,  but  it  is  not  restricted  at  present. 


In  preparation  of  "Synopsis  of  the 
Herpetology  of  Mexico,"  a  problem  has 
arisen:  whether  or  not  to  apply  the  name 
Eumeces  capito  Bocom-t  (1879:429-431, 
pi.  22D,  Figs.  8,  8a-8c)  to  some  Mexican 
species.  The  origin  of  the  only  specimen 
(holotype,  no.  5531  of  the  Museum  Na- 
tional d'Histoire  Naturelle,  Paris)  is  in 
doubt,  for  although  the  published  locality 
is  "la  cote  oriental  des  Etats-Unis,"  the 
several  registers  in  the  Museum  in  Paris 
give  still  other  indications:  "Mexique" 
and  "Amerique  septentrionale."  The  col- 
lector is  also  unknown,  for  M.  S.  Bracon- 
nier,  by  whom  Bocourt  {loc.  cit.)  indicat- 
ed the  specimen  was  "donne,"  was  merely 
an  assistant  in  the  Museum,  never  par- 
ticipating in  any  expedition;  he  simply 
made  this  and  other  specimens  available 
for  study  by  Bocourt  and  other  scientific 
personnel. 

Taylor's  exhaustive  monograph  (1936: 
28,  231-2,  506)  of  Eumeces  sheds  no  light 
upon  the  problem  of  the  identity  of  E. 
capito,  as  the  type  was  not  examined,  and 
by  description  alone  the  name  could  not 
be  allocated.  Taylor  did  note  the  possi- 
bility that  the  name  applies  to  his  Eu- 
meces inexpectatus,  1932,  although  he 
thought  it  "probable  that  it  is  based  upon 
an  aberrant  specimen  of  jasciatus"  (p. 
232). 

We  have  thus  been  motivated  to  re- 
examine the  only  specimen  constituting 
the  hypodigm  for  Eum.eces  capito^  being 
made  available  by  the  junior  author  for 
study  by  all  of  us.  It  proves  to  be  a  typi- 
cal, mature  example  of  Eumeces  xanthi 
Giinther  (1889:218).  The  specimen  is  in 
excellent  condition,  slightly  softened,  but 
not  notably  damaged,  discolored,  or  faded. 
Salient  data  are:  snout- vent  72  mm,  hind 
leg  28  mm,  foreleg  19  mm,  axilla-groin 
40  mm,  tail  69  mm  (30  mm  regenerat- 
ed);  24  scale  rows  around  midbody;  54 


scales  from  parietals  to  above  anus;  16-16 
lamellae  under  4th  toe;  a  conspicuous 
patch  of  enlarged  postfemoral  scales;  one 
postnasal;  two  postmentals;  parietals  nar- 
rowly separated  posteriorly  by  tip  of  in- 
terparietal; frontonasal  broadly  contacting 
frontal;  upper  secondary  temporal  quad- 
rangular, dorsal  and  lateral  edges  nearly 
parallel,  separated  from  7th  (posterior) 
supralabial  by  contact  of  primary  tem- 
poral and  lower,  subtriangular  secondary 
temporal;  two  pairs  of  nuchals;  median 
preanals  overlapped  by  lateral  scales;  a 
somewhat  modified,  slightly  keeled  lateral 
postanal;  median  subcaudals  twice  as  wide 
as  adjacent  scales;  a  distinct,  brown  later- 
al stripe  on  4th  scale  row  and  edges  of 
adjacent  3rd  and  5th  rows;  a  dorsolateral 
light  stripe  occupying  most  of  the  3rd 
scale  row;  and  a  lateral  light  stripe  occu- 
P3'ing  the  lower  half  of  the  5th  scale  row; 
no  evidence  of  a  median  light  stripe  on 
head,  and  its  only  evidence  on  trunk  the 
absence  of  dark  pigment  on  the  adjacent 
halves  of  the  2  median  scale  rows;  other 
dorsal  scale  rows  with  some  dark  flecking 
on  the  base  of  each  scale.  Other  features 
as  indicated  in  the  accompanying  figures. 
Most  of  the  characters  of  this  specimen 
conform  with  those  detailed  by  Taylor 
(1936:239-243,  Fig.  33,  PL  15)  for  Eu- 
meces xanthi,  and  indeed  the  holotype  of 
E.  capito  closel}'  resembles  one  of  the  syn- 
types  figured  on  his  Plate  15  (Fig.  3), 
although  the  pigment  loss  has  not  attained 
the  level  there  depicted  for  a  76  mm  speci- 
men. Critical  are  the  modified  lateral 
postanals,  the  enlarged  postfemorals,  the 
low  number  (24)  of  scale  rows,  and  the 
position  of  the  dorsolateral  light  stripe  on 
the  3rd  scale  row,  in  addition  to  the  single 
postnasal  and  paired  postmentals.  No 
American  species  of  Eumeces  have  en- 
larged postfemorals;  only  a  few  Asiatic 
species   possess   them,    and   all   except   E. 


^Department    of    En\-ironmental,    Population,    and    Orgnnismic   Biology,    University   of  Colorado,    Boulder,   80302,    U.S.A. 
^Museum   National    d'Histoire   Naturelle.    25    Rue  Cuvier.    Paris.    75005,    France. 


109 


110 


GREAT  BASIN  NATURALIST 


Vol.  35,  No.  1 


Fig.  1.     Holotype  of  Eumeces  capita  Bocourt,  dorsal  view. 


xanthi  are  eliminated  from  consideration 
by  number  of  scale  rows  or  by  the  post- 
nasal-postmental  characters. 

The  only  notable  discrepancy  between 
the  holotype  of  E.  capita  and  Taylor's 
account  of  E.  xanthi  is  the  low  number  of 
dorsals  (54)  in  the  former,  as  compared 
with  the  range  (56  to  60)  for  the  latter. 
However,  northern  examples  of  E.  xanthi 
tend  to  have  fewer  dorsals  (56-59,  com- 
pared with  59-60  for  southern  examples) ; 
nevertheless,  northern  examples  tend  to 
have  22  scale  rows,  whereas  southern 
ones  usually  have  24.  We  suspect  that 
the  type  of  E.  capita  was  taken  in  more 
northern  parts  of  the  species'  range,  con- 
forming more  closely  with  the  geographic 
variant  that  bears  the  name  Eumeces 
pekinensis  Stejneger  (1924:120),  type  lo- 
cality Hsin-Lung-Shan  district,  imperial 
hunting  grounds,  Chihli  Province,  665  mi 
N  Peking,  China,  than  with  the  geograph- 
ic group  represented  by  Eumeces  xanthi 
(type  locality  Ichang,  Hupeh,  China). 
The  contact  of  frontonasal  with  frontal 
that  occurs  in  the  type  of  E.  capita  con- 
forms with  Stejneger's  description  and 
figures  (1925:49-51,  fig.  2)  for  the  three 
types  of  E.  pekinensis  and  with  Taylor's 
figure  (1936:242,  fig.  33),  in  which  the 
two  prefrontals  are  in  contact  and  there- 
fore separate  frontal  and  frontonasal. 
However,  the  figured  specimen  is  from 
the  same  district  as  the  types  of  E.  pe- 


kinensis; presumably  the  character  is  not 
taxonomically  significant.  On  the  con- 
trary, the  number  of  dorsals  and  number 
of  scale  rows  may  indeed  be  significant. 


/x\ 


I 


Fig.   2.     Dorsal  view  of  head  of  the  holotype 
of  Eumeces  capita  Bocourt. 


March  1975 


SMITH,  ET  AL.:    LIZARD  IDENTITY 


111 


Fig.  3.  Lateral  view  of  the  head  of  the  holo- 
type  of  Eumeces  capita  Bocourt.  The  minute 
spheres  abundantly  evident  in  this  figure,  and 
less  abundant  in  Figs.  2  and  4,  are  air  bubbles. 
All  photos  were  taken  of  the  specimen  under 
water. 

at  least  subspecifically;  more  material  will 
be  required  to  establish  the  nature  of  the 
variation  that  occurs  in  the  species.  Cer- 
tainly the  low  number  of  dorsals  in  the 
type  of  E.  capita  casts  no  doubt  upon 
proper  allocation  with  E.  xanthi^  although 
it  may  be  important  in  naming  the  geo- 
graphic races  of  that  species  at  some  time 
in  the  future. 

We  are  not  aware  of  more  recent 
studies  of  E.  xanthi  that  would  shed  any 
hght  upon  the  geographic  variation  of  that 
species.  The  related  species  E.  tamdaoen- 
sis  Bourret  (1937:19-21,  fig.  5)  is  very 
similar  and  may  well  be  referable  to  E. 
xanthi  as  a  geographic  race,  but  the  origi- 
nal description  does  not  note  presence  or 
absence  of  enlarged  postfemorals;  a  pe- 
culiar head  pattern  of  juveniles  may  be 
distinctive,  as  well  as  the  greenish  color 
above  and  below  in  life.  Eumeces  coreen- 
sis  Doi  and  Kamita  (1937:211-215,  figs.) 
is  not  closely  similar,  being  related  more 
closely  to  E.  chinensis  (no  postnasal,  no 
enlarged  postfemorals).  However,  the 
wide  range  of  E.  xanthi  (and  its  close 
relatives  E.  tamdaoensis,  E.  elegans  and 
E.  tunganus)  suggests  that  a  polytypic 
species  or  a  species  complex  may  be  in- 
volved, the  nomenclature  of  which  is  far 
from  stable. 

Because  of  this  primitive  state  of  knowl- 
edge of  variation  in  the  xanthi  subgroup 
(unique  in  having  enlarged  postfemorals) 
of  the  Asiatic  members  of  Taylor's  fascia- 
tus  group,  we  regard  it  unwise  to  restrict 
the  type  locality  of  E.  capita;  clearly  the 
published  designation  of  eastern  United 
States  is  in  error,  and  China  probably  em- 
braces the  lizard's  actual  origin,  but  the 


final    fixation    remains    in    the   hands    of 
future  workers. 

In  like  fashion  we  are  reluctant  to  sug- 
gest that  the  name  Eumeces  capita  be  sup- 
pressed in  order  to  preserve  the  name 
Eumeces  xanthi  despite  the  facts  that  (1) 
E.  xanthi  has  been  used  for  85  years 
whereas  (2)  E.  capita  has  never  been 
used,  except  for  its  types,  since  it  was  pro- 
posed 95  years  ago.  Even  E.  xanthi  was 
not  clearly  fixed  with  a  recognizable  spe- 
cies until  1936,  however,  when  Taylor 
demonstrated  that  it  is  the  same  as  the 
more  familiar  (even  though  more  recently 
described)  Eumeces  pekinensis  Stejneger 
(1924).  Present  decisions  are  not  limited 
an}^  longer  by  a  rigid  (and  ambiguous) 
namen  ablitum  rule.  That  rule  is  replaced 
by  this  statement:  "A  zoologist  who  con- 
siders that  the  application  of  the  law  of 
Priorit}'  would  in  his  judgment  disturb 
stability  or  universality  or  cause  confusion 
is  to  maintain  existing  usage  and  must  re- 
fer the  case  to  the  Commission  for  a  de- 
cision under  the  Plenary  Powers."  It  is 
to  be  noted  that  the  requirement  to  justify 
suspension  of  the  Law  of  Priority — use  of 
xanthi  by  at  least  five  •  different  authors 
in  at  least  10  publications  after  Taylor's 
1936  fixation— would  be  difficult  to  meet 
(see  ICZN,  1972:  185-186).  Convinced 
that  stability  of  nomenclature  is  not  a  fac- 
tor to  be  considered  in  the  present  context, 
we  recommend  that  E.  xanthi  be  replaced 
as  a  species  name  b}'^  its  senior  synonym 
E.  capita,  recognizing  that  it  is  quite  likely 
that  the  name  E.  xanthi  may  well  be  util- 
ized in  the  future  for  a  subspecies  of  E. 
capita.  Since  E.  capita  is  the  earliest  name 
applied  to  any  member  of  the  subgroup 
characterized  by  enlarged  postfemorals,  we 


Fig.  4.  Posterior  view  of  hind  leg  of  the  holo- 
type  of  Eumeces  capita  Bocourt,  showing  the 
patch    of   enlarged   scales   on   thigh. 


112 


GREAT  BASIN  NATURALIST 


Vol.  35,  No.  1 


suggest  that   it   be   designated   the   capito 
subgroup. 

Our  conclusion  not  to  recommend  sup- 
pression is  reinforced  to  a  certain  extent 
by  realization  that  the  failure  of  recogni- 
tion of  the  identit}'  of  E.  capito  for  almost 
a  hundred  years  is  not  to  be  attributed 
wholh'  to  Bocourt,  for  his  description  is 
exemplary  in  detail  and  illustration  (even 
though  the  critical  postfemoral  scale  char- 
acter was  not  noted)  and  appeared  in  a 
widely  known  work.  The  erroneous  lo- 
cality was,  of  course,  the  misleading  fac- 
tor, but  certainly  not  a  unique  one;  many 
species  have  been  properly  allocated  des- 
pite totally  misleading  type  localities.  A 
succession  of  extraordinary  circmnstances 
that  prevented  subsequent  workers  from 
reexamining  the  holotype  is  responsible 
for  the  name's  long  histor}'  as  a  nomcn 
duhium.  Had  the  name  been  proposed 
with  a  totally  inadequate  description  or  in 
a  very  obscure  outlet  onl}-  recently  dis- 
covered, it  might  be  construed  as  appro- 
priate that  it  remain  in  oblivion;  but  un- 
der the  circumstances  as  they  actually 
exist,  it  is  fully  appropriate  that  Bocourt's 
name  be  given  its  impartial  place  in  no- 
menclature, subject  only  to  the  automatic 
provisions  of  the  Code.  The  present  de- 
ficienc}^  of  an  effective  tjpe  locality  can 


readily  be  remedied  at  the  appropriate 
time,  when  an  arbitrary  designation  can 
be  proposed  in  conformance  with  detailed 
knowledge  of  geographic  variation  that  is 
now  lacking. 

Literature    Cited 

Bocourt,  Marie-Firmin.  1879.  Etudes  sur  les 
reptiles.  Miss.  Sci.  Mexique,  Rech.  ZooL. 
Livr.    6:361-440,   pis.   21-22,   22A-22D. 

BouRRET.  R.  1937.  Notes  lienDetologiques  sur 
rindochine  francaise.  Bull.  Gen.  Inst.  Publ. 
9:1-26,  figs.  1-5.  pi. 

Doi,  H.,  .AND  T.  K.AMITA.  A  new  species  of 
Eumeces  from  West  Corea.  Zool.  Mag.,  Tokyo 
49:211-215,  ill. 

GiJNTHER,  A.  1889.  Third  contribution  to  our 
knowledge  of  reptiles  and  fishes  from  the 
Upper  Yangtsze-Kiang.  Ann.  Mag.  Nat.  Hist. 
(6)4:218-229. 

ICZN  (International  Commission  on  Zoological 
Nomenclature).  1972.  Seventeenth  Interna- 
tional Congress  of  Zoology.  Monaco,  24-30 
September.  1972.  Bull.  Zool.  Nomencl.,  29: 
168-189. 

Stejneger,  L.  1924.  Herpetological  novelties 
from  China.  Occ.  Pap.  Boston  Soc.  Nat.  Hist. 
5:119-121. 

.      1925.     Chinese   amphibians   and   reptiles 

in  the  United  States  National  Museum.  Proc. 
U.S.   Nat.   Mus.  66:1-115,  figs._  1-4. 

Taylor,  E.  H.  1936.  A  taxonomic  study  of  the 
cosmopolitan  scincoid  lizards  of  the  genus 
Eumeces  with  an  account  of  the  distribution 
and  relationships  of  its  species.  Kansas  Univ. 
Sci.  Bull.  23:1-643,  figs.  1-84,  pis.  1-43. 


STUDIES  IN  NEARCTIC  DESERT  SAND  DUNE  ORTHOPTERA. 

PART  XV.  EREMOGEOGRAPHY  OF  SPANIACRIS 

WITH  BIOECOLOGICAL  NOTES 

Ernest  R.  Tinkham^ 

Abstract. —  Four  decades  of  the  author's  records  indicate  that  Spaniacris  deserticola  (Bruner)  is 
confined  within  the  periphery  of  the  Colorado  Desert.  It  is  usually  found,  near  or  within  a  few 
hundred  feet  of  sea  level,  marking  the  shore  line  of  ancient  Lake  Cahuilla  (e.xcept  for  the  Dale 
Lake  record).  The  preferred  host  plant  is  Coldenia  palmeri  growing  on  the  lower  fringes  of  bajadas, 
with  C.  plicata  on  drift  sand  being  second  in  preference.  Spaniacris  can  tolerate  sand  and  rock  tem- 
peratures of  60  C.  (believed  to  be  a  maximum  for  Colorado  Desert  life).  Mating  takes  place  at  that 
and  lower  temperatures.  When  they  are  disturbed  while  on  the  tops  of  host  plants,  their  flight  is 
low  and  direct  and  of  short  duration,  and  they  come  to  rest  on  the  torrid  soil  for  long  periods  of 
time.  The  female,  much  larger  than  the  male,  can  sustain  the  male  in  flight  while  mating.  The  study 
verified  spatial  longevity  of  Spaniacris  at  Indio,  California,  after  approximately  70  years  and  for  the 
Kane  Springs  area  after  52  years. 

Spaniacris  deserticola  (Bruner,  1906), 
one  of  the  rarest  of  Nearctic  Desert  grass- 
hoppers, was  based  on  a  single  female 
collected  at  Indio  by  H.F.  Wickham. 

In  1931,  Morgan  Hebard  (1937:376) 
instructed  me  to  make  special  efforts  to 
locate  S.  deserticola.  Late  in  the  after- 
noon of  25  August  1931,  a  large  colony 
was  located  on  a  low  sand  ridge  some 
ten  miles  east  of  Coyote  Wells  and  about 
a  mile  or  so  east  of  Plaster  City,  Cali- 
fornia. A  torrid  day  had  been  spent 
hunting  for  the  elusive  creature  on  the 
blistering  mesas.  One  had  to  blink  con- 
stantly to  keep  one's  eyes  somewhat  moist, 
and  it  was  so  hot  that  the  ubiquitous  grass- 
hopper Trimerotropis  p.  pallidipennis  flew 
from  the  top  of  one  creosote  bush  to 
another.  At  8:00  that  night,  after  sun- 
down, it  was  still  122  F  in  El  Centro. 
Hebard  records  that  I  collected  13  males 
and  7  females  for  him  that  day  and  that 
my  brother  collected  12  males  and  9  fe- 
males for  me.  In  1940  the  colony  was 
still  there,  and  on  14  August  I  took  8 
males  and  10  females.  In  May  1961  I 
could  not  find  any  trace  of  the  colony. 
During  those  intervening  years  the  High- 
way Department  had  made  a  barrow  pit 
out  of  the  sand  ridge  and  had  destroyed 
the  habitat. 

During  the  period  1949-1973  I  took  2 
males  and  1  female  in  early  June  1953 
at  or  near  the  motith  of  Palm  Canyon 
at  a  location  northward  and  across  High- 
way 111  from  the  Smoke  Tree  Ranch 
(both  locations  had  disappeared  by  1972 
due  to  residential  developments)  and  P. 
H.  Timberlake  took  a  female  here  on  24 
June  1952  and  a  male  on  21  June  1953. 


On  22  May  1954,  Dr.  John  Goodman 
and  I  were  collecting  on  the  east  side  of 
dry  Dale  Lake,  25  miles  east  of  Twenty 
Nine  Palms  and  found  a  female  nymph, 
probably  in  the  last  stadium,  on  barren 
playa,  where  sand  drifted  across  the  grav- 
el road.  In  May  1973  inspection  showed 
this  area  so  blasted  and  eroded  by  vio- 
lent sand  storms,  that  the  only  surviving 
vegetation  was  some  ancient,  dying  creo- 
sotes with  their  crowns  supported,  like 
mangroves,  on  long  exposed  roots.  It  is 
believed  that  this  colony  has  been  ex- 
terminated. It  was  the  only  one  at  a 
considerable  elevation,  as  all  others  lie 
close  to  sea  level. 

On  14  June  1964,  I  examined  a  rather 
level  sweep  of  sand  (air  temperature 
119  F)  well  covered  with  sand  mat 
{Coldenia  plicata)  some  miles  west  of 
Rice,  California,  and  one  several  miles  east 
of  the  turnoff  to  the  Iron  Mountain 
Pimiping  Station.  The  first  plant  ex- 
amined contained  a  female  Spaniacris 
(juietly  resting  ainong  the  leaves  not  more 
than  several  inches  above  the  torrid  sands. 

On  21  June  1964,  along  the  old  high- 
way about  4  miles  west  of  Thousand 
Palms,  I  found  a  single  male.  On  29  July 
1974  I  reexamined  this  area  with  sand 
mat  margining  the  south  edge  of  the  road 
and  scattered  on  the  sand  but  could  find 
no  Spaniacris,  although  that  week  I  had 
located  six  new  colonies  some  miles  east 
of  Thousand  Palms. 

On  29  June  1970  Jim  Davis  took  a 
pair  of  Spaniacris  in  Thousand  Palms 
Canyon.  The  next  day  I  found  Spania- 
cris at  that  locality  confined  to  Coldenia 
palmeri    bushes    bordering    several    hun- 


m-H\    Date    Palm    Avenue,    Indio,    California    02201. 


113 


114 


GREAT  BASIN  NATURALIST 


Vol.  35,  No.  1 


dreds  of  feet  of  an  old  gravel  road  near 
the  base  of  the  gravelly  hills.  I  took  2 
females  and  one  male.  Associated  were: 
Anconia  integra  on  Atriplex  canescens 
var.  linearis  and  A.  polycarpa;  Xeracris 
minimus  on  sandpaper  weed  (Petalonyx 
thurbcri) ;  Ciholacris  parviceps  on  the 
road  and  Tytthotyle  maculata,  the  Mal- 
pais  lubber,  nearby  among  the  boulders  of 
the  brief  bajada  at  the  foot  of  the  hills. 
One  male  was  taken  there  1  July  1971, 
and  one  male  and  one  female  on  3  July 
1972;  there  were  no  Spaniacris  there  in 
1973,  a  very  dry  year.  In  1974  this  area 
was  designated  Colony  No.  6.  (I  shall  re- 
port on  Colony  No.  6  later.) 

On  the  hot  evening  of  11  June  1973, 
Jim  Davis  and  I  made  a  night  collecting 
trip  along  the  roads  to  the  Borrego  Desert. 
One  female  Spaniacris  was  taken  on  the 
road  6  miles  west  of  Salton  City,  just  east 
of  the  radar  tower;  one  pair  was  taken 
one  mile  west  of  the  tower;  and  one 
crushed  female  was  taken  two  miles  west. 
In  the  arroyo  area  of  a  broad  Pleistocene 
valley,  about  13.5  miles  southeast  of  Bor- 
rego Springs  by  road  and  within  half  a 
mile  of  Highway  78,  another  female  was 
found  on  the  paved  road  at  night.  In  the 
late  evening  of  10  Aug.  1974,  I  surveyed 
the  adjacent  arroyo  area,  both  sides  of 
the  road,  but  found  no  evidence  of  Spania- 
cris. The  Borrego  area,  based  on  the 
parched  conditions  and  poor  showing  of 
flowers,  apparently  got  little  of  the  day- 
long drizzle  that  drenched  Coachella 
Valley  on  7  Jan.  1974. 

Theodore  J.  Cohn  contributed  some  of 
his  collection  records  as  follows:  "Imperial 
Co.,  Ca.,  3  mi.  E  Plaster  City,  at  the  Oy- 
ster Beds  Turnoff,  25  June  1965,  T.  J. 
Cohn,  1  male,  2  females.  On  ground  near 
road  around  noon,  hot  as  a  firecracker.  I 
have  stopped  there  many  times  since  and 
have  never  seen  them  again.  Lots  of 
Anconia  around  in  other  years. 

"Sonora,  Mexico,  22.3  mi.  SE  San  Luis 
Rio  Colorado,  20  June  1965,  T.  J.  Cohn, 
No.  25.  Base  of  sand  dunes  but  not  in 
them.  1  juv.  female,  last  instar.  Xeracris 
and  Coniana  also  found  here.  These  are 
the  first  set  of  dunes  east  of  San  Luis. 

"Sonora,  Mexico,  76  mi.  SE  San  Luis 
Rio  Colorado  (14  mi  NW  Los  Vidrios). 
14  June  1966,  T.  J.  Cohn,  No.  16.  Very 
sparse  vegetation  on  moderately  sandy 
soil,  not  far  from  stabilized  dunes.  This 
was  around  a  corral  a  few  hundred  yards 


south  of  the  road,  near  the  next  set  of 
dunes,  1  female.  Lots  of  what  I  think  is 
Coniana  here." 

The  sand  dunes  22.3  miles  SE  San  Luis 
are  in  large  part  in  Arizona;  the  Inter- 
national fence  terminating  on  some  rocky 
hills  astride  the  International  Boundary. 
The  best  dunes  are  in  Arizona  immed- 
iately north  of  these  hills.  I  have  collec- 
ted there  on  many  occasions  but  have 
never  found  evidence  of  Spaniacris.  This 
should  be  the  best  location  to  discover 
Spaniacris  in  Arizona.  At  the  76-mile 
location,  a  high  ridge  of  aeolian  dunes 
sweeps  far  inland  into  the  area  from  the 
southwestern  coastal  regions. 

Notes  on  the   1974   Survey 

Colony  No.  1.  Found,  25  July  3  miles 
north  of  Indio,  California,  on  Monroe 
Street  on  C.  palmeri  on  the  north  side  of 
the  dyke  protecting  the  Coachella  Valley 
branch  of  the  All  American  Canal 
within  fifty  feet  of  the  paved  road.  The 
Colony  was  at  the  southern  edge  of  a  con- 
siderable bajada  that  gently  slopes  down 
from  the  Indio  Hills  over  an  area  of  many 
miles.  This  location  is  directly  south  of 
the  Curtis  Desert  Palms  Oasis,  whose  ex- 
istence is  threatened  by  the  huge  Massey 
Rock  and  Sand  gravel  pit.  The  plant  life 
is  typical  of  that  described  fully  under  No. 
2  below  but  is  not  as  extensive.  Thus,  the 
presence  of  Spaniacris  deserticola  for  Indio 
was  verified  almost  70  years  after  Bruner 
described  it  in  1906;  I  have  not  been  able 
to  find  the  date  of  Wickham's  collections 
prior  to  1906. 

Colony  No.  2.  That  afternoon,  at  the 
powerline  crossing  of  Washington  Street, 
4  miles  north  of  Interstate  10  and  about 
10  miles  northwest  of  Indio,  I  found  a 
much  larger  colony,  here  designated  No. 
2.  The  location  was  similar  to  that  of  No. 
1,  representing  the  southern  edge  of  the 
long  bajada  margining  the  south  side  of 
the  Indio  Hills.  The  desert  vegetation 
consists  of  shrubs,  living  ephemerals,  and 
the  dead  skeletons  of  spring  ephemerals. 

The  desert  shrubbery  was  composed  of 
many  widely  scattered  Coldenia  palmeri 
growing  along  the  narrow,  poorly  paved 
powerline  road  as  well  as  along  the  mar- 
gins of  a  shallow  dry-  wash  with  fewer 
scattered  clumps  of  creosote  {Larrea  di- 
varicata),  sandpaper  weed  (Petalonyx 
thurheri),  Burrobush  (Franseria  dumosa), 


March  1975 


TINKIIAM:    DESERT   GRASSHOPPER 


115 


<uid  even  more  rarely  encelia  {Encelia 
j(irinosa),  desert  sweet  {Bebbia  juncea), 
( lieesebush  (Hymenoclea  salsola)  and  Cal- 
ifornia dalea  (Dalea  calif ornica) .  Living 
t'[ihemerals  were:  inflated  stem  {Eriogo- 
tiurn  inf latum) ,  Spanish  needle  (Palofoxia 
linearis),  velvet  rosette  {Psathy rotes  ra- 
/nosissima),  Stillingia  spinulosa,  and  Ti- 
des troernia  oblongi folia;  all  but  Tidestro- 
er/iia  were  in  bloom.  Dry  spring  ephe- 
inerals  were:  desert  gold  (  Geraea  ca- 
iiescens),  forget-me-nots  {Cryptantha  cos- 
tiita  and  maritima),  Thomas  buckwheat 
[I-.riogonum  Thoniasi),  brown-eyed  prim- 
rose {Oenothera  clavaeformis) ,  blazing 
star  (Mentzelia  multiflora) ,  and  perhaps 
others.  The  presence  of  the  flowering 
cphemerals  in  July  and  August  on  the 
blazing  Colorado  Desert  can  be  explained 
(iiily  by  the  slow  drizzling  rain  (1.56 
inches)  on  7  Jan.  1974,  which  penetrated 
to  at  least  three  feet.  On  29  July,  I 
foimd  damp  sand  one  foot  down.  Al- 
though this  soil  moisture  w^as  a  great  boon 
in  reviving  dying  shrubs  (transects 
anywhere  will  show  65-75  percent  dead 
or  partly  dead),  germination  of  ephemeral 
seeds  was  poor  because  of  the  cold  Janu- 
ary weather.  The  deep  penetration  of 
moisture  accounts  for  the  flowering  of 
Palofoxia  and  Eriogonum  in  midsummer 
and  undoubtedly  was  responsible  for 
breaking  the  diapause  of  Spaniacris  and 
other  acridid  eggs. 

On  25  July  the  two-acre  area  surveyed 
( approx.  100  by  80  yds.)  contained  at 
least  6  pairs  of  Spaniacris  of  which  I  took 
3  males  and  2  females,  leaving  the  rest 
for  propagation.  Next  day,  26  July, 
4:15  to  5:00  pm,  slightly  hazy,  tempera- 
ture 45  C,  soil  57  C.  Quite  a  few  adults 
noted  but  none  taken. 

On  1 7  Aug.  a  hot  wind  was  blowing 
from  the  southeast.  At  5:00  pm,  air  tem- 
perature was  44  C,  soil  54  C;  5:20  pm, 
air  44  C,  soil  53  C.  Photographed  Spania- 
cris with  Tri-X  film.  Found  Coniana 
snowi  and  for  the  first  time  Anconia  In- 
tegra female  on  Coldenia  palmeri.  Xera- 
cris  minimus  also  sometimes  found  but 
more  often  taken  from  Petalonyx  thurb- 
eri. 

On  20  Aug.  5:00  to  6:30  pm,  air  38  C, 
soil  54  C.  Found  only  a  few  pairs  in  the 
original  two-acre  area.  Extended  the  col- 
ony southeastward  for  3/10  mile,  where 
I  found  two  pairs,  and  westward  across 
Washington    Street    at    least    100    yards. 


where  I  found  two  more  pairs,  one  pair 
in  copula,  at  6:30  pm.  Total  area  covered 
by  Colony  No.  2,  at  least  one  half  mile 
in  length  and  about  100  yards  in  width. 
Photographed  the  species  with  Pana- 
tomic-X  film  as  Tri-X  was  much  too 
fast  for  glaring  light  of  the  desert. 

On  24  Aug.  5:30  to  6:30  pm  visited  the 
site  with  Chas.  Neeley,  photographer.  At 
5:30  pm,  air  41  C,  soil  51  C.  First  male 
found  within  a  few  feet  of  the  road.  One 
area  four  feet  square  formed  by  two  con- 
tiguous C.  palmeri  plants  held  two  females 
and  one  pair  mating;  this  is  the  greatest 
concentration  ever  found  for  this  species. 
On  this  date  the  original  two-acre  area 
contained  eight  females  and  nine  males 
despite  the  fact  that  three  males  and  two 
females  were  removed  on  25  July.  Since 
that  date  all  specimens  left  for  propaga- 
tion of  the  species.  Little  change  in  vege- 
tation noted  since  25  July.  Fairly  hot, 
slight  breeze  from  the  west. 

On  18  Sept.,  1:15  to  2:00  pm,  air  41  C, 
soil  51  C.  One  male  found  near  road,  one 
female  at  least  200  feet  away.  This  fe- 
male recognized  by  the  slightly  shorter 
tegmina.  Trimerotropis  p.  pallidipennis 
present  for  the  first  time  this  summer; 
four  females  noted.  These  had  probably 
migrated  into  the  area  as  no  nymphs  of 
any  acridids  had  been  noted  previously. 

On  29  Sept.,  1:15  to  2:00  pm,  fairly  hot 
wind,  temperature:  air  39.5  C,  soil  52  C. 
Examined  all  C.  palmeri  bushes  in  the 
original  two-acre  area.  On  the  91st  plant 
the  female  noted  on  18  Sept.  was  found 
and  some  distance  away  on  the  104th 
plant  found  a  pair,  in  copula,  missed  on 
18  Sept.  Did  not  find  the  male  found  18 
Sept.  near  the  road,  which  had  been  at 
least  200  feet  away  from  the  others.  It  is 
obvious  that  Spaniacris  is  quite  localized 
in  its  movements  especially  in  the  late 
summer. 

On  4  Oct.,  brisk  west  wind,  air  31.5  C, 
soil  54  C  on  sunny  side  of  a  C.  palmeri 
plant  in  wind-sheltered  position.  Examined 
about  120  plants;  apparently  only  one 
female  surviving  at  this  late  date;  this  one, 
the  one  noted  on  18  and  29  Sept.,  was  still 
in  the  same  area.  I  found  female  by 
waving  my  net  back  and  forth  near  each 
plant.  While  doing  so,  I  was  aware  of 
the  slightest  movement  of  something 
dropping  an  inch  or  two  from  the  end 
of  a  branch  to  the  ground.  This  female 
was  hiding  under  the  plant  on  the  sandy 


116 


GREAT  BASIN  NATURALIST 


Vol.  35,  No.  1 


soil.  This  weis  the  first  time  I  had  ob- 
served this  habit  and  the  first  record  of 
this   species   for  October. 

On  11  Oct.,  11:20  to  11:40  am,  much 
cooler.  Had  time  only  to  check  over  those 
bushes  mostly  likely  to  hold  Spaniacris 
perched  on  top  of  plant  as  they  almost 
alw^ays  were.  Found  none;  believe  colony 
had  died  out  for  1974.  On  this  late  date 
vegetation  was  surprisingly  like  that  noted 
in  late  July,  with  E.  inflatum,  Palofoxia 
linearis,  and  P.  ramosissima  still  bloom- 
ing, but  Stillingia  has  been  dead  and 
brown  since  mid-September. 

Colony  No.  3.  Located  25  July,  about 
one  mile  northwest  of  Colony  No.  2,  on 
the  south  side  of  Washington  Street  as  it 
swings  west  to  join  the  east  end  of  Ramon 
Road  at  the  turnoff  to  Thousand  Palms 
Canyon.  The  colony  comprised  about 
600  yards  of  C.  palmeri  on  the  east  and 
west  slopes  of  a  gentle  hill  with  a  few 
plants  of  Dalea  californica  and  D.  emoryi 
here  and  there.  The  road  crosses  the  south 
marginal  area  of  the  Indio  Hill's  bajada. 
The  300  yards  of  the  east  slope  contained 
at  least  five  pairs  of  Spaniacris;  one  pair 
taken;  w^est  slope  at  least  two  pairs;  one 
pair  taken.  When  disturbed  this  species 
never  flies  to  another  C.  palmeri  bush 
but  always  alights  on  the  very  hot  soil. 
Flight  is  low  and  direct,  the  wings  show- 
ing rather  deep  azure.  On  the  food  plant 
it  usually  orients  the  long  axis  of  its  body 
parallel  to  the  rays  of  the  sun  so  that  us- 
ally  only  the  face  is  exposed  to  the  full 
effect  of  the  very  hot  sun.  Under  the  cir- 
cumstances this  reduces  exposure  to  the 
minimum,  and  the  very  long  legs  help 
elevate  the  body  from  the  extremely  hot 
soil  tempartures.  Spaniacris  is  not  wary: 
the  cautious  photographer  can  move  the 
macrolens  of  his  camera  to  within  8  inches 
of  the  grasshopper.  Trimerotropus  p. 
pallidipennis  seldom  permits  approach 
closer  than  10  feet. 

Colony  No.  4.  Found  29  July,  located 
just  east  of  the  Thousand  Palms  (Canyon 
Turnoff.  About  one-fifth  of  an  acre  f)f 
C.  palmeri  fairly  widely  scattered  with 
a  few  creosote  interspersed;  2:  30  pm,  air 
37  C;  (had  been  overcast  in  am),  soil  58 
C.  This  small  area  (100  x  100  ft)  re- 
vealed at  least  9  females  and  11  males, 
some  of  which  were  on  very  hot  ground. 
Most  of  the  females  were  on  the  plants. 
While  I  was  taking  temperature  readings, 
one  pair,  in  copula,  sat  on  a  rock  (58  C) 


for  at  least  five  minutes  without  showing 
any  signs — as  other  acridids  quickly  do — 
that  the  rock  was  hot.  Evidence  seems  to 
indicate  that  Spaniacris  can  tolerate  more 
heat  than  can  any  other  animal  living 
on  the  torrid  Colorado  Desert.  Temper- 
ature one  foot  above  soil  level  was  45  C. 
Still  on  margin  of  the  bajada. 

Colony  No.  5.  Found  25  July,  one  half 
mile  west  of  Turnoff  to  Thousand  Palms 
Canyon,  4:30  to  5:00  pm,  air  45  C,  soil  55 
C.  Margin  of  the  same  bajada  but  sandier 
than  in  previous  colonies.  Flora:  C.  pal- 
meri with  scattered  creosote,  D.  califor- 
nica, cheesebush,  Dicoria  canescens,  Atri- 
plex  c.  linearis,  and  dead  sand  verbena 
{Abronia  villosa) .  One  pair  in  copula,  fe- 
male could  fly  only  a  few  feet  with  the 
male.  Took  mating  pair  and  left  2  males 
and  one  female. 

Colony  No.  6.  Visited  once  on  25  July. 
This  is  the  location  mentioned  previously 
under  dates  of  1970-1973,  within  the 
mouth  of  Thousand  Palms  Canyon.  Ob- 
served 1  female  and  1  male  perched  on  C. 
palmeri  along  margin  of  old  gravel  road 
previously  described.  Did  not  disturb. 
Noted  Anconia  Integra  on  Atriplex  c.  li- 
nearis, Xeracris  minimus,  and  flushed  a 
female  Tytthotyle  maculata  out  of  lux- 
uriant creosote  bush.  No  temperature 
readings  were  made  on  the  first  day  of 
1974  survey. 

On  Saturday  afternoon,  27  July,  I  con- 
tinued my  Spaniacris  survey.  Dri^■ing 
northward  on  Washington  Street  and  then 
westward,  I  passed  colonies  2,  3,  and  4, 
then  turned  right  on  Thousand  Palms 
Canyon  Road  to  pass  Colony  6  and  reach 
the  Dillon  Road  junction  in  about  four 
and  one-half  miles.  En  route  I  examined 
400  yards  of  fine  habitat  of  C.  palmeri 
on  both  sides  of  the  road  about  1.5  miles 
north  of  Paul  Wilhelm's  Oasis,  but  found 
no  specimens.  As  this  was  about  as  fine 
a  habitat  as  I  had  seen,  I  wondered  why 
no  Spaniacris.  Apparently,  since  it  is  so 
rare,  localized,  and  sedentary  in  habits 
and  because  this  location  is  several  hun- 
dred feet  higher  than  Colony  6,  it  has  not 
been  able  to  ])enetrate  up  through  this 
rather  broad  and  low  canyon. 

A  survey  of  Dillon  Road  from  the  Thou- 
sand Palms  Canyon  junction  (approx- 
imately 600  feet  elevation)  southeasterly 
1 1  miles  to  where  sand  is  first  encountered 
at  about  sea  level,  revealed  no  Spaniacris. 

Colony  No.  7.    Shortly  beyond,  at  11.7 


March  1975 


TINKHAM:    DESERT   GRASSHOPPER 


117 


miles  from  the  Dillon-Thousand  Palm 
Can^'on  junction,  I  examined  200  yards 
of  C.  plicata  and  found  on  one  ])lant  one 
male  Spcmiacris  and  1  female  Coniana 
sjiowi  resting  on  top  of  the  low  bush  with- 
in one  foot  of  each  other.  Air  temperature 
at  6:00  pm,  45  C  with  hot  wind  blowing, 
II'  C  when  calm,  leaf  surface  44.2  C.  This 
location  was  3.6  miles  northwest  of  free- 
way and  about  5  miles  northwest  of  Coa- 
chella. 

Both  sides  of  Dillon  Road,  edged  with 
sparse  patches  of  Coldenia  palmeri  and  C. 
plicata,  were  examined  from  Colony  No. 
7  to  the  junction  with  Interstate  10  (3.4 
miles)   and  no  Spaniacris  were  found. 

On  29  Jul}',  I  examined  old  Highway 
99  (now  Vamer  Road)  from  Thousand 
Pahiis  west  to  Garnet  Hill  (13.3  mil»s) 
and  found  no  trace  of  Spaniacris  on  Col- 
denia palmeri  and  C.  plicata,  which  often 
formed  nice  colonies  along  the  margin  of 
the  road. 

On  10  Aug.  I  drove  along  Highway 
86  to  the  Kane  Springs  area,  then  west- 
ward on  Highway  78,  but  failed  to  find 
evidence  of  Spaniacris  other  than  at  Col- 
ony No.  8,  noted  below. 

Colony  No.  8.  At  about  7.5  miles  from 
Kane  Springs,  6  miles  northwest  of  the 
junctions  of  Highways  78  and  86,  I  ex- 
amined 100  }^ards  of  C.  plicata  growing 
on  south  sandy  side  of  the  road  and  found 
a  female  Spaniacris  that  flew  about  30 
feet,  its  azure  wdngs  showing  in  low 
direct  flight,  to  land  on  the  bare  clay  soil 
typical  of  the  area.  Later  I  examined  a 
small  flat  wash  with  C.  plicata,  creosote, 
and  smoke  tree  (Dalea  spinosa)  about 
six  miles  northwest  of  Kane  Springs  that 
admirably  fits  Hebard's  (1937:377)  loca- 
tion where  he  took  this  species.  Nothing 
was  found  here. 

Because  of  drought  conditions  there  was 
very  little  Coldenia  west  on  Highway  78 
and  what  there  was  occurred  chiefly  at 
the  sea  level  line  some  6  to  7  miles  west 
of  the  Kane  Springs  junction.  Late  that 
night,  at  about  10:30  pm,  I  examined  by 
Coleman  lantern  light  the  area  immed- 
iately east  of  the  radar  tower  where  Spa- 
niacris  had  been  encountered  on  the  night 
of  11  June  1973,  but  found  nothing.  I  had 
examined  about  10  acres  of  this  area  in 
late  afternoon  of  20  July  1974;  but,  al- 
though it  was  in  interesting  habitat,  I 
could  find  no  Spaniacris.  Farther  on,  4 
miles  west  of  Salton  City,  I  examined  300 


yards  of  roadside  Coldenia  plicata  with 
Coleman  lantern  and  found  nothing  but 
Coniana  snowi.  I  had  also  examined  this 
area  on  July  20. 

Summary  and  Conclusions 

Biology.  The  adult  Spaniacris  is 
now  known  from  early  June  until  early 
October.  Records  of  nymphs  in  Ma}'  rep- 
resent the  two  last  nymphal  stadia.  No 
small  nymphs  have  ever  been  found.  The 
number  of  ova  laid  per  feinale  is  small,  so 
the  population  potential  is  likewise  small. 
Habits.  The  low  direct  flight  of  rather 
short  duration  is  diagnostic  for  the  genus. 
During  flight  the  wings  appear  darker 
azure  than  they  appear  on  close  inspec- 
tion. Flight  is  almost  always  from  the  top 
of  the  Coldenia  plants  to  the  desert  soil, 
where  they  will  rest  immovable  many 
minutes,  seemingh'  able  to  tolerate  per- 
fectly the  torrid  soil  temperatures.  Spania- 
cris is  not  a  wary  grasshopper  and  can 
be  slowly  and  easily  approached  to  within 
six  inches  for  macrolens  photography. 
However,  cpiick  lateral  movements  will 
alarm  them  and  cause  flight.  The  female 
of  this  s])ecies  is  probably  the  only  grass- 
hopper that  can  sustain  flight,  carrying 
the  male,  while  mating.  I  never  failed  to 
find  mating  pairs  in  the  larger  colonies 
during  afternoon  and  late  evening  hours. 
No  observations  were  made  in  the  early 
morning.  Mating  was  observed  froin  late 
July  to  late  September.  Oviposition  was 
not  observed.  During  the  hotter  portions 
of  the  day,  from  late  morning  until  late 
evening,  females  oriented  themselves  so 
that  the  long  axis  of  the  body  was  paral- 
lel to  the  rays  of  the  sun.  In  such  positions 
onl}'  the  front  of  the  long  face  w^as  ex- 
posed to  the  full  effect  of  the  rays.  Onl}' 
on  rare  occasions,  and  that  in  the  late  eve- 
ning, did  I  observe  males  and  females  rest- 
ing on  the  shady  side  of  their  host  plant 
(Fig.  4). 

Habitat.  The  marginal  fringes  of  great 
bajadas,  where  the  soil  is  partly  of  rock, 
sand,  and  cla}',  seem  to  be  the  preferred 
habitat.  Spaniacris  has  been  taken  in 
purely  drift  sand  locations  near  Rice 
(Figs.  1  and  2),  Dale  Lake,  and  the  north- 
western Sonora,  Mexico  (records  of  Dr. 
Cohn).  The  preferred  altitudinal  habitat 
appears  to  be  at  or  slightly  abo^e  the  sea 
level  contour  line  in  the  Colorado  Desert; 
no   captures   have   been   made  below   sea 


118 


GREAT   BASIN    NATURALIST 


Vol.  35,  No.  1 


..^^Efc^i^- 


m 


Figs  1-8.  Habit  and  liabitat  photographs  of  Spaniacris  deserticola:  1,  Habitat  of  drift  sand, 
with  C.  plicata  in  foreground,  some  miles  west  of  Rice.  Ca..  14  June  1964,  119  F  m  shade;  2,  Fe- 
male resting  in  top  of  C.  plicala  at  Fig.  1  locality;  3,  Female  (by  Chas  C.  Neeley).  24  Aug.  1974, 
late  evening  at  Colony  No.  2;  4,  Female  resting  on  C.  palmeri  in  shade  of  settmg  sun.  Colony  No. 
4,  29  July  1974,  taken  with  Vivitar  Strobe  Flash  No.  292  with  white  cloth  filter;  5,  Female  on  hot 
gravelly  soil.  Colony  No.  2.  20  Aug.  1974.  soil  surface  54  C;  6,  Male.  Colony  No.  6,  30  June 
1970;  7,  Mating  pair.  Colony  No.  4,  29  July  1974,  resting  on  rock  surface,  58  C,  2:15  pm;  8, 
Habitat.  Colony  No.  4,  C.  palmeri  in  foreground;  some  Creosote  in  background,  looking  north  up 
Thousand  Palms  Canyon.  29  July  1974,  2:30  pm.  All  photographs  taken  by  Ernest  R.  Tink- 
ham  except  No.  3,  which  is  by  Chas.  C.  Neeley. 


March  1975 


TINKHAM:    DESERT   GRASSHOPPER 


119 


level.  The  only  records  of  considerable 
elevation  (±  1000  feet)  appear  to  be  the 
captures  at  Dale  Lake  and  near  Rice. 
There  are  no  records  for  sand  dune  situ- 
ations. Thus,  the  sea  level  contour  rep- 
resents the  ancient  shore  line  of  Pleisto- 
cene  fresh  water  Lake  Cahuilla,  which 
dried  up  hundreds  of  years  ago  to  leave 
the  salt  deposits  of  the  Salton  sea  depres- 
sion before  flooding  in  1903.  It  is  obvious, 
because  Spaniacris  is  still  largely  con- 
fined to  the  old  beach  line,  that  its  powers 
of  dissemination  must  be  rather  poor.  This 
seems  partly  accounted  for  by  its  seden- 
tary habits  (as  noted  under  Colony  No. 
()),  by  its  low  egg  potential,  b}"  the  restric- 
ted habitat  of  its  chief  host  plant  (C. 
palmeri),  by  the  torrid  temperatures,  by 
the  great  aridity  of  its  habitat,  and,  per- 
haps, b}'  other  factors  as  well. 

Host  Plants.  Spaniacris  appears  to  be 
strictly  confmed  to  Coldenia  palmcri  and 
C.  plicata.  The  Palmer  coldenia  seems  to 
be  the  preferred  host  because  its  distribu- 
tion is  closely  related  to  the  sea-level  beach 
line  of  ancient  lake.  Other  factors  in  the 
choice  of  host  plants  appear  to  be  the  size 
and  greater  height  of  C.  Palmeri,  which 
provides  green  foliage  all  summer  long, 
and  the  edaphic  conditions  which  provide 
i^reater  protection  than  the  sand  (inhab- 
ited by  C.  plicata)  for  the  eggs  during 
drought-induced  diapauses.  Very  little  is 
known  about  the  diapause  in  the  eggs  of 
desert  acridids.  On  the  other  hand,  sand 
appears  to  be  a  poor  medium  for  the  pro- 
tection of  eggs,  especially  if  laid  close  to 
the  sand   surface. 

Temperature  Tolerance.  Spaniacri.^  ap- 
pears to  tolerate  torrid  soil  and  rock  tem- 
j^eratures  of  140  F  (60  C),  as  portrayed 
by  the  mating  pair  in  Photo  7,  without  ex- 
hibiting any  of  the  signs  of  intolerance 
(such  as  the  lifting  of  tarsi)  that  most 
other  desert  acridids  would  soon  portray 
under  such  situations.  Its  extremely  long 
logs  (for  an  acridid)  must  be  partly  re- 
sponsible for  this  heat  tolerance.  Thus, 
the  evidence  seems  to  indicate  that  Spania- 
cris can  tolerate  higher  temperatures  than 
any  other  life  form  associated  with  the 
Colorado,  our  hottest  desert. 

Spatial  Longevity.  The  evidence  pro- 
vided by  Colonies  1  and  8  indicates  that 
Spaniacris  has  survived  in  the  Indio  area 
for  75  years,  and  in  the  location  a  few 
miles  northwest  of  Kane  Springs  for  50 
years,  which  is  more  than  half  the  time 


of  American  occupation  of  the  territory. 
However,  in  the  areas  near  Dale  Lake 
and  Palm  Springs  and  west  of  Thousand 
Palms  there  is  evidence  that  the  destruc- 
tive practices  of  man  have  extirpated 
three  colonies  of  Spaniacris  within  the 
past  two  decades.  These  practices  in- 
clude scraping  off  large  areas  of  desert 
shrubbery,  asphalt  paving  of  huge  areas, 
and  many  other  methods  well  knowji  to 
conservationists,  all  of  which  helped  pro- 
duce the  increasing  droughts  and  sand- 
storms of  the  past  two  decades.  It  is  also 
apparent  that  Spaniacris  survived  many 
millennia  in  the  Colorado  Desert  before 
the  advent  of  the  white  man. 

Faunal  Designation.  Spaniacris  ap- 
pears strictly  confined  to  the  Colorado 
Desert,  its  zonal  distribution  being  con- 
fined largely  to  the  old  beach  line  of 
Pleistocene  Lake  Cahuilla,  which  is  also 
at  sea  level. 

Orthopteran  Associates.  The  chief  as- 
sociates are:  Coniana  snowi  especially  on 
sandier  habitats  dominated  by  C.  plicata; 
Xeracris  jnininnis.  showing  a  preference 
for  PctaJonyx  thurhcri;  and,  rarely,  the 
saltbush  grasshopper  (Anconia  integra), 
which  prefers  Atriplex  spp.  Also  present 
are  soil-dwelling  species  such  as  the  Ar- 
royo Grasshopper  (Cibolacris  parviceps), 
the  ubiquitous  Trimerotropis  p.  pallidi- 
pcnnis,  rarely  the  Malpais  lubber  {Tyt- 
thotyle  maculata)  as  in  Colony  6,  and  the 
ground   mantid    {lAtancutria   minor) . 

Enemies.  Since  the  1974  surve}-  dem- 
onstrated that  many  colonies  exist  along 
road  margins  favored  by  C.  paJmeri  and 
(where  sandier)  C.  plicata,  evidence  in- 
dicates that  many  Spajiiacris  are  crushed 
by  passing  cars,  especially  at  night,  when 
there  seems  to  be  considerable  nocturnal 
activity  among  desert  acridids.  As  the 
crested  lizard  is  strictly  herbivorous,  there 
seems  to  be  no  evidence  that  it  would 
feed  on  Spaniacris;  but  the  presence  of  ze- 
bratails  and,  rarely,  sand  dune  lizards  of 
the  genus  IJma  in  sandy  habitats  where  C. 
plicata  is  growing  may  jiose  a  threat  to 
the  early  stages  of  Spaniacris,  about  which 
nothing  is  known. 

Rkfi:hences 

Bruner,  L.  1906.  Ramona  (not  of  Casey  1886 
rColcoptcral ) ,  Biol.  Ccntr.  Amer.,  Orth.  2: 
186-187. 

Heiwrd,  M.     1937.     Studios  in  Orthoptera  which 


120  GREAT   BASIN    NATURALIST  Vol.  35,  No.  1 

occur  in  North  America  north  of  the  Mexican  figs.  4,  5. 

Boundary.  IX.  A  new  generic  name  for  the  Rehn,  J.  A.  G.,  and  H.  J.  Grant,  jr.  1960.  An 
group  Anconiae  (Oedipodinae,  Acrididae).  additional  tribe  of  the  Romaleinae  (Orth:  Ac- 
Trans.    Amer.    Ent.    Soc.    63(103)  :375-378,    23,  rididae).   Notulae  Naturae  327:1-4,  5   text-figs. 


ROOSTING   BEHAVIOR   OF  MALE  EUDERMA   MACU LATUM 
FROM  UTAH 


Richard  M.  Poche^  and  George  A.  Ruffnei-- 

Abstract. —  Eight  spotted  bats  {Euderma  maculatum)  were  captured  and  released  along  the  Fort 
Pierce  Wash,  Washington  County,  Utah,  in  August  1974.  Observations  indicated  that  Euderma  roosts 
in  cracks  and  crevices. 


According  to  Easterla  (1970,  1973), 
the  spotted  bat  {Euderma  maculatum) 
probably  utilizes  cracks  and  crevices  as 
roosting  sites  in  Big  Bend  National  Park, 
Texas.  Observations  by  Poche  (1974)  in- 
dicate that  the  spotted  bat  inhabits  similar 
daytime  retreats  in  Utah. 

Between  12  and  15  August  1974,  eight 
spotted  bats  were  netted  over  Fort  Pierce 
Wash,  Utah.  All  were  sexed  and  marked 
using  a  wing  perforating  nmnbering  sys- 
tem. Only  one  female  (apparently  post- 
partum) was  obtained,  and  six  of  the  seven 
males  captured  were  scrotal. 

The  mean  weight  for  spotted  bats  cap- 
tured was  13.9  g,  with  a  range  of  14.8  to 
13.6  g.  After  marking,  the  bats  were  re- 
leased individually,  and  the  path  of  flight 
was  followed  with  binoculars.  The  first 
individual  set  free  on  13  August  disap- 
peared into  a  narrow  crack  along  the 
steep-walled  canyon.  Esterla  (1973)  re- 
ported similar  postrelease  behavior  in  Big 
Bend  National  Park. 

A  second  Euderma  flew  west  of  the  Fort 
Pierce  ruins  when  released  and  landed  on 
the  near-vertical  walls  of  the  Navajo  sand- 
stone cliffs,  whereupon  the  bat  walked 
about  easily  in  search  of  a  crevice.  Easter- 
la (1972)  'and  Parker  (1952)  previously 
reported  Euderma  walking  over  horizontal 
surfaces;  but  the  bat  that  we  released 
searched  over  the  vertical  wall  with  great 
facility.  J.  S.  Findley  (pers.  comm.)  re- 
ported similar  observations  of  captive  spot- 
ted bats  walking  over  volcanic  rocks  which, 
however,  are  more  porous  than  sandstone. 
As  the  second  bat  climbed  the  wall,  it  ap- 
peared to  use  its  ears  as  probes  in  search- 
ing for  a  crevice.  After  approximately 
two  minutes  of  crawling  about,  the  bat 
crept  into  a  narrow  crack  about  3  cm  wide. 

A  third  spotted  bat,  when  released  on 
14  August,  flew  south  of  the  wash  and 
went  out  of  view  behind  a  large  boulder. 
After  five  minutes  of  searching,  we  lo- 
cated the  animal  beneath  a  rock  about  50 


cm  in  diameter.  It  apparently  had  backed 
underneath  the  fallen  rock.  The  bat  was 
well  concealed  and  ordinarily  would  not 
have  been  noticed. 

The  fourth  bat  we  released  flew  under 
a  large  boulder  on  the  side  of  a  steep  in- 
cline. It  was  found  hanging  by  its  feet  at 
a  15  degree  angle  on  the  side  of  the  rock. 

The  fifth  bat,  liberated  on  14  August, 
flew  north  of  the  wash  and  up  the  steep 
slopes.  This  animal  landed  on  the  face  of 
the  cliff,  and  like  the  second  individual, 
it  engaged  in  seeking  out  a  crack  into 
which  it  could  withdraw.  Because  of  the 
observation  distance,  it  was  impossible  to 
detect  which  crack  the  bat  selected.  We 
climbed  the  cliff  and  in  fifteen  minutes 
located  the  Euderma.  The  bat  had  moved 
into  a  narrow  angling  fracture  and  was 
detected  by  blowing  into  the  crack.  This 
procedure  produced  a  loud  clicking  sound 
by  the  bat,  typical  for  the  species  when 
disturbed. 

On  15  August  1974,  a  sixth  spotted  bat 
(female)  was  netted,  and  numerous  ecto- 
parasites were  noted.  The  bat  escaped 
from  the  holding  bag  while  on  the  hood 
of  our  truck.  Another  male  collected  the 
same  morning  hosted  numerous  mites. 
These  were  later  identified  as  Cryptonys- 
sus  desuliorious.  and  this  was  the  first  re- 
port of  an  ectoparasite  associated  with  the 
spotted  bat  (Radovsky  and  Poche,  1975). 
The  male  also  had  a  large  (3  to  5  mm) 
swollen  infection  on  the  right'  forearm, 
near  the  elbow.  This  individual  was  re- 
leased at  2000  hours  at  Fort  Pierce,  and  it 
flew  approximately  30  m  down  the  north 
side  of  the  wash  and  landed  in  a  depres- 
sion on  Navajo  sandstone.  It  immediately 
climbed  about  the  near-vertical  cliff  face, 
looking  for  a  crevice.  As  expected,  the 
poUex  appeared  to  serve  as  the  main  tool 
for  grasping. 

After  several  unsuccessful  attempts  to 
locate  a  crack  large  enough  to  crawl  into, 
the  Euderma  appeared  to  be   "nervous." 


^Enviionmontal     Sciences     Division.     Stoanis-Roger, 
-Museum    of    NorthciTi    .'\rizona,    Harold    S.    Coltr 


Inc.,     P.O.   5888,   nenvcr,   Colorado  80217. 
n    Rcscardi  Center.  Flagslaff,  Arizona  86001 


121 


122 


GREAT    BASIN    NATURALIST 


Vol.  35,  No.  1 


While  we  were  attempting  to  photograph 
the  bat,  it  flew  from  the  wall  to  a  ledge 
with  an  overhang  of  3  meters.  At  this 
time,  we  abandoned  pursuit  rather  than 
harass  the  bat  any  further. 

Poche  and  Baillie  (1974)  and  Poche 
(1975)  reported  observations  indicating 
that  the  natural  roost  of  the  spotted  bat  in 
the  Utah-Arizona  region  is  small  cracks 
and  crevices.  The  findings  presented  here 
lend  evidence  to  the  validity  of  these  earli- 
er notions.  The  fact  that  three  bats  were 
observed  in  active  search  for  factures 
while  they  were  suspended  almost  verti- 
cally, further  supports  the  suggestion. 

We  wish  to  express  our  thanks  to  the 
Nevada  Power  Compan}'  for  financing 
this  study.  C.  E.  Baker  and  D.  J.  Schmidly 
provided  helpful  comments  in  reviewing 
the  original  draft. 


LiTER.'VTURE  Cited 

E.'XSTERLA,  D.  A.  1972.  First  records  of  the  spot- 
ted bat  in  Texas  and  notes  on  its  natural  his- 
tory.   Am.  Midi.  Nat.  83(l):306-08. 

.     1973.     Ecology     of     the     18     species    of 

Chiroptera  at  Big  Bend  National  Park,  Texas. 
Northwest  Mo.  State  Univ.  Studies  34(4): 
Part  II. 

P.ARKER,  H.  C.  1952.  Two  new  records  of  the 
spotted  bat  in  California.  J.  Mammal.  33(4): 
480-82. 

Poche,  R.  M.,  and  G.  L.  Baillie.  1974.  Notes 
on  the  spotted  bat  (Euderma  rnaculatum) 
from  southwest  Utah.  Great  Basin  Nat.  34: 
254. 

Poche,  R.  M.  1975.  New  record  of  Euderma 
rnaculatum  from  Arizona.  J.  Mammal  (In 
Press) . 

Radovsky,  R.  J.,  AND  R.  M.  Poche.  1975.  First 
report  of  an  ectoparasite  {Cryptonyssus  desul- 
torius,  Acari:  Mesostigmata:  Macronyssidae) 
associated  with  the  spotted  bat.  J.  Med.  Ent. 
(In  Press). 


THE  NEST  AND  LARVA  OF  DIPLOPLECTRON  BRUNNEIPES 
(CRESSON)    (HYMENOPTERA:    SPHECIDAE) 

Howard  E.  Evans^ 

Abstract. —  Diploplectron  brunneipes  (Cresson)  makes  a  shallow  nest  in  compact  clay-sand  con- 
taining at  least  two  cells.  It  is  provisioned  with  immature  Heteroptera.  The  larva  resembles  that  of 
Astata  in  a  general  way  but  differs  in  several  particulars. 


Wasps  of  the  genus  Diploplectron  es- 
(  ape  frequent  detection  because  of  their 
small  size  (4-7  mm)  and  secretive  behav- 
ior. For  many  years  the  genus  was  poorly 
understood,  but  in  1972  there  appeared 
two  important  papers:  Parker  presented 
a  revision  of  the  15  New  World  species, 
with  notes  on  the  biology  of  4  of  them; 
and  Kurczewski  published  a  fairly  detailed 
study  of  the  nesting  behavior  of  one 
of  these  (D.  peglowi  Krombein).  The 
present  paper  includes  brief  observations 
on  a  previously  unstudied  species,  D.  brun- 
neipes (Cresson),  as  well  as  the  first  de- 
scription of  a  larva  of  this  genus.  The 
latter  is  of  some  importance,  as  knowledge 
of  the  larvae  of  this  subfamily  (Astatinae) 
has  previously  been  based  only  on  the 
genus  Astata,  and  there  are  some  dis- 
crepancies in  the  published  information 
on  that  genus. 

These  observations  were  made  along 
the  shores  of  Terry  Lake,  just  north  of  the 
city  of  Fort  Collins,  Colorado.  D.  brun- 
neipes was  not  uncommon  during  July 
and  August  1974,  especially  in  strips  of 
bare,  flat  soil  on  top  of  a  bank  bordering 
the  beach  proper.  A  number  of  females 
were  seen  walking  and  flying  about  low 
vegetation,  apparently  hunting.  Only  one 
nest  was  found.  This  was  located  in  the 
center  of  a  bare  strip  of  rather  hard- 
packed  sandy  clay.  On  24  July  a  female 
was  seen  walking  in  a  circuitous  path 
holding  a  small  bug  in  her  mandibles. 
After  a  few  moments  she  plunged  into  a 
small,  open  hole  having  a  diameter  of  2.5 
mm  and  having  no  evidence  of  a  mound 
of  soil  around  it. 

The  female  was  captured  when  she 
emerged  from  this  hole  a  few  minutes 
later.  The  burrow  was  found  to  penetrate 
the  soil  at  about  a  60  degree  angle  with 
the  surface.  Two  cells  were  located,  at 
depths  of  5.5  and  6.5  cm,  the  two  cells 
being  4  cm  apart  and  about  9  cm  from 


the  entrance.  Each  cell  contained  6  im- 
mature bugs,  Uhleriola  floralis  (Uhler) 
(Lygaeidae)  [det.  J.  A.  Slater]  and  a 
small  larva,  one  of  which  was  reared  to 
maturity  and  is  described  below.  Both  of 
these  cells  had  been  closed  off  with  a  bar- 
rier of  sand.  Since  the  wasp  had  just 
brought  in  prey,  there  must  have  been  a 
cell  in  the  course  of  being  provisioned  (or 
prey  stored  in  the  burrow),  but  this  was 
not  found.  1  he  cells  were  broadly  ellipti- 
cal, measuring  about  3x4  mm.  The  bugs, 
all  approximately  the  same  instar,  were 
in  some  cases  on  their  backs,  in  other 
cases  on  their  sides. 

Description  of  Larva 

Length  (measured  in  usual  curved  posi- 
tion) 6.2  mm;  maximum  width  2.3  mm; 
maximum  height  (4th  abdominal  seg- 
ment) 2.4  mm.  Body  fusiform,  middle  seg- 
ments somewhat  himiped  dorsally,  4th 
abdominal  segment  more  humped  than 
any  other;  anus  terminal  and  supraanal 
and  subanal  lobes  equally  developed  (Fig. 
2).  Pleural  lobes  rather  weakly  devel- 
oped; division  of  segments  into  dorsal  an- 
nulets indistinct  posterior  to  middle  of 
body.  Integument  smooth  and  glistening, 
under  high  power  seen  to  have  a  very  few 
minute  setae,  chiefly  on  the  dorsum  and 
pleural  lobes  of  the  more  anterior  seg- 
ments, also  sparse,  minute  spinules  on 
parts  of  the  venter  and  pleura.  First  pair 
of  spiracles  slightly  larger  than  the  others; 
atrium  somewhat  pear  shaped,  sparsely 
lined  with  anastomosing  ridges;  peritreme 
distinct;  opening  into  subatrium  simple, 
unarmed  (Fig.  5). 

Head  0.9  mm  wide,  0.7  mm  high  (ex- 
clusive of  labrum)  (Fig.  1).  Head  largely 
unpigmented,  except  mandibles  and  pleu- 
rostomal  thickenings  brownish;  center  of 
front  with  paired,  small  depressions;  pari- 
etal bands  very  weak.  Antennal  orbits  el- 
liptical, papillae  only  slightly  longer  than 
wide  at  base.  Head  with  only  a  very  few 


^Department   of  Zoology  and   Entomology,   Colorado  State  University,    Fort    Collins,    Colorado    80523.      Part    of    a    study    of 
the    comparative    behavior    of    solitary    wasps,    supported    by    the  National  Science  Foundation,   grant  GB-43790. 


123 


124 


GREAT   BASIN    NATURALIST 


Vol.  35,  No.  1 


Figs.  1-5.     Diploplectron     brunneipes,     mature dible,   ventral    (posterior)    aspect;   5,   anterior  tho- 
larva:    1,  head;   2,  body,   lateral  view;   3,   labrumracic  spiracle,  high  magnification, 
(left  side)    and  epipharynx   (right  side);  4,  man- 


small  setae.  Labrum  0.33  mm  wide,  with  a 
strong  V-shaped  median  emargination; 
surface  with  about  20  small  setae,  also 
with  a  fe  w^  small  marginal  sensilla, 
mainly  laterally;  epipharynx  with  6 
strong  sensillae  surrounding  the  emargin- 
ation and  some  weaker  ones  basal  of  these, 
otherwise  clothed  with  very  fine  spinules 
except  medially  (Fig.  3).  Mandibles  stout, 
with  5  strong  teeth,  one  of  them  ventral 
of  the  most  apical  tooth  (Fig.  4);  upper 
surface  with  a  single  minute  seta.  Maxil- 


lae short,  directed  mesad,  inner  surface 
roughened  but  not  distinctly  spinulose; 
galeae  very  much  more  slender  than  ])al- 
pi.  Hypopharynx  inconspicuous  and  evi- 
dently not  spinulose.  I^abium  without  se- 
tae or  spinules,  spinnerets  blunt,  consider- 
ably exceeding  the  palpi. 

Discussion 

The  nest  of  D.   hrumieipes  closely  re- 
sembles that  of  D.  peglowi  Krombein,  as 


March  1975 


EVANS:    WASP   NEST   AND  LARVA 


125 


described  by  Parker  (1972)  and  Kurczew- 
ski  (1972),  although  evidently  in  flatter 
and  more  compact  soil  than  that  species 
usually  occupies.  The  angle  of  the  bur- 
row, depth  and  size  of  the  cells,  and  open 
nest  entrance  are  similar  in  the  two  spe- 
cies, as  are  the  type  of  prey  and  manner 
of  prey  carriage. 

The  larva  is  basically  similar  in  struc- 
ture to  that  of  Astata  as  described  by 
Evans  (1958),  although  differing  in  some 
details  from  descriptions  provided  by  earli- 
er workers.  The  paired  spinnerets, 
humped  fourth  abdominal  segment,  ter- 
minal anus,  short  antennal  papillae,  stout 
mandibles,  mesally  directed  maxillae, 
and  smooth  integimient  together  clearly 
define  the  Astatinae  as  distinct  from  other 
subfamilies.  One  other  feature  that  I 
used  to  define  the  subfamily,  the  pres- 
ence of  numerous  setae  on  the  mandibles, 
does  not  hold  up,  and  it  represents  an  im- 
portant difference  between  the  larvae  of 
Astata  and  Diploplectron.  Other  differ- 
ences include  the  5-toothed  mandibles  of 
Diploplectron,  the  sparser  head  setae,  and 
the  lack  of  conspicuous  labral  sensory 
cones. 

The  larva  of  Astata  feeds  in  an  inverted 
position  in  the  cell,  the  egg  having  been 


laid  on  the  ventral  side  of  a  bug  placed 
with  its  dorsum  upward  in  the  bottom  of 
the  cell  (Evans,  1957).  I  had  assumed 
that  the  humped  4th  abdominal  segment 
represented  a  pseudopod  that  assisted  the 
larva  in  feeding  in  this  unusual  position. 
However,  the  species  of  Diploplectron 
place  the  bugs  in  the  cell  in  various  posi- 
tions, and  according  to  Kurczewski  (1972) 
the  bug  bearing  the  egg  is  found  either 
on  its  side  or  with  its  venter  upward.  Of 
course  the  middorsal  hump  may  still  serve 
to  assist  the  larva  in  pushing  itself  about 
from  prey  to  prey,  though  it  seems  less 
suitably  adapted  for  this  type  of  feeding. 

Literature  Cited 

Evans,  H.  E.  1957.  Ethological  studies  on  dig- 
ger wasps  of  the  genus  Astata  (Hymenop- 
tera,  Sphecidae).  J.  N.  Y.  Ent.  Soc.  65:159- 
185. 

.     1958.     Studies   on   the   larvae   of   digger 

wasps  (Hymenoptera,  Sphecidae).  Part  IV: 
Astatinae,  Larrinae.  and  Pemphredoninae. 
Trans.   Ainer.   Ent.   Soc.   84:109-139. 

Kurczewski,  F.  E.  1972.  Obsei-vations  on  the 
nesting  behavior  of  Diploplectron  peglowi 
Krombein.  Proc.  Ent.  Soc.  Washington  74: 
385-397. 

Parker,  F.  D.  1972.  On  the  subfamily  Astat- 
inae, Part  VII.  The  genus  Diploplectron  Fox 
(Hymenoptera:  Sphecidae).  Arm.  Ent.  Soc. 
Amer.   65:1192-1203. 


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TABLE  OF  CONTENTS 

Evolution  of  the   sceloporine  lizards    (Iguanidae).   Kenneth   R.   Larsen  and 

Wilmer    W.    Tanner    1 

New    synonymy   and   new   species   of   American    bark    beetles    (Coleoptera: 

Scolytidae).    Stephen  L.  Wood  21 

Genetics,  environment,  and  subspecies  differences:   the  case  of  Polites  sabu- 

leti    (Lepidoptera:    Hesperiidae) .    Arthur  M.  Shapiro  33 

Life    history    and    ecology    of   Megarcys  signata    (Plecoptera:    Perlodidae), 

Mill  Creek,  Wasatch  Mountains,  Utah.    Mary  R.  Cather  and  Arden  R. 

Gaufin   39 

Records  of  stoneflies   (Plecoptera)   from  Nevada.    Mary  R.  Cather,  Bill  P. 

Stark,    Arden    R.    Gaufin    49 

Growth    of    Plecoptera    (stonefly)    nymphs    at    constant,    abnormally    high 

temperatures.    Joseph   M.   Branham,  Arden  R.  Gaufin,  and  Robbin  L. 

Traver   51 

Water  balance  and  fluid  consumption  in  the  southern  grasshopper  mouse, 

Onychomys  torridus.  Vernon  C.  Bleich  and  Orlando  A.  Schwartz  ....  62 
A    systematic    study    of    Coenia    and    Paracoenia    (Diptera:    Ephydridae). 

Wayne    N.     Mathis    65 

Environmental  factors  in  relation  to  the  salt  content  of  Salicornia  pacifica 

var.  utahensis.    D.  J.  Hansen  and  D.  J.  Weber 86 

New   records  of  stoneflies    (Plecoptera)    from   New  Mexico.    Bill  P.   Stark, 

Theodore  A.  Wolff,  and  Arden  R.  Gaufin  97 

The    authorship   and   date   of   publication   of   Siren   intermedia    (Amphibia: 

Caudata).  Hobart  M.  Smith,  Rosella.B.  Smith,  and  H.  Lewis  Sawin  ....  100 
New  mites  from  the  Yampa  Valley  (Acarina:  Cryptostigmata:  Oribatulidae, 

Passalizetidae) .     Harold   G.   Higgins  and   Tyler  A.   Woolley   103 

The   identity   of   Bocourt's   lizard   Eumeces  capita   1879.   Hobart  M.   Smith, 

Rozella  B.  Smith,  and  Jean  Guibe  109 

Studies   in  nearctic  desert  sand  dune  Orthoutera.    Part  XV.    Eremography 

of  Spaniacris  W\\\v  biological  notes.    Ernest  R.  Tinkham  113 

Roosting   behavior  of   male   Euderma   maculatum  from   Utah.    Richard   M. 

Poche  and  George  A.  Ruffner  121 

The   nest   and   larva   of  Diploplectron  brunneipes   (Cresson)    Hymenoptera: 

Sphecidae).    Howard  E.   Evans   123 


HE  GREAT  BASIN  NATURALIST 


L|ne35No.2  June  30,1975 

v5-  G    ^6.  ^ 


Brigham  Young  Universitv 


LJBRARY 

OCT  6    19/iD 

HARVARD 

u<\jjve;f^sjvy 


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,  Zoology;  Stanley  L. 

Welsh,  Botany;  Clayton  M.  White,  Zoology. 
Ex  Officio  Editorial  Board  Members.  A.  Lester  Allen,  dean.  College  of  Biological  and 

Agricultural  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  continuously  published  from  one  to  four  times  a  year  since  then  by  Brigham 
Young  University,  Provo,  Utah.  In  general,  only  original,  previously  unpublished 
manuscripts  pertaining  to  the  biological  natural  history  of  the  Great  Basin  and  western 
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Manuscripts.  All  manuscripts  and  other  copy  for  the  Great  Basin  Naturalist 
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The  Great  Basin  Naturalist 

Published  at  Provo,  Utah,  by 
Brighaini  Young  University 


Volume  35 


June  30,  1975 


No.  2 


A  REVISION  OF  THE  PHACELIA  CRENULATAE  GROUP 
{IIYDROPHYLLACRAE)  FOR  NORTH  AMERICA 

N.  Duane  Atwood^ 

Abstract. —  This  taxononiic  monograph  of  tlie  Phacelia  crenulatae  gioup  for  North  America 
recognizes  35  species  and  7  varieties.  A  hrief  discussion  of  the  history,  general  morphology,  phyl- 
ogeny.  and  cytology  is  given.  All  entities  are  separated  by  a  comprehensive  key.  followed  by  a  list 
of  types,  synonyms,  dcsci-iptions.  and  general  habitat.  Distribution  maps  and  illustrations  are  in- 
cluded. One  vaiiety  is  desciibed  as  new.  The  body  of  this  wotk  is  based  on  herbarium  specimens  and 
extensive  field  observations  and  collections  made  throughout    nuich    of    North    America. 


Introduction 

The  Crenulatae  group  of  Phacelia  be- 
longs to  the  subgenus  Phaceha,  section 
Phacelia,  and  may  be  distinguished  from 
other  members  of  the  section  by  the  four- 
seeded  capsule  and  excavated  ventral  sur- 
face of  the  seeds.  Many  of  the  species 
are  viscid  and  ill-scented  desert  plants 
confined  mostly  to  western  North  Amer- 
ica and  Mexico.  The  remaining  species 
occur  in  South  America  and  in  the  mid- 
western  and  the  west  central  parts  of  the 
United  States. 

The  lack  of  phenologic,  edaphic,  morph- 
ologic, and  distributional  data  plus  the 
description  of  additional  species  since  the 
monograph  by  Voss  (1937)  have  neces- 
sitated a  thorough  study  of  the  group. 
Very  little  was  known  about  the  species 
occurrmg  in  Texas,  New  Mexico,  Arizona, 
Utah,  and  California  or  species  in  Mexico 
and  South  America.  Previous  revisions 
were  based  entirely  on  herbarium  spec- 
imens, and,  possibly  because  of  this  fact, 
numerous  errors  and  misconceptions  ap- 
peared in  them.  Therefore  it  'became 
necessary  to  conduct  extensive  field  work 
wherein  most  of  the  entities  were  ex- 
amined  in  living  condition. 

Phacelia  integrifolia  Torr.  was  the  first 
species  of  the  Crenulatae  group  to  be  de- 
scribed.   It  was  collected  by  James  in  June 


1820  but  was  not  described  until  1826.  The 
next  taxon  was  described  in  1835  when 
Hooker  named  P.  congesta  from  plants 
collected  by  Drummond  in  the  vicinity  of 
Galveston  Bay,  Galveston  County,  Tex- 
as. In  1848  Nuttall  described  P.  gland- 
ulosa  from  specimens  collected  the  same 
year,  "about  Ham's  Fork  Colorado  of  the 
West."  Hooker  (1851)  assigned  P.  gland- 
iilosa  to  the  genus  Eutoca.  Since  1849, 
41  nominate  species  have  been  described. 
Of  the  78  names  proposed  in  this  group, 
fewer  than  half  of  them  are  recognized 
as  valid   taxa  in  the  present  study. 

Materials  and  Methods 

Research  materials  for  this  study  have 
come  from  two  major  sources:  a  large 
quantity  of  plants  borrowed  from  herbaria 
in  Germany,  Mexico,  and  the  United 
States,  and  from  field  studies  made  during 
the  growing  season  of  1968  in  Utah,  Ar- 
izona, and  Wyoming;  in  1969  in  Arizo- 
na, California,  Colorado,  Utah,  and  Wy- 
oming; and  during  1970-1971  in  Texas, 
Mexico,  Arizona,  New  Mexico,  Califor- 
nia, and  Utah. 

Measurements  of  such  large  plant  parts 
as  stems,  leaves,  and  inflorescences  were 
made  with  a  metric  ruler.  Small  struc- 
tures such  as  calyces,  flowers,  capsules, 
and  seeds  were  measured  with  the  aid  of 


^Research    associate,    Department    of    Botany,    Brighani    Young    University,    Provo,    Utah    84602. 


127 


128 


GREAT   BASIN    NATURALIST 


Vol.  35,  No.  2 


an  ocular  micrometer  fitted  to  a  stereo- 
scopic microscope.  At  the  end  of  each  de- 
scription a  list  of  specimens  examined  in 
the  study  of  each  entity  is  given.  The 
number  following  the  description  indicates 
the  total  number  of  specimens  seen,  while 
the  number  in  ])arentheses  indicates  the 
number  of  collections  made  by  the  author. 
The  standard  abbreviations  of  herbaria, 
with  a  few  exceptions,  are  those  of  Lan- 
jouw  and  Stafleu  (1964).  These  indicate 
the  herbaria  from  which  specimens  were 
examined.  The  type  specimens  examined 
were  photographed  and  the  photographs 
deposited  in  the  herbarium  at  Brigham 
Young  University;  they  are  indicated  by 
an  exclamation  mark  following  the  her- 
barium symbol  designation  in  the  list  of 
synonyms. 

B  Botanisclicr    Garten    and    Botanisches 

Museum.  Berlin-Dahlem.  Germany 

BRY  Brigham    Young     University.    Provo, 

Utah 

CAS  California  Academy  of  Sciences.  San 

Francisco.  California 

GH  Gray   Herbarium,    Harvard    Universi- 

ty. Cambridge.  Massachusetts 

JEPS  Jepson  Herbarium.  University  of  Cal- 

ifornia. Berkeley,  California 

MEXU  Herbario  Nacional  del  Institute  Bio- 
logia.  Universidad  Nacional  de  Mex- 
ico 

NY  New    York    Botanical    Garden.    New 

York.   New  York 

POM  Pomona    College    Herbarium.    Clare- 

mont.   California 

RM  Rock}'     Mountain     Herbarium,     Lar- 

amie.  Wyoming 

RSA  Rancho    Santa    Ana    Botanic    Garden. 

Claremont,  California 

UC  University    of    California.     Berkeley, 

California 

UNM  University  of  New  Me.xico.  Albu- 
quercjue.   New   Me.xico 

US  United      States      National      Museum. 

Washington,   D.C. 

UT  University   of   Utah    Herbarium.    Salt 

Lake  City,  Utah 

UTC  Intermountain      Herbarium.      Logan. 

Utah 

WTS  West    Texas    State    LTniversity.    Can- 

yon.  Texas 

Seeds  provide  important  distinguish- 
ing features,  and,  therefore,  the  Electron 
Scanning  Microscope  facilities  at  Brigham 
Young  University  were  used  to  photo- 
graph them.  They  were  air-dried  and 
mounted  on  polished  brass  sj)ecimen  stubs 
with  Elmer's  glue  which  had  been  di- 
luted one  part  glue  to  three  parts  water. 


The  specimens  were  coated  with  gold 
(200-300  angstroms)  using  a  rotating 
stage  and  examined  with  a  Hitachi  SSM-2 
microscope.  All  specimens  were  exam- 
ined with  the  microscope  beam  voltage 
set  at  20  kv. 

General  Morphology 

Members  of  the  Crenulatae  group  are 
annual,  biennial,  or  perennial  herbaceous 
plants.  They  possess  a  simple  taproot, 
which  varies  in  size  depending  on  the 
species  and  even  on  the  individual  plant. 
The  stem  pattern  varies  considerably  be- 
tween species  but  basically  consists  of  an 
axis  which  is  usually  foliate  and  bears 
several  to  many  scorpioid  cymes.  The 
majority  of  species  are  annuals  which 
complete  their  life  cycle  in  two  to  three 
months.  However,  the  seeds  of  some  spe- 
cies, such  as  P.  corrugata,  generally  germ- 
inate in  the  fall  and  produce  a  rosette  of 
leaves.  This  rosette  is  small  at  first  but 
continues  to  grow  during  the  warmer 
periods  of  the  winter  months.  Then  in 
the  spring  it  produces  a  flowering  shoot. 
These  are  winter  annuals.  The  biennial 
species,  as  well  as  some  annuals,  gener- 
ally have  thick  stems  and  produce  a  basal 
rosette  of  leaves.  The  perennial  species 
have  a  thickened  woody  caudex  which 
produces  one  to  several  herbaceous 
branched  or  simple  stems.  These  are 
terminated  by  a  series  of  scorpioid  cymes. 

Leaves 

The  leaves  vary  from  simple  to  bipin- 
nately  compound,  with  a  series  of  inter- 
mediate types.  About  half  the  species  have 
simple  leaves  with  the  margins  crenate 
to  dentate,  irregularly  serrate,  or  incised. 
Those  taxa  having  compound  leaves  are 
usually  fjuite  distinct  from  the  foregoing, 
but  A'ariations  between  the  two  types  exist. 
The  margins  of  some  leaves,  such  as  those 
of  P.  cofistancei,  P.  intergrifolia,  and  P. 
ircls/iii,  are  often  revolute.  A  basal  rosette 
is  usually  present  in  biennial,  perennial 
and  some  robust  annual  species.  The 
basal  and  lower  cauline  leaves  are  typ- 
ically larger  and  longer-petiolate  than  the 
gradually  reduced  upper  cauline  leaves. 
Leaf  pubescence  varies  depending  on  the 
spec  i(\s.  but  the  leaves  possess  either  one 
or,  nior(»  often,  a  combination  of  pub- 
escence types.  In  general,  leaf  characters 
have  not  been   used  to  delineate  species, 


197: 


ATWOOD:   phaci:lia  crenui.atae  group 


129 


since  other  less  variable  and  more  im- 
portant taxonomic  characters  are  avail- 
able. 

Inflorescence 

Heckard  (1960)  indicated  that  agree- 
ment is  generally  lacking  as  to  the  exact 
terminology  used  to  describe  a  branched 
system  of  scorpioid  cymes.  HoweA'er,  the 
inflorescences  are  generally  best  described 
as  being  composed  of  compound  scor])ioid 
cymes.  In  some  species,  such  as  P.  coeru- 
Ica  and  P.  hombycina,  the  inflorescences 
appear  to  be  racemose.  The  flower  cluster 
is  actually  a  false  raceme  because  the 
flowers  are  all  borne  on  one  side  of  the 
peduncle.  In  most  species  the  inflores- 
cence is  open  but  with  terminally  con- 
gested clusters.  In  P.  pa/mcri.  P.  utahcn- 
sis,  and  P.  vossii  the  inflorescence  is  con- 
gested into  a  spicate  thyrsus.  The  inflores- 
cences are  generally  more  glandular  than 
the  stems  and  leaves.  The  individual 
cymes  of  P.  integrifolia  elongate  to  as 
much  as  2.1   dm  in  JFruit. 

Flower 

Corolla:  The  flowers  are  crowded  along 
a  coiled  peduncle  that  uncoils  as  flower- 
ing advances.  The  shape,  size,  and  color 
of  the  corollas  are  taxonomically  im- 
portant. The  corollas  are  funnelform  to 
rotate  or  campanulate  in  shape,  and  blue, 
purple,  violet  or  lavender  in  color.  One 
series  of  taxa  has  white  to  lavender  tubu- 
lar corollas.  The  corolla  lobes  are  nor- 
mally entire  or,  at  the  most,  merely  cre- 
nulate  and  finely  pubescent.  However, 
in  P.  neomexicana  and  its  relatives  the 
lobes  are  either  fimbrate  or  denticulate. 
The  pedicels  are  commonly  less  than  3 
mm  long  with  exceptions  in  P.  pedicellata 
and  P.  scariosa,  in  which  the  pedicels  are 
6  and  8  mm  long  respectively. 

Corolla  scales:  Corolla  scales  are  present 
in  all  species  of  this  group  and  occur  in 
pairs  at  the  base  of  each  filament.  The 
variation  in  size,  shape,  and  attachment 
of  the  scales  offers  some  variation,  but  as 
a  whole  it  is  not  as  useful  in  delineating 
species  as  are  other  characters. 

Androecium:  The  filaments  are  attached 
at  the  base  of  the  corolla  tube  and  vary  in 
length  depending  on  the  taxa  involved. 
Even  in  individual  plants  filament  length 
varies  considerably.  The  filaments  are 
glabrous  in  all  species.    The  anthers  are 


dorsifixed,  ca.  1  mm  long  and  0.5  mm 
broad,  and  ojjen  their  full  length  by  two 
longitudinal  slits.  The  ])ollen  has  not  been 
studied  systematically.  The  stamens  as 
well  as  the  style  in  /-•.  coerulca.  P.  den- 
licLilata.  and  P.  anelsonii  are  included 
within  the  corolla,  or  nearly  so.  This  fea- 
ture has  been  given  taxonomic  im])ortance, 
but  it  varies  in  some  ])opulations,  as  noted 
in  P.  coerulea  and  /-*.  denticulata.  In 
these  instances,  the  stamens  are  barely  ex- 
serted  from  the  tube.  Some  confusion  may 
occur  in  keying  out  collections  of  plants 
that  are  in  early  anthesis,  since  the  sta- 
mens of  most  species  are  folded  in  the  bud 
and  become  exserted  only  when  the  flower 
is  fully  opened. 

Gynoecium:  The  gynoecium  consists  of 
an  ovoid  to  subglobose  or  oblong,  usually 
puberulent  and  conmionly  glandular 
ovary.  The  jiersistent  bifid  style  is  term- 
inated by  small  stigmatic  areas.  The  bi- 
furcation of  the  style  varies  from  two- 
thirds  to  three-fourths  of  its  length,  with 
the  lower  undi^•ided  ]:)ortion  being  pub- 
escent. The  ovary  is  1 -celled  or  incom- 
pletely 2-celled  b>'  union  of  the  placentae. 
Four  ovules  are  commonly  produced; 
however,  sometimes  one  is  reduced  in 
size  or,  less  frequently,  lacking  altogether. 
This  condition  ajipears  only  si)oradically 
and  is  probably  influenced  by  environ- 
mental and  nutritional  factors. 

Calyx:  The  calyx  is  five-parted  to  the 
base,  or  nearly  so.  The  lobes  vary  in  size 
and  shape  from  species  to  species.  There 
is  consistent  variation  in  flowering  and 
fruiting  calyces  with  those  in  fruit  being 
larger  and  sometimes  scarious,  as  in  P. 
scariosa. 

Seeds 

The  seeds  are  geminate,  elliptic  to  ob- 
long and  ovoid,  and  generally  cymbiform 
in  shape.  Size,  shape,  and  surface  mark- 
ings are  diagnostically  important.  The 
seeds  are  unique  in  having  the  ventral 
surface  excavated  on  one  or  both  sides  of 
a  prominent  ridge.  However,  the  seeds 
of  P.  bakeri  have  the  dorsal  surface  flat, 
with  only  a  faint  longitudinal  groove 
down  the  center.  On  the  ventral  surface, 
the  raphe  is  elevated  above  the  normally 
excavated  portions,  thus  giving  the  seed 
a  triangular  shape  in  cross  section.  In 
other  taxa,  the  dorsal  surface  may  be 
transversely    ridged,    as    in    P.    arizonica, 


130 


GREAT  BASIN  NATURALIST 


Vol.  35,  No.  2 


P.  palmeri,  and  P.  popei,  and  reticulate 
to  scabrous  in  P.  congesta  and  P.  rupestris. 
P.  howelliana,  P.  serrata,  and  P.  utahen- 
sis  haAe  the  dorsal  surface  smooth  and 
shiny  ^^^th  faint  alveolations,  while  the 
seeds  of  P.  pediccllata  are  tuberulate.  The 
remaining  species  are  alveolate  (pitted). 
The  ridge  is  corrugated  on  one  side  in 
over  half  the  taxa,  while  the  remainder 
lack  corrugations.  The  seed  margins  can 
be  entire  as  in  P.  alba,  P.  denticulata, 
and  others,  or  corrugated  along  part  or 
all  of  the  marginal  edge.  Corrugated  mar- 
gins are  well  represented  in  seeds  of  P. 
bombycina,  P.  coerulea,  and  P.  corrugata. 

Color  variations  occur,  but  brown  pre- 
dominates as  in  P.  neomexicana  and  P. 
pediccllata.  In  such  taxa  as  P.  constancei, 
P.  pallida,  and  P.  palmeri  the  seeds  are 
black,  while  in  P.  bombycina,  P.  coerulea, 
and  P.  formosula,  they  are  dark  brown. 
Reddish  or  reddish  browTi  seeds  are  typ- 
ical of  P.  glandulosa,  P.  rafaelensis,  and 
P.  utahensis. 

The  smallest  seeds  occur  in  P.  coulteri 
(1.6  mm  long),  and  the  largest  known  are 
in  P.  denticulata  (4  mm  long) .  The  light- 
colored,  glutinous  thickened  band  spoken 
of  by  Voss  (1937)  is  a  feature  that  de- 
velops during  the  ontogeny  of  the  seeds. 
When  immature,  the  seeds  are  either  dark 
and  turn  light  in  color  through  a  mottling 
pattern,  or  they  are  light  and  become 
dark  when  mature.  The  descriptions  of 
seeds  in  this  treatment  are  based  on  ma- 
ture examples.  They  are  considered  the 
most  important  single  feature  in  delin- 
eating taxa,  and  on  the  basis  of  the  size, 
shape,  and  surface  characters,  several  dis- 
tinct groups  can  be  arranged  in  an  appar- 
ent phylogonetic  order.  These  groupings 
are  supported  by  other  morphological  fea- 
tures as  well,  namel}'  leaf  shape,  pub- 
escence, corolla  shape  and  color,  and  du- 
ration of  the  plant.  The  branching  pat- 
tern of  the  stem  and  inflorescence,  the 
type  of  calyx  segments,  and  the  stamens 
and  style,  whether  included  or  exserted, 
are  also  useful  features. 

Vesture 

Consi(lerabl(>  confusion  exists  as  to  the 
terminology  used  in  describing  the  ves- 
ture of  plants.  Ihe  terms  employed  here- 
in are  defined  in  Ai)pen(Hx  II  to  lend  uni- 
formity to  their  interpretation.  The  pub- 
escence often  consists  of  two  or  more  types 


of  intermixed  hairs.  There  are  two  main 
types  of  trichomes,  each  exhibiting  vari- 
ation in  size:  (1)  simple,  unicellular  tri- 
chomes, which  vary  in  length  and  rigidity 
and  may  be  erect,  straight,  or  appressed; 
and  (2)  a  stipitate-glandular  type,  which 
is  usually  multicellular.  The  stalk  in  the 
stipitate-glandular  type  varies  in  length 
and  in  number  of  cells.  Sometimes  the 
gland  is  sessile,  or  nearly  so,  and  the  stalk 
is  often  flattened. 

Phylogeny 

Constance  (1963)  indicates  that  the 
family  appears  to  be  a  collection  of  mor- 
phological and  geographical  odds  and 
ends,  held  together  by  floral  and  capsular 
features.  He  states,  "I  am  not  prepared  to 
offer  a  complete  system  for  Phacelia." 
However,  Constance  (1963)  appears  to 
have  arrived  at  the  most  natural  grouping 
of  the  genus  Phacclia  by  recognizing  three 
subgenera,  Cosmanthus,  Howellanthus, 
and  Phacelia.  The  latter  is  the  largest  and 
most  complex  of  the  three  and  has  been 
subdivided  by  Constance  (I.e.)  into  the 
following  species-groups:  Crenulatae,  Eu- 
glypta,  Gymnobythus,  Miltitzia,  Pulchel- 
lae,  Tanacetifoliae,  and  Whitlavia.  The 
Crenulatae  group,  revised  by  Voss  (1937), 
was  the  most  complete  study  of  the  group 
])rior  to  the  ])resent  work.  Gillett  (1960b) 
indicates.  "The  current  infrageneric  clas- 
sification of  PJuwclia  is  generally  consid- 
ered to  be  inadequate  .  .  .  and  that  con- 
siderably more  evidence  must  be  accumu- 
lated before  the  various  species  groups  can 
be  accorded  classification  that  properly  re- 
lates them  to  each  other."  The  author 
agrees  that  natural  generic  and  infra- 
generic  relationships  cannot  be  ])roposed 
imtil  additional  morphological,  distribu- 
tional, and  cytological  data  have  been  ac- 
cumulated. Howe^■er,  he  is  ]irepared  to 
offer  a  tentative  phyogenetic  sunmiary  of 
relationships  within  the  Crenulatae  grouj). 
These  data  are  subject  to  change  as  ad- 
ditional research  may  warrant. 

Those  species  occurring  from  Mexico 
to  South  America  present  a  problem  in 
th(>  formulation  of  a  complete  phylogen- 
etic  scheme.  These  southern  taxa  a])pear 
to  !)(>  the  most  jirimitiAO  and  are  certainly 
file  least  understood  of  all  the  Cremdatae 
grou]).  Most  are  known  only  from  the 
t^■pe  collections.  Il  wo"uld  seem  likely  that 
the    Cremdatae    group    has    been    derived 


June  1975 


AT^VOOD:     I'HACELIA    CRENULATAF.   CiROUP 


131 


congesta 


crenulata 


welshi 


neomexicana 


Fig.    1     A    phylogenetic    arrangement    of   the 
complexes  in  the  Crenulatae  group. 

from  some  form  of  Phacelia,  past  or  pres- 
ent, somewhere  in  Mexico  or  South  Amer- 
ica. The  modern  sj^ecies  suggest  several 
avenues  of  migration  from  Mexico,  which 
have  contributed  to  the  ])resent  diversity 
in  morphology  and  distribution.  Further- 
more, the  morphological,  cytological,  and 
distributional  relationships  of  the  subgen- 
eric  and  sectional  groups  of  Phacelia  sug- 
gest either  a  polyphyletic  origin  or,  if  a 
monophyletic  one,  then  a  derivation  pos- 
sessing several  major  lines  of  development. 
Those  main  lines  of  development  occur- 
ring within  the  Crenulatae  group  are  out- 
lined in  Figures  1-7.  The  species  are 
grouped  together  and  arranged  on  the 
basis  of  similar  morphological  features 
and  distribution.  The  following  discussion 
is  given  to  indicate  which  characters  are 
considered  to  be  advanced  or  primitive 
in  this  group. 

Seeds  provide  the  most  important  char- 
acters in  differentiating  entities.  The  most 
primitive  species,  which  occur  in  Mexico, 
all  have  small  seeds,  which  suggests  that 
large  seeds  are  probably  a  derived  feature. 
This  character  seems  to  follow  a  south-to- 
north  trend  with  the  largest  seeds  occur- 
ring to  the  north.  There  have  been  several 
avenues  of  specialization  with  the  primi- 
tive seeds  having  more  surface  markings 
and    being    thicker    and    narrower.     The 


seeds  of  most  taxa,  except  in  P.  bakeri  and 
P.  argillacea,  uniformly  have  the  ventral 
surface  excavated  on  both  sides  of  a  prom- 
inent ridge.  These  latter  entities  have  the 
raphe  elevated  above  the  usually  exca- 
vated portions  and  would  appear  to  have 
diverged  from  the  more  typical  form. 
Seeds  with  a  corrugated  ridge  appear  to  be 
primitive,  and  those  with  pitted  (alveo- 
late) and  entire  margins  appear  to  be  ad- 
vanced. Light  brown  seeds  are  apparently 
primitive,  and  dark  brown,  black,  and  reel- 
dish  types  are  apparently  derived.  Retic- 
ulate, transversely  ridged,  and  smooth-sur- 
faced seeds  are  also  probably  derived. 

The  corolla  has  developed  along  three 
basic  lines.  Primitive  plants  are  those 
having  blue  to  purple  colored  campan- 
ulate  corollas  and  exserted  stamens  and 
styles.  However,  some  of  the  less  advanced 
species  have  small,  pale,  campanulate 
corollas  with  included  stamens  and  styles. 
Thirdly,  pale  to  white  tubular  corollas  are 
present  in  the  more  advanced  entities. 
These  advanced  forms  have  less  attractive 
flowers  and  long  exserted  stamens  and 
styles.  The  corolla  lobes  have  developed 
along  two  major  lines,  with  the  P.  neo- 
mexicana complex  having  denticulate  or 
erose  margins  and  the  remainder  having 
entire    margins.     The    former    feature    is 

bakeri 
glandulosa  ^  argillacea 


formosub 


denticulata 


coulter 


Fig.  2.     A   phylogenetic   arrangement  of  the 
species  in  the  neomexicana  complex. 


132 


GREAT  BASIN  NATURALIST 


Vol.  35,  No.  2 


splendens 


rafaelensis 


utahensis 


Fig.  3.     A   phylogenetic   arrangement  of  the 
species  in  the  welshii  complex. 

probably  advanced,  entire  margins  being 
primitive. 

The  scarious  calyx  segments  of  P.  scar- 
iosa  and  P.  pedicellata  seem  to  be  an  ad- 
vanced feature.  This  is  supported  by  the 
fact  that  the  calyx  lobes  are  persistent  in 
fruit  and  probably  aid  in  dispersal.  Small, 
narrow  calyx  segments  are  considered  to 
be  primitive. 


palMeri 


The  spicate  thyrsus  type  of  inflores- 
cence of  the  perennial,  biennial,  and  some 
robust  annual  species  is  apparentl}"  ad- 
vanced, while  the  variously  branched  sys- 
tems developed  in  most  annual  and  some 
biennial  taxa  appear  to  be  primitive. 

The  primitive  taxa  do  not  follow  the 
generalization  that  the  perennial  habit  is 
more  primiti^  e  than  the  biennial  or  an- 
nual type.  The  majority  of  entities  are  ro- 
bust annuals,  probably  an  inherited  fea- 
ture; biennial  and  perennial  types  ap- 
pear to  be  derived.    The  primitive  taxa 


integrifolia 


howel  liana 

onelsonii 


intePMedia 


coerulea    boMbycina 


constcncei  robusta 


pallida 


Fig.  4.     A   phylogenetic   arrangement   of   the 
species  in  the  palmeri  complex. 


Fig.  5.     A   phylogenetic   arrangement  of  the 
species  in  the  crenulata  complex. 

possess  an  erect,  usually  branched,  stout 
stem  with  a  compound  scorpioid  inflores- 
cence. Some  advanced  biennial  and  pe- 
reimial  entities  have  become  specialized 
in  the  development  of  a  spicate  thyrsus  in- 
florescence. 

Entire  or  subentire  leaves  are  appar- 
ently derived  from  compound  leaves.  The 
narrowly  revolute  type  exhibited  by 
P.  constancei  is  considered  to  be  a  special- 
ization, while  the  basal  rosette  common  in 
the  biennials  and  robust  annuals  appears 
to  be  a  feature  that  has  been  retained 
during  the  phylogenetic  development  of 
leaves. 

Members  of  the  Crenulatae  group  are 
probably  monophyletic  and  have  devel- 
oped  along   six   major   lines.     These   are 


June  1975 


AT  WOOD:     PHACELIA    CRENULATAE   GROUP 


133 


rupestris 


congesta 


\7 


infundibuliforMis 


Fig.  6.     A   phylogeiietic   arrangement   of  the 
species  in  the  congesta  complex. 

treated  as  complexes  but  are  not  accorded 
taxonomic  status. 

The  hypothetical  ancestor (s)  of  the 
Crenulatae  group  were  apparently  robust, 
densely  glandular  annuals  with  a 
branched  stem,  compound  leaves  and  in- 
florescences, nonscarious  sepals,  and 
broadly  campanulate  blue  or  purple  cor- 
ollas. The  seeds  were  light  brown,  small, 
excavated  on  both  sides  of  the  corrugated 
ridge,  with  thick  entire  margins,  cymbi- 
form,  and  elliptic  to  oblong  in  shape.  Bi- 
ennial and  peremiial  types  developed  later 
in  the  phylogeny  of  the  group.  The  prim- 
itive members  of  the  neoniexicana  complex 
are  apparenth"  the  most  primitive  and  are 
considered  to  be  closest  to  the  ancestral 
forms.  The  palmeri  complex  arose  some- 
what later  and  extended  more  to  the  west 
of  the  neomexicana  complex  in  its  migra- 
tion northward.  The  congesta  complex 
had  its  origin  somewhere  in  north  central 
Mexico  and  ])ossibly  arose  from  the  multi- 
ovulate  P.  infundibuliformis  or  some  sim- 
ilar form.  The  crenulatae  and  scariosa 
complexes  probably  arose  from  taxa  now 
extant  in  western  Mexico.  These  six 
complexes  are  discussed  and  outlined  in 
Figures  1-7. 


pattern  of  this  complex  has  been  north- 
ward out  of  Mexico  through  New  Mexico 
to  Wyoming  and  Montana.  P.  popei  and 
P.  arizonica  have  developed  from  a  com- 
mon ancestor,  as  indicated  by  their  sim- 
ilarity in  seed,  pubescence,  and  vegetative 
features.  The  same  is  true  of  P.  fonnosula^ 
P.  glandulnsa,  P.  bakeri,  and  P.  argillaceci; 
however,  the  latter  two  have  become  spe- 
cialized in  the  development  of  noncorru- 
gated  seeds.  This  is  the  only  specializa- 
tion away  from  the  typically  excavated 
seed  type  present  in  the  rest  of  the  Cren- 
ulatae group. 

Welshii  complex 

This  complex  is  characterized  by  the 
large  reddish  seeds,  showy  corollas,  and 
generally  long  exserted  stamens  and  style. 
All  taxa  are  narrowly  restricted  endemics 
occurring  in  Utah,  Arizona,  and  western 
Colorado.  P.  welshii  is  considered  to  be 
the  most  primitive  on  the  basis  of  its 
smaller,  somewhat  brownish  seeds  and 
branched  habit.  P.  utahensis  and  P.  splen- 
dens  probably  had  a  common  ancestor  but 
have  adapted  to  different  edaphic  situa- 
tions and  have  therefore  been  isolated  and 
selected  out;  judged  on  its  robust  bran- 
ching habit  and  glandular  pubescence, 
the  former  is  probably  more  primitive. 
P.  rafaelensis  is  related  to  P.  utahensis 
and  may  have  been  derived  from  it. 
P.  serrata  is  the  most  advanced  species  as 
characterized  by  its  smaller,  lighter-col- 
ored corollas,  and  shortly  exserted  sta- 
mens and  styles. 

Palmeri  complex 

P.  vossii  and  P.  pallida  are  the  most 
primitive  and,  along  with  P.  robusta,  are 
restricted  to  the  south  central  part  of  the 
U.S.  and  adjacent  Mexico.  The  remaining 
two  species,  occurring  in  Utah,  Arizona, 


Neomexicana  complex 

This  complex  is  characterized  by  non- 
corrugated  seeds,  densely  glandular  pub- 
escence, light  brown  seeds  (except  in 
P.  glandulosa),  compound  leaves,  and 
branched  habit.  P.  coulteri  is  closest  to  the 
ancestral  species.  P.  alba,  P.  denticulata, 
and  P.  neomexicana  are  related  to  P. 
coulteri  but  possess  the  advanced  features 
of  small,  white  to  pale-colored  corollas, 
and    less    robust   habit.     The   migrational 


pedicel  lata 


Fig.  7.     A   phylogenetic   arrangement   of   the 
species  in  the  scariosa  complex. 


134 


GREAT  BASIN  NATURALIST 


Vol.  35,  No.  2 


and  Nevada,  are  disjunct  from  the  others. 
The  species  are  distinguished  by  their 
pale  tubular  corollas,  small  black  seeds, 
and  perennial  or  biennial  habit. 
P.  robusta  is  related  to  P.  pallida  but  is 
considered  to  be  advanced  on  the  basis  of 
its  larger,  reddish  seeds.  P.  palmeri  pos- 
sesses the  advanced  features  of  less-divided 
leaves  and  thin-margined  seeds. 

Crenulata  complex 

Although  not  lacking  in  a  glandular 
pubescence,  this  complex  displays  more 
divergence  from  this  primitive  feature 
than  does  any  other  complex.  There  ap- 
pears to  be  a  bilateral  development,  with 
P.  crenulata  and  its  relatives  becoming 
specialized  with  a  mixed  pubescence  of 
long  stipitate  glands  and  short  hairs  and 
dark  brown  seeds.  P.  anelsonii  and  P. 
coerulea  have  small  corollas  with  included 
stamens  and  style.  The  former  is  more  ad- 
vanced and  possesses  a  thyrsoid  inflores- 
cence and  more  or  less  scarious  sepals. 
The  other  line,  of  which  P.  corrugata  is 
the  primitive  taxon,  is  characterized  by 
light  brown  or  dark  brown  seeds,  short 
stipitate  glands,  and  yellowish  stems. 
P.  hoivellicma  and  P.  integrifolia  are  the 
most  highly  developed  species  in  this  line, 
the  latter  apparently  being  the  most  ad- 
vanced on  the  basis  of  its  lavender  corollas 
and  large,  noncorrugated  seeds.  The 
former  has  large,  dark  brown  seeds  and 
bicolored  corollas.  The  entire  complex, 
with  the  exception  of  P.  intergrifolia,  po- 
ssesses distinctly  corrugated  seeds. 

Congesta  complex 

This  complex  is  related  to  and  has  pos- 
sibly been  derived  from  /-•.  injwidihuli- 
formis  or  some  form  close  to  it.  P.  infuii- 
dibuliforniis  cHffers  from  other  taxa  in  this 
complex  only  in  its  multiovulate,  nar- 
rowly oblong  capsule  and  overall  vege- 
tative appearance.  The  small  white  cor- 
ollas, barely  exserted  stamens  an(]  style, 
and  perennial  habit  of  P.  rupestris  in- 
dicate that  it  is  the  most  advanced  species 
in  this  complex.  The  species  of  the  con- 
gesta complex  occur  in  the  cast  conlral 
part  of  the  range  of  the  Crenulatae  gronjx 

Scariosa  complex 

This  most  advanced  complex  is  char- 
acterized by  distinctly  scarious  sepals,  bi- 
colored   corollas,    and    large,    corrugated. 


transversely  ridged  seeds.  The  species  oc- 
curs in  the  southwesternmost  part  of  the 
range  of  the  grouj). 

Distribution  and  Ecology 

Members  of  the  Crenulatae  group  oc- 
cur mostly  in  western  North  America. 
The  remaining  species,  P.  boliviano 
Brand,  P.  pinnatifida  Griseb.  ex  Wedd., 
and  possibly  others,  occur  in  Peru,  Bolivia, 
and  Argentina.  The  center  of  distribution 
in  North  America,  based  on  the  greatest 
concentration  of  taxa,  is  Arizona  and  New 
Mexico. 

In  general,  members  of  this  group  are 
desert  plants  that  occur  from  near  sea 
level  to  5,000  feet  elevation  (up  to  11,000 
feet) .  Some  are  restricted  to  a  particular 
geologic  formation,  such  as  P.  utahensis, 
which  grows  only  on  the  Arapian  Shale 
formation.  P.  baker i  is  restricted  to  mon- 
tane or  subalpine  regions  on  talus  or  al- 
pine slopes  in  Colorado,  whereas  P.  integ- 
rifolia occurs  mostly  in  deep  sand.  P. 
splendens  is  endemic  to  gypsiferous  soil  in 
western  Colorado  and  northwestern  New 
Mexico.  The  majority  of  taxa  occur  in  the 
Lower  Sonoran  zone  and  are  restricted 
to  an  isolated  mountain  range  or  valley. 
P.  corrugata,  P.  denticulata,  P.  glandulosa, 
and  others  occur  in  the  Upper  Sonoran 
zone  and  generally  have  a  wider  distri- 
bution. P.  denticulata  is  limited  by  the 
continental  divide,  occurring  only  on  its 
eastern  side.  P.  congesta,  P.  pedicellata, 
P.  arizonica,  P.  denticulata,  and  P.  rupe- 
stris are  able  to  survive  in  the  shade  of 
overhanging  ledges  or  as  an  understory 
of  trees  and  shrubs. 

An  important  isolating  mechanism 
that  helps  to  account  for  the  wide  distri- 
bution is  the  seasonal  ^ariati()n  in  phe- 
nology. The  palmeri  (omplex  flowers  in 
late  sunnner  and  fall,  whereas  members 
of  the  welshii  com])lex  flower  in  spring 
and  early  summer.  The  foetid  odor  of 
some  species  is  known  to  attract  beetles, 
while  bees  and  other  insects  are  imj:)ortant 
pollinating  agents  in  other  species. 

The  light,  cymbiform  seeds  are  prob- 
ably wind-dispersed.  Tn  addition  to  wind, 
birds  are  probabh'  an  important  dispers- 
ing agent. 

(Cytology 

Cave  aufi  Constance  (1042.  1944.  1947. 
1950.  1959)  and  Constance  (1963)  have 


June  1975  atwood:   phacelia  crknulatae  (;roup  135 

made   chromosome  comits   on   about  half  alternate,   entire  to  bipinnate,   sometimes 

the   Crenulatae   group,    all   of   which   are  revolute,    sessile    to    long    petiolate;    in- 

11=11.      The     uncounted     members     are  florescence  of  terminal,  axillary,  or  thyr- 

mostly      narrowly      restricted      endemic  soid,    compound,    scorpioid    cymes;    calyx 

plants  and  include  the  following:  P.  and-  divided  nearly  to  the  base,  elliptic  to  lin- 

sonii.  P.   hakeri.  P.   boUviana.  P.  bomby-  ear,    oblanceolate   or    spatulate,    variously 

cina,  P.  constancei,  P.  coulteri,  P.  formo-  jnibescent  and    sometimes   accrescent;   co- 

sula,  P.  glandulosa,  P.  howcUiana,  P.  in-  rolla  white  or  lavender  to  blue,  tubular, 

frgrifolia,  ^  ar.    tcxana,  P.   intermedia.  P.  campanulate  to  rotate-campanulate,  a  pair 

pallida.  P.  serrata.  P.  utahensis,  P.  vossii.  of  variously  shaped  scales  attached  to  the 

and  P.  welshii.  base  of  each  filament,  these  partially  free 

from  or  completely  attached  to  the  tube, 

filament,   or  adjacent   scale;    stamens   ex- 

Taxonomic  Treatment  ^^^^^j   ^^  included   within  the   tube,   and 

Phacelia  Kiis.  Gen.  129.  1789.  inserted  at  the  base  of  the  corolla  tube; 

Subgenus  Phacelia   Constance.   Britt.    15:278.  Style  exserted  or  included  withm  the  tube, 

1963.  bifid    1/2-3/4   its   length,    capsule   nearly 

Section    Phacelia    Brand.    Das    Pflanzenreich  bilocular  by  union  of  the  placentae,  OVoid 

IV.  251:72.  1913  jq    subglobose,    variously    pubescent    and 

Group   Crenulatae   Constance.    Britt.    15:279.  niostly   glandular;   mature   seeds  4    (1,  2 

or   4    in    P.    amabilis   and    P.    congesta)^ 

Annual,    biennial,    or    j:)erennial    herbs  light    brown   to   black,    favose,    reticulate, 

from    a    taproot;    stems    simple    to    much  entire  to  corrugated  or  tranversely  ridged, 

branched,    erect,    ascending    or    prostrate,  excavated   on  both   sides   of  a   prominent 

leafy,    puberulent    to    hispid,    strigose    or  ridge    (except   in   P.    bakeri)    and   mostly 

\ariously    glandular;    leaves    jirevailingly  cymbiform. 

Key  to  the  species  of  the  Crenulatae  group 

la.      Stamens  and  style  included  or  nearly   so    ----- 2 

lb.     Stamens  and  style  exserted  2  mm  or  more   - - 4 

2a.     Corolla  tubular,  light  blue,  lobes  denticulate;    plants    of    Colorado    and 

Wyoming    15.   P.   denticulata 

2b.     Corolla  campanulate  or  rotate-campanulate,    lobes    entire    or    at    most 

crenulate    - — 3 

3a.     Plants  brittle,   breaking  easily;   corolla  3-4  mm  long,  pale  mauve  to  light 

blue;  mature  seeds  dark  brown  9.  P.  coerulea 

3b.     Plants    not   brittle;    corolla    ca.    6   mm    long,    lavender    or    white;    seeds 

brown -- 4.  P.  anelsonii 

4a.     Pedicels  shorter  than  the  calyx;  sepals  not  scarious  in  fruit  6 

4b.     Pedicels  filiform  (at  least  as  long  as  the   calyx  lobes);    sepals   scarious  in 

fruit;    leaves    pinnately    compound,  the  divisions  broad  5 

5a.  Sepals  less  than  3  times  longer  than  broad;  mature  seeds  2.5  mm  long 
or  less;  plants  of  lower  Baja  California  and  southwestern  Sonora, 
Mexico  - - - 30.  P.  scariosa 

5b.  Sepals  3  or  more  times  longer  than  broad;  mature  seeds  2.5  mm  long  or 
more;  plants  of  central  Baja  California  north  to  California,  Ari- 
zona,  and   Nevada   25.  P.  pedicellata 

6a.     Corolla  over  4  mm  long,  white  or  variously   colored   13 

6b.     Corolla  small   (4  mm  long  or  less),  white,  blue,  or  lavender 7 

7a.     Plants  prostrate,  diffusely  branched    (at  the  base);  mature  seeds  1.8-1.9 

mm  long,  ovate,  transversely  ridged;  corolla  white  - 6.  P.  arizonica 


136  GREAT  BASIN  NATURALIST  Vol.  35,  No.  2 

7b.     Plants  erect,  mature  seeds  mostly  over    2    mm    long,    if    smaller    then 

not    with    the    above    combination    of    characters 8 

8a.  Corolla  lobes  entire;  mature  seeds  with  the  ridge  corrugated  or  the  dor- 
sal   surface    reticulate 11 

8b.     Corolla  lobes  erose  or  denticulate;  mature     seeds     pitted,     margins     and 

ridge  entire 9 

9a.     Corolla  white  or  pale  colored,  3-4  mm  long 1.  P.  alba 

9b.     Corolla  blue  or  purple,  4-5  mm  long  10 

10a.  Stems  thick,  robust;  corolla  4-5  mm  long,  bluish  purple;  mature 
seeds  1.6-1.9  mm  long;  plants  endemic  to  the  states  of  Hidalgo  and 
Zacatecas,  Mexico 13.  P.  coulteri 

10b.     Stems  weak;  corolla  4  mm  long,  blue;  mature  seeds  3.2-3.3  mm  long; 

plants  of  Arizona   and  New  Mexico    22.    P.    neomexicana 

11a.  Corolla  white;  mature  seeds  2.1-2.7  mm  long,  the  ridge  not  corrugated, 
dorsal  surface  reticulate;  plants  from  southeastern  Arizona  eastward 
29.   P.   rupestris 

lib.     Corolla  blue  to  light  violet;  seeds  2.5-3.2  mm  long,  the  ridge  corrugated 

_._._ 12 

12a.  Corolla  bicolored  (tube  white,  lobes  blue  to  lavender),  campanulate; 
seeds  ovate,  ridge  corrugated,  margins  entire;  plants  from  western  Ar- 
izona   westward    3b.  P.  ambigua  var.  minutiflora 

12b.  Corolla  not  bicolored,  light  blue  to  lavender,  rotate;  seeds  elliptic  to  ob- 
long, ridge  and  margins  corrugated;  endemic  to  Coconino  County, 
Arizona    31.   P.  serrata 

13a.     Corolla  distinctly  tubular,  white  or  pale  colored 14 

Hb.  Corolla  campanulate,  purple,  blue,  lavender,  or  white  (appearing  tu- 
bular  in    some   pressed    specimens) -. 19 

14a.     Plants  annual  or  biennial  (possibly  perennial  in  P.  pallida) ;  north  of  Nu- 

evo  Leon,  Mexico — -. 15 

14b.     Plants  perennial;  endemic  to  the  state  of  Nuevo  Leon,  Mexico 

34.    P.    vossii 

15a.     Seeds  brown  or  reddish  brown,  2.9  mm  long  or  more 16 

15b.     Seeds  black,  2.9  mm  long  or  less  17 

16a.     Seeds   3.5-4   mm   long;    cauline   leaves    sessile     (or    nearly    so),    auric- 

ulate;  plants  of  Utah  -.. 27.    P.    rafaelensis 

16b.     Seeds  2.9-3.7  mm  long;   cauline  leaves    distinctly    petiolate,    not    auric- 

ulate;  plants  of  Kansas,  Oklahoma,  Texas,  and  Mexico 28.  P.  rolmsta 

17a.     Inflorescence    thyrsoid;    stems    solitary  or  if  branched  then  near  the  base 

..^. 24.    P.    palmeri 

17b.     Inflorescence  open;  stems  branched  throngiiont,  especially  at  base  18 

18a.      Leaves    revolute,    narrowly    linear   or     lanceolate      (less     than     1.5     cm 

wide);    plants  of  Utah   and   Arizona  11.  P.  constancei 

18b.     Leaves    not    revolute.    broadly   oblong    or    lanceolate    (mostly    over    1.5 

cm  wide);    plants  of  Texas  and   adjiKciit    Mfwico    23.    P.   pallida 

19a.      Leaves   pinnately    or    bipinnately    compound,    fiiu^iy   dissected    20 

19b.     Leaves  simple  or  if  compound  not  fiiu^l\'  so.  the  divisions  broad   (over  5 

mm    wide)    25 


June  1975  atwood:  phacelia  crenulatae  group  137 

20a.     Corolla  violet;  plants  endemic  to  Jackson    County,    Colorado    

16.    P.   formosula 

20b.     Plants    not   as    above    21 

21a.  Pubescence  of  the  leaves  mostly  unicellular,  j)uberulent  to  hispid;  plants 
native  from  western  and  southern  New  Mexico,  southward  and  east- 
ward  (except  in  P.  orgillacea  which  is  endemic  to  Utah)    22 

21b.     Leaves     mostl}'     with     multicellular,     stipitate-glandular   hairs;    native 

from  central  New  Mexico  northward  and  westward 24 

22a.  Flowers  violet  to  light  blue;  seeds  more  or  less  excavated  on  one  side  of 
the  ventral  ridge;  ])lants  endemic  to  the  Green  River  Shale  formation 
in  Utah  County,  Utah 5.    P.    argillacea 

22b.     Flowers  blue  to  purple;  seeds  excavated    on    both    sides    of   the   ventral 

ridge - 23 

23a.     Seeds  1.8  mm  long,  ovate;  ultimate   leaf    divisions    mostly    less    than    4 

mm  wide - 26.  P.  popei 

23b.     Seeds  over  1.8  mm  long,  elliptic  to  oblong;  ultimate  leaf  divisions  mostly 

over  5  mm  wide 10.    P.    congesta 

24a.     Seeds  not  excavated  ventrally;   plants  east  of  the  Continental  Divide  in 

Colorado   and   New  Mexico 7.  P.  hakcri 

24b.     Seeds  excavated  ventrally;  plants  west    of    the    Continental    Divide    in 

Wyoming,  Montana,  and  Idaho  17.  P.  glandulosa 

25a.     Corolla   not  distinctly   bicolored,  blue,   purple,  or  white  28 

25b.     Corolla   distinctly  bicolored,  the  tube  white  or  yellow,  the  lobes  blue 26 

26a.  Cauline  leaves  sessile,  auriculate,  plants  robust,  0.8-5.8  dm  tall,  en- 
demic to  Sanpete  and  Sevier  counties,  Utah 33.  P.  utahcnsis 

26b.     Cauline  leaves  distinctly  petiolate;  plants  not  especially  robust,  less  than 

2.7  dm  tall,  more  eastern  in  distribution 27 

27a.  Stems  branched  at  base;  leaves  simple,  strigose  and  glandular;  corolla 
tube  white;  seeds  corrugated  on  the  margins  and  ridge,  dorsal  sur- 
face   smooth    18.   P.   howelliana 

27b.  Stems  simple  or  branched  above;  leaves  essentially  glabrous,  some  of 
the  lower  usually  compound;  corolla  tube  yellowish;  seeds  essen- 
tially lacking  corrugations,  dorsal  surface  dee])ly  j)itted  32.  P.  splendens 

28a.  Corolla  white;  plants  endemic  to  Saline  Valley,  Inyo  County,  Califor- 
nia  :: 2.  P.  amabilis 

28b.  Corolla  pale  blue,  purple,  or  lavender  (rarely  white);  plants  wide- 
spread or  if  endemic  not  as  above 29 

Corolla   lavender;   seeds   lacking  ventral    corrugations 

19a.  P.  integrifoUa  var.  integrifolia 

Corolla  pale  blue  to  purple;  seeds  corrugated  ventrally 30 

Stamens  and  style  exserted  4  mm  or  less;  mature  seeds  2.9  mm  long  or 

less 31 

Stamens  and  style  exserted  over  4  mm;  mature  seeds  over  2.0  mm  long 

32 

31a.  Mature  seeds  2.2-2.5  mm  long;  dark  brown;  plants  very  brittle  an- 
nuals   8.  P.  bombycina 

31b.     Mature  seeds  2.7-2.9  mm  long;   brown;     plants    not    especially    brittle 

20.  P.  intermedia 


138  GREAT  BASIN  NATURALIST  Vol.  35,  No.  2 

32a.     Mature  seeds  corrugated  only  on  the    ridge;    pubescence    of   the    stems 

densely  hispid,  glandular  above 3a.  P.  ambigua  var.  ambigua 

32b.     Seeds  with  the  margins  and  ridge  corrugated;    pubescence    of   the    stems 

mostly    glandular,    sometimes   finely  so 33 

33a.     Glandular  pubescence  of  the  stems  and   herbage  mostly  multicellular  34 

33b.  Glandular  pubsecence  of  the  stems  and  herbage  stipitate  but  not  multi- 
cellular   35 

34a.  Corolla  lavender  to  purple,  4.5-6  mm  long;  anthers  blue  green;  stems 
often  reddish;  mature  seeds  reddish  brown,  2.4-3  mm  long;  plants  of 

southeastern  New  Mexico  and  adjacent  Texas  

19b.  integrifolia  Torr.  var.  tcxana 

34b.  Corolla  blue,  6  mm  long  or  longer;  anthers  always  yellow;  stems  green 
or  yellow  green;  mature  seeds  light    brown,    3.1-4    mm    long;    plants 

of  northwestern  New  Mexico,  Arizona,  Colorado,  and  Utah 

12.  P.  corrugata 

35a.  Leaves  narrowly  oblong,  mostly  less  than  1  cm  wide,  glandular  pub- 
escence of  the  leaves  and  lower  part  of  the  stems  short  stipitate,  the 
nonglandular  hairs  mostly  fine,  retrorse  ..  14a.  P.  crenulata  var.  angustifolia 

35b.     Leaves  mostly  well  over  1   cm  wide,  glandular  pubescence  of  the  leaves 

and  stems  stipitate-multicellular,   nonglandular  hairs  spreading  36 

36a.  Stems  usually  reddish  at  least  below;  mature  seeds  with  a  dark  cen- 
ter dorsally  and  lighter  margins;  plants  of  western  Utah  and  Ar- 
izona   westward    to    California    and    Nevada 

14b.  P.  crenulata  I'orr.  var.  crenulata 

36b.     Stems  green;  mature  seeds  uniform  in  color  dorsally;  plants  endemic  to 

Coconino  County,  Arizona  '. 37 

37a.  Corolla  lavender  to  white,  campanulate,  5-6  mm  long;  leaves  un- 
dulate   to    regularl}'    dentate,    oblong  to  lanceolate;   seeds  uniformly 

brow2i;  plants  endemic  to  Kane  and    Garfield   counties,   Utah  

21.   P.   mammillarensis 

37b.  Corolla  dark  blue,  tubular  to  funnelform,  5-8  mm  long;  leaves  irreg- 
ularly crenate  to  dentate,  lanceolate;  seeds  brown  ventrally  and 
reddish  dorsally;  plants  endemic  to  Coconino  County,  Arizona  .... 
35.    P.    ivelshii 

1.  Phacelia  alba  Rvdberg  July  1899,  Townsend  and  Barber,   129,   (ny); 

-p-^.    o       '  Isotypes    (cAs,  mexu,  pom,  rm,  uc,  uc,  us). 

^"  Phacelia  glandulosa  Nutt.   ssp.   eu-glandulosa 

Phacelia  alba  Rydb..  Bull.  Torr.  Bot.  Club  28:30.  Brand  var.  elatior  Brand.   Das  Pflanzcnreich 

1901.  IV.  251:82-83.   1913.  in  part.  Holotype:   Wyo- 

Holotype:    Colorado:    Costilla   Co.:    Sangre  de  ining-'    Albany    Co.:    Jolm,    12   August    1900, 

Christo    Creek,    2    July    1900,    Rydberg    and  A.  Nelson.  8053.   (gh,  pom,  rm). 

Vroeland,      5755,       (ny?);      Isotype       (rm/). 

Paratypes:    Colorado:     Clear    Creek    County:  Plants     annual.     0.5-7     (hn     tall;     stems 

Valley    of    Upper    Arkansas    River,    1873.    J.  simple    to    much    branched,    erect    or    as- 

™f«-„^9'  (NY);  head  waters  of  Clear  Creek,  ,o,Kli,,o.    loafy,    puberulent,    setose    to   hir- 
1861.  C.  Parry,  314  (gh,  ny);  New  Mexico:  ^         ^ ,      ^.    •.  ■    '       ,        i    i  •    n       • 

Lincoln  Co.:  "  Ruidoso   Creek.   White   Moun  ''^'^^  •'"^'   stipitate-glandular,  especially  in 

tains,  1  July  1895,  E.  Wooton,  s.n..   (ny).  the  ijiflorescence;  leaves  irregularly  lobed 

Phacelia  neomexicana  Thurber  ox  Torr.  var-.  to    bipiniiate,    2-10    (.  ni     wide,    strigose    tO 

alba    (R.ydb.)    Brand.   Das   Pflan/.eiueich    IV.  setose,     slightly     to     moderately     stipitate- 

251:83.  1913.  glandular,  long  petiolate  below  to  sessile 

Phacelia  neomexicana  Thurl^er  ex  To'. .  var.  ;,j.  ^ubsessile  above;  inflorescence  of  dense 

couUeri  subvar.  folisissima  Brand.  Das  Pflan-  .  .       ,  ,       -  •    .  , 

zenreich  IV.  251:84.  1913.  Holotype:  Mexico:  terminal      compound      scorpioid      cymes, 

state  of  Chihuahua:   near  Colonia  Garcia.    13  densely  glandular  and    puberulent  to  hir- 


June  1975 


ATWOOD:     PHACELIA    CRENULATAK    GROUP 


139 


sute,  the  cymes  1-2  cm  long  in  flower  to 
8  cm  long  in  fruit,  pedicels  0.8-1  mm  long; 
sepals  linear  to  oblanceolate,  3.5-4  mm 
long,  0.5-1  mm  wide,  finely  glandular 
and  somewhat  setose;  corolla  campanulate, 
white  (sometimes  pale  purple),  3-4  mm 
long  and  broad,  lobes  pubescent  and  den- 
ticulate; capsule  ovoid  to  subglobose,  3- 
3.3  mm  long,  2.5-2.9  mm  wide,  puber- 
ulent  and  quite  glandular  (the  partition 
oblanceolate) ;  mature  seeds  elliptic  to  ob- 
long, light  to  dark  brown,  2.4-3  mm  long, 
1.4-1.5  mm  wide,  uniformly  alveolate 
throughout  and  cymbiform,  the  ventral 
surface  shallowly  excavated  on  both  sides 
of  the  ridge  and  lacking  corrugations,  the 
margins  thick  and  entire  (Fig.  9).  Col- 
lections: 207  (5);  representative:  C.  Parry 
314,  1861  (gh,  ny);  D.  Atwood  1975, 
1962,  1963a  (bry.  ny.  us);  M.  Jones  511 
(pom). 

Habitat. —  Dry  clay-loam  or  sandy 
draws  and  flats,  fields,  meadows,  and 
gravelly  hillsides.  From  6,000  to  9,500 
feet.  Growing  on  the  short  grass  prairie, 
sagebrush  belt,  and  pinyon-juniper  com- 
munities at  the  lower  elevations  and  as- 
sociated with  aspen,  spruce,  fir,  or  pine 
forests  at  the  higher  elevations.  Late 
May  to  early  October. 

Distribution. —  Laramie  and  Albany 
COS.,  Wyoming,  southward  through  cen- 
tral Colorado,  New  Mexico,  and  adjacent 
Chihuahua,  Mexico,  westward  to  Gra- 
ham and  Apache  cos.,  Arizona,  and  Sevier, 
Wayne,  Garfield,  and  Washington  cos., 
Utah  (Map  1). 

The  material  cited  by  Brand  (1913) 
for  P.  glandulosa  ssp.  eu-glandulosa  var. 


Fig.  8.     Phacelia   alba    Rydberg.    L.    Higgins 
2229  (bry). 


Fig.  9.     Dorsal  and  ventral  view  of  the  seeds 
of  P.  alba  Rydberg.  D.  Atwood  1975  (bry). 


140 


GREAT  BASIN  NATURALIST 


Vol.  35,  No.  2 


elatior  Brand    (Nelson   8053)    belongs  to 
P.  alba. 


County,   California,   along  Hunter   Creek 
at  an  elevation  of  1,800  feet. 


2.  Phacelia  amabilis  Constance 

Phacelia    amabilis    Constance.    Madrono    7:56-59. 

1943. 

Holotvpe:  California:  Inyo  County:  Saline 
Valley,  21  April  1942,  A.  Alexander  and  L. 
Kellogg  2681    (uc!);  Isotype    (uc). 

Plants  annual;  stems  stout,  branched 
above,  stipitate-glandular,  puberulent,  and 
hispid;  leaves  petiolate,  oblong  to  oblong- 
ovate,  8-15  cm  long,  3-5  cm  wide,  pin- 
natifid,  upper  leaves  reduced  and  less 
deepl}"  divided;  inflorescence  of  com- 
pound scorpioid  cymes,  the  cymes  5-12 
cm  long,  pedicels  2-3  mm  long;  sepals 
lanceolate,  3-5  mm  long,  1-2  mm  wide; 
corolla  broadly  campanulate,  white,  7-8 
mm  long,  8-12  mm  wide;  stamens  and 
style  exserted  5  mm  or  more;  capsule 
ovoid,  3-4  mm  long,  2-3  mm  wide;  im- 
mature seeds  apparently  2  or  4,  3-4  mm 
long,  thin  and  pale,  ventral  surface  ex- 
cavated on  each  side  of  the  prominent 
ridge;  collections:  1  (0);  representative: 
A.  Alexander  and  L.  Kellogg  2681    (uc). 

Distribution  and  Habitat. —  Appar- 
ently   endemic    to    Saline    Valley,    Inyo 


3.  Phacelia  amhigua  Jones 

Plants  annual,  0.2-5.7  dm  tall;  stems 
simple  to  much  branched,  hispid,  pub- 
erulent, and  stipitate-glandular;  leaves 
simple  to  pinnately  compound,  petiolate 
to  sessile  above,  the  margins  various,  stri- 
gose  to  hispid  and  stipitate-glandular,  0.5- 
13  cm  long,  0.5-4.5  cm  wide;  inflores- 
cence of  compound  scorpioid  cymes,  the 
cymes  elongating  to  12  cm  in  fruit,  pub- 
escence as  for  the  stem;  sepals  elliptic  to 
oblanceolate,  2.7-5.1  mm  long,  1-1.3  mm 
wide,  puberulent,  hispid,  and  stipitate- 
glandular;  corolla  campanulate  to  rotate- 
campanulate,  purple  or  dull  lavender, 
4-10  mm  long  and  broad,  pubescent;  sta- 
mens and  style  exserted  2-10  mm;  style 
bifid,  pubescent  below;  capsule  globose  to 
subglobose,  3-3.5  mm  long,  2.5-3.4  mm 
wide,  puberulent  and  glandular;  mature 
seeds  4,  ovate,  reddish  to  brown,  2.5-3.3 
mm  long,  1.5-1.8  mm  wide,  alveolate, 
cymbiform,  the  ventral  surface  excavated 
on  both  sides  of  the  ridge,  the  ridge  cor- 
rugated on  one  side. 


1.  Corolla  4  mm  long  or  less;  style  6.5  mm 
than  4  mm  long 

1.  Corollas  over  4  mm  long;  style  9  mm 
mm  long 


Key  to  the  varieties  of  P.  amhigua 

long  or  less;   calyx  in   fruit  less 


var.   minutiflora 

more;    calyx   in   fruit  4 
var.    amhigua 


3a.  var.  amhigua 

Phacelia  ambigua  Jones.  Contr.  West.  Bot.   12:52. 

1908. 

Holotype:  California:  San  Bernardino  Co.: 
Needles,  5  May  1884,  M.  Jones  3822.  (pom!); 
duplicates  (ny,  rm,  uc,  us).  Paratypes:  Ari- 
zona: Coconino  Co.;  Hole  in  the  Bock.  13 
April  1894.  M.  Jones  s.n..  (?):  Nevada:  Lin- 
coln Co.:  Calienfe.  29  April  1904,  M.  Jones, 
s.n.,   (?). 

Phacelia  crenulata  Torr.  in  Wats.  var.  am- 
bigua (Jones)  Macbride.  Contr.  Gray  Herb. 
49:25.   1917. 

Plants  annual,  0.5-5.7  dm  tall;  stems 
simple  to  much  branched,  usually  more 
leafy  toward  base;  leaves  0.5-13  cm  long, 
0.5-4.5  cm  wide,  slrigose  to  hispid,  us- 
ually only  with  scattered  glands,  reduced 
from  the  base  upward;  cymes  elongating 
to  12  cm  in  fruit;  sepals  3.2-5.1  mm  long, 
1-1.3    mm     wide;     corolla     campanulate. 


purple  to  blue,  5-10  mm  long  and  wide, 
pubescent;  stamens  and  style  exserted  9 
mm  or  more;  style  bifid  2/3  its  length, 
puberulent  and  glandular  below;  capsule 
3.3-3.5  mm  long,  3-14  mm  wide,  puber- 
ulent and  stipitate-glandular;  mature 
seeds  3.3  mm  long,  1.5  mm  wide.  Col- 
lections: 250  (30);  representative:  E.  Pal- 
mer 625  (ny);  M.  Jones  5018  (ny,  pom, 
RM.  uc.  us);  C.  Pringle  s.n.  (cas,  gh, 
ny);  J.  Howell  3504  (rsa);  D.  Atwood 
2210,  2220a,  2294,  2296,  2303,  2310,  2319, 
2353  (bry). 

Habitat. —  Growing  on  a  wide  \  ariety 
of  soils  in  the  lower  Sonoran  Desert  from 
490  to  5,000  feet  elevation.  February  to 
mid-June. 

Distribution. —  vSouthern  Nevada  and 
south wostoni    Utah    in    Washington    Co., 


June  1975 


ATWOOD:    PHACELIA   CRENULATAE   GROUP 


141 


N 

^^" 

n 

1 

:               -J 

••• 

1 

L  ^ 

• 
/          t 

• 

1          V 

1 

)     • 

/ 

? 

Map  1.  Southern  Wyoming,  Utah,  Colorado. 
Arizona,  New  Mexico,  and  Chihuahua,  Mexico. 
Distribution  of  P.  alba  Rydberg. 

southward  through  Arizona  (except  for 
Navajo  and  Apache  cos.,)  and  southeast- 
ern California    (Map  2). 

Phacelia  arnbigua  has  been  treated  as 
a  variety  of  P.  crenulata,  which  it  closely 
resembles.  However,  the  former  species 
appears  to  be  more  uniform  throughout  its 
range  than  does  the  latter.  Also,  to  treat 
P.  arnbigua  at  infraspecific  rank  would 
require  the  inclusion  of  other  closely  re- 
lated taxa,  namely  P.  bombycina  W.  &  S. 
and  P.  amabilis  Constance.  In  order  to 
understand  the  complete  relationships  be- 
tween these  taxa  additional  field  and 
greenhouse  studies  are  needed.  These 
studies  may  demand  nomenclature 
changes,  but  until  such  studies  are  carried 
out  the  present  treatment  provides  a  more 
uniform  arrangement  of  the  entities  in- 
volved. 

3b.  var.  minutiflora  (Voss  in  Munz) 

Atwood  comb.  nov. 

Fig.  10 

Phacelia    minutiflora    Voss    in    Munz,    Man    So. 

Calif.  Bot.  409,  600.  1935. 

Holotype:    California:    Imperial   Co.:    2  miles 
north     of     Cargo     Muchacho     Mountains.     5 


April  1932,  P.  Munz  and  L.  Hitchcock  12141 
(pom!). 

Phacelia  crenulata  Torr.  in  Wats.  var.  minuti- 
flora  (Voss)   Jeps.,  Fl.  Calif.  3:266.     1943. 

Plants  annual,  0.2-4.5  dm  tall;  stems 
simple  or  variously  branched,  hispid, 
puberulent  and  stipitate-glandular  (es- 
pecially in  the  inflorescence) ;  leaves  0.5- 
11  cm  long,  0.5-3.5  cm  wide,  strigose  to 
hispid  and  often  quite  glandular,  the 
lower  with  longer  petioles  than  the  up- 
per reduced  leaves,  cymes  elongating  to 
7  cm  in  fruit;  sepals  2.7-3.8  mm  long,  1 
mm  wide,  corolla  rotate-campanulate, 
tube  white,  lobes  lavender  to  blue,  pub- 
escent, 4  mm  long  and  broad;  stamens  and 
style  exserted  2  mm  or  less;  style  bifid 
2/3  its  length,  puberulent  and  glandular 
below;  capsule  3-3.2  mm  long,  2.5-3  mm 
wide,  puberulent  and  glandular;  mature 
seeds  2.5-3.2  nun  long,  1.3-1.8  mm  wide 
(Fig.  11).  Collections:  89  (4);  repre- 
sentative: E.  Palmer  626  (ny);  T.  Kear- 
ney and  R.  Peebles  10941,  10963,  11016 
(us);  D.  Atwood  2320,  2355,  2352,  2341 
(bry);  I.  Wiggins  9669   (rsa,  uc). 

Habitat. —  Sandy  to  rocky  desert  flats, 
washes  and  slopes  from  near  sea  level  to 
2,200  feet.  Common  in  Larrea,  Ambrosia. 
Atriplex,  Fouguieria,  and  Ccrcidiuin  com- 
munities.   Late  December  to  late  April. 


Map  2.  Utah,  Nevada,  Arizona,  California, 
and  adjacent  Mexico.  Distribution  of  P.  ambigua 
Jones:    O  var.   ambigua;   var.   minutiflora.    # 


142 


GREAT  BASIN  NATURALIST 


Vol.  35,  No.  2 


.^ 


^v(  ;i^  \     m 


Fig.   10.     Phacclia  ambigua  Jones  yav.  minutiflora  (Voss  in  Muiiz)  Atvvood.  D.  Atwood  2341   (bry). 


Distribution. —  Southwestern  Arizona 
in  Maricopa,  Pima,  and  Yuma  cos.,  and 
westward  to  San  Bernardino  Co.,  Califor- 
nia, south  into  Baja  California  and  Sonora, 
Mexico  (Map  2). 

1.  Phacclia  anclsonii  Macbride 
Fig.  12 

Phacelia  anelsonii   Macbride,   Contr.    Gray   Herb. 
49:26.     1917. 

Holotype:  Nevada:  Lincoln  Co.:  Meadow  Val- 


ley  Wash.   28   April    1902,   L.   Goodding   635 

(rm!). 

Erect  annual,  1-5.5  dm  high;  stems 
terete,  usually  simple  covered  \\dth  brown- 
ish stipitate  glands,  leafy  throughout; 
leaves  narrowly  to  broadly  oblong,  pin- 
natcly  cleft,  1.5-8  cm  wide,  pubescence 
brownish,  stipitate-glandular,  with  a  few  j 
non-glandular  hairs,  jiedicels  from  3  cm 
long  on  the  lower  part  of  the  stem  to 
nearly  sessile  on  the  upper  part,  the  pin- 


June  1975 


ATWOOD:     PHACELIA   CRENULATAE   GROUP 


143 


plant  which  Goodding,  when  he  des- 
cribed his  P.  foetida  .  .  .  took  to  represent 
P.  palmeri  Wats.,  a  very  different  plant 
with    exserted    stamens    and    corrugated 


Fig.  11.  Dorsal  and  ventral  view  of  the 
seeds  of  P.  ambigua  Jones  var.  minuiiflora  (Voss 
in   Munz)    Atvvood.    D.   Atwood   2320    (bry). 

nae  somewhat  reduced  toward  the  base 
of  leaf,  margins  crenate;  inflorescence 
racemose  to  paniculate,  usually  terminal 
on  the  upper  half  of  the  stem,  some- 
times on  leafy  lateral  branches,  individual 
cymes  1-5  cm  long,  setose  and  glandular 
pubescent;  corolla  light  violet  or  white, 
rotate-campanulate,  6  mm  long  and  wdde; 
sepals  oblanceolate  to  spatulate,  3-6  mm 
long,  1-2  mm  wide,  setose  to  glandular, 
1-2  mm  longer  than  the  capsule;  stamens 
included,  anthers  yellow;  style  included, 
3.5-4.8  mm  long,  shorter  than  the  stamens, 
cleft  2/3  its  length,  glandular  and  puber- 
ulent  at  the  base;  capsule  oval,  3.3-3.7 
mm  long,  glandular  spotted  throughout 
and  pilose  on  the  upper  half;  mature  seeds 
4,  oblong,  2.7-3.4  mm  long,  1-1.3  mm 
wide,  light  browai,  margins  entire,  ventral 
surface  strongly  alveolate,  divided  by  a 
prominent  ridge,  ridge  corrugated  along 
one  side,  dorsal  surface  aheolate  (Fig.  13). 
Collections:  30  (1);  representative:  L. 
Goodding  635  (rm);  R.  Barneby  2937 
(cAS,  rsa)  :  S.  Welsh,  D.  Atwoood,  and  E. 
Mathews  9542  (bry);  L.  Higgins  499 
(bry). 

Habitat. —  Commonly  in  shady  places 
at  the  base  of  sandstone  and  limestone 
cliffs  or  among  rocks  and  in  sandy  to  grav- 
elly washes,  2,000  to  5,000  feet  elevation. 
Usually  locally  scattered,  April  to  May. 

Distribution. —  Lincoln  Co.,  Nevada, 
southward  to  Washington  Co.,  Utah,  Inyo 
and  San  Bernardino  cos.,  California  (Map 
3). 

Macbride  (1917)  in  his  original  des- 
cription of  P.  anelsonii  savs,  "this  is  the 


Fig.  12. 
Ripley  and  R. 


Phacelia     anelsonii     Macbride.     H. 
Barneby  3496  (cas). 


144 


GREAT  BASIN  NATURALIST 


Vol.  35,  No.  2 


seeds."  It  is  not  known  from  whence 
Macbride  drew  this  conclusion,  but  it 
supports  the  observations  of  the  author 
that  P.  foetida  is  the  same  entity  as 
P.  palmeri. 

P.  anelsonii  is  related  to  P.  crenulata 
and  P.  amabilis  as  indicated  by  both 
leaves  and  seed  characters.  P.  anelsonii 
is  easily  distinguished  from  them  by  the 
included  stamens  and  style,  a  feature 
which  it  has  in  conunon  with  P.  coerulea 
and  P.  denticulaia.  Detailed  observations 
of  the  type  specimens,  as  well  as  other 
collections,  revealed  the  presence  of  cor- 
rugations along  one  side  of  the  ridge  in 
some  seeds  of  P.  anelsonii.  Both  P.  cren- 
ulata and  P.  amabilis  have  the  corrugated 
ridge.  Macbride  (1917)  and  Voss  (1937) 
have  indicated  that  the  seeds  lack  cor- 
rugations. 

John  Thomas  Howell  was  the  first  to 
report  this  taxon  for  Washington  Co., 
Utah,  and  San  Bernardino  Co.,  California 
(1941),  and  for  Inyo  Co.,  Cahfornia 
(1942).  Two  collections  from  Washing- 
ton Co.,  Utah  (R.  Barneby  2937  and  B. 
Wood  140),  are  more  robust  in  habit  and 
have  unusually  large  parts. 

5.  Phacelia  argillacea  Atwood 

Phacelia  argillacea  Atwood.  Phytologia  26(6): 437. 

1973. 

Phacelia  glandulosa  Nutt.  var.  argillacea  At- 
wood in  Welsh  &  Moore,  nomen.  nudum. 
Holotype:  Utah  Co.:  Spanish  Fork  Canj-on. 
Clear  Creek  ca.  6  mi  west  of  Soldier  Summit. 
18  August  1971.  D.  Atwood  et  al.  3091 
(bry);  Isotypes,  to  be  distributed;  Pleasant 
Valley    Junction    (Colton).    Wasatch    Moun- 


N 

1 

• 

I 

^^\^ 

1 

• 

• 

•  • 

^s^ 

k 

• 

• 

Fig.   13.     Dorsal    and    ventral    view    of    tlie 
seeds  of  P.  anelsonii  Macbride.  R.  Barneby  2937 

(CAs). 


Map  3.  Southwestern  Utali,  southern  Ne- 
vada, and  California.  Distribution  of  P.  anelsonii 
Macbride. 

tains,  August  1883,  M.  E.  Jones  s.n.  (us!); 
Isotypes  (c.^s,  cas,  ny,  ny,  pom,  uc,  uc). 
Paratypes:  Utah:  Utah  Co.:  Clear  Creek 
near  Soldier  Summit.  6  July  1894,  M.  E. 
Jones   5591    (ny,   pom,   uc). 

Plants  annual,  1-3.6  dm  tall;  stems 
finely  })ubescent;  leaves  oblong  in  outline, 
pinnatifid,  0.8-5  cm  long,  0.5-1.5  cm  wide, 
strigose,  petiolate;  inflorescence  of  com- 
pound scorpioid  cymes,  stipitate-glandular 
and  setose  to  hirsute,  pedicels  0.7-1  mm 
long,  cymes  elongating  to  7.5  cm  in  fruit; 
sepals  elliptical  to  oblanceolate,  2-3.8  mm 
long,  1  mm  v\dde,  stipitate-glandular  and 
hirsute  to  setose;  corolla  campanulate. 
bluish  violet,  ca.  5  mm  long  and  broad, 
lobes  pubescent;  stamens  and  style  ex- 
serted  en.  7  nmi;  capsule  subglobose,  3.2- 
3.3  nun  long,  2.3-2.4  mm  wide,  glandular 
and  setose;  mature  seeds  4,  brown,  ovate 
to  elliptic,  2.4  mm  long,  1.1  mm  wide, 
pitted,  the  ridge  curved  and  more  or  less 
excavated  on  one  side.  Collections:  3 
(1);  representative:  M.  Jones  s.n.  (cas. 
NY,  POM,  RM.  uc,  us);  M.  Jones  5591 
(ny,  pom,  uc);  D.  Atwood  3091    (bry). 

Habitat. —  The  species  ap])arentl^ 
grows  on  gravelly  hillsides  of  the  Green- 
river  formation  between  (i.5()0  and  7.000 
feet,  July  to  August. 

DisTiuBUTiON. —  Known  only  from 
Utah  Co.,  Utah. 

This  species  is  related  to  both  P.  inland- 
iilosa  and  to  P.  bakeri  but  can  be  (Hstin- 
guished  by  its  more  nearly  glabrous  herb 
age,    smaller    capsule,    flowers,    and    dil 
ferent  seeds. 


June  1975 


ATWOOD:    PHACELIA  CRENULATAE  GROUP 


145 


6.  Phacelia  arizonica  A.  Gray 
Fig.  14 

Phacelia  arizonica  A.   Gray,   Syn.   Fl.   II,    1:394. 

1878. 

Lectotype:    Arizona:    Maricopa  Co.:   plains  of 

the  upper  Gila,  15  April  1880,  E.  Greene  s.n. 

(gh!);  duplicate  (c^s). 

Phacelia   popei   Torr.    &   Gray  var.   arizonica 

(Gray)   Voss,  Bull.  Torr.  Bot.  Club  64:94-95. 

1937. 

Low  diffusely  branched  (at  the  base), 
prostrate  annual,  0.2-2.1  dm  tall  (up  to  4 
dm  in  extreme  forms);  stems  slender,  1- 
many,  hispid  to  villous  and  glandular, 
often  reddish;  leaves  larger  and  denser 
at  the  base,  oblong  to  linear  in  outline, 
sessile  above  to  short  petiolate  below 
deeply  lobed  to  pinnatifid,  1-5  cm  long, 
0.5-2  cm  wide,  densely  strigose,  inflores- 
cence terminal,  cymes  densely  flowered, 
the  flowers  subsessile  (pedicels  to  0.5  mm 
long),  densely  covered  with  small  gland- 
ular and  short  simple  hairs,  with  some 
longer  simple  hairs  intermixed;  sepals 
oblong  to  elliptic,  2.5-4  mm  long,  1  mm 
wide,  hirsute;  corolla  campanulate,  white 
or  rose  white,  (often  reddish  in  bud),  3-4 
mm  long  and  broad,  lobes  pubescent  and 
shallowly  erose;  stamens  and  style  ex- 
serted  3-4.5  mm;  style  bifid  3/4  its  length, 
the  lower  1/2  puberulent  and  sometimes 
finely  glandular;  mature  seeds  4,  ovate 
and  more  or  less  cymbiform,  brown,  1.8- 
1.9  mm  long,  1.2-1.3  mm  wide,  alveolate, 
ventral  surface  excavated  on  both  sides  of 
the  ridge,  dorsal  surface  alveolate  and 
transversely  ridged  (Fig.  15).  Collections: 
104  (2);  representative:  C.  Pringle  s.n. 
(ny);  L.  Goodding  1035  (ny,  uc,  us); 
M.  Jones  28501,  28503  (pom);  D.  Atwood 
2200a  (bry,  cas,  ny,  wts,  b);  L.  Higgins 
2814  (bry);  D.  Atwood  2186  (b,  bry,  cas, 

NY,   wts). 

Habitat. —  Common  along  roadsides, 
sandy  flats,  and  gravelly  hillsides,  from 
1,600  to  2,500  feet.  Often  growing  with 
Prosopis.  Quercus,  Junipcrus,  or  grass  com- 
munities, mid-February  to  late  June. 

Distribution. —  Arizona  from  Mari- 
copa Co.  southward  into  Sonora,  Mexico, 
and  eastward  to  Sierra  and  I^una  cos..  New 
Mexico  (Map  4). 

This  species  was  treated  as  a  variety  of 
P.  popei  T.  and  G.  by  Voss  (1937),  but  is 
easily  distinguished  as  a  species  on  the 
basis  of  its  smaller,  white,  glandless  co- 
rolla, prostrate  habit,  less-dissected  leaves. 


and  nearly  leafless  stems.  However,  the 
two  are  similar  in  seed  characters  and  in 
the  much-branched  habit.  P.  popei  has 
erect  to  ascending  and  more  rigid  stems 
in  contrast  to  the  slender,  prostrate  stems 
of  Phacelia  arizonica.  Gray  (1878)  did 
not  designate  a  type  specimen.  Therefore, 
the  author  has  chosen  the  collection  of 
Greene,  "on  the  plains  of  the  upper  Gila," 
as  the  lectotype. 

7.  Phacelia  bakeri  (Brand)  Macbride 
Fig.  16 

Phacelia   bakeri    (Brand)    Macbride.   Contr.    Gray 

Herb.  n.  ser.  49:24.     1917. 

Phacelia  crenulata  Torr.  e.x  S.  Wats.  var. 
bakeri  Brand.  Das  Pflanzenreich  IV.  251:78. 
1913.  Holotype:  Colorado:  Ouray  Co.: 
Ouray,  10  August  1901,  C.  F.  Baker  758 
(gh!);  duplicates  (ny.  pom,  rm,  uc,  us). 
Phacelia  glandulosa  Nutt.  subsp.  eu-glandu- 
losa  Brand  var.  australis  Brand,  Das  Pflan- 
zenreich IV.  251:82-83.  1913,  in  part.  Lecto- 
type: Colorado:  Conejos  Co.:  Cumbers.  7  Sep- 
tember 1899.  C.  F.  Baker  549  (us!);  dupli- 
cates (gh.  ny.  pom,  rm,  rm).  Paratypes: 
Colorado:  El  Paso  Co.:  Manitou,  15  July 
1903.  C.  Clements  47.1  (rm),  in  part;  Hins- 
dale Co.:  Lake  Fork  River,  Lake  City.  July 
1893.  C.  A.   Purpus  618    (uc,  uc). 

Annual,  0.5-4.8  dm  tall;  stems  simple  or 
branched,  with  multicellular  stipitate 
glands,  pilose  to  somewhat  hirsute;  leaves 
pinnately  divided,  the  pinnae  irregularly 
crenate  to  dentate,  2-8  cm  long,  0.5-3  cm 
wide,  reduced  upwards,  petioles  0.5-4  cm 
long,  dorsal  surface  strigose,  ventral  sur- 
face strigose,   glandidar  and   setose  along 


•                      N 

• 

••• 

• 
• 

% 

Map  4.  Soutliern  Arizona  and  adjacent  So- 
nora, Mexico,  and  western  New  Mexico.  Distribu- 
tion of  P.  arizonica  Gray. 


146 


GREAT  BASIN  NATURALIST 


Vol.  35,  No.  2 


Fig.  14.     Phacelia  arizonica   Gray.   L.   Higgins  2814  (bry). 


June  1975 


ATWOOD:    PHACELIA   CRENULATAE  GROUP 


147 


Fig.  15.     Dorsal    and    ventral    view    of    the 
seeds  of  P.  arizonica  Gray.  D.  Atwood  2186  (bry). 

the  veins;  inflorescence  of  compound 
scorpioid  c>TTies,  terminal  on  the  main 
stem  and  lateral  branches,  setose  to  puber- 
ulent  and  glandular,  pedicels  1-2  mm 
long;  sepals  oblanceolate  to  narrowly  spat- 
ulate,  1-1.5  mm  longer  than  the  capsule, 
setose  to  puberulent  and  glandular;  co- 
rolla campanulate,  violet  to  dark  blue, 
7-8  mm  long,  5-7  mm  wide,  pubescent; 
stamens  exserted  5-9  mm,  anthers  green- 
ish, filaments  bluish;  style  exserted  bifid 
2/3  its  length  and  pubescent  on  the  lower 
1/3;  capsule  oblong  to  oval,  2.5-4  mm 
long,  3-3.2  mm  wide,  setose  and  gland- 
ular; mature  seeds  elliptic,  brown,  2.7-3 
mm  long,  1.3-1.6  mm  wide,  ventral  sur- 
face pitted  with  a  central  ridge  gradually 
tapering  toward  the  margins,  excavations 
lacking  on  each  side  of  ridge,  dorsal  sur- 
face flattish  with  a  faint  longitudinal 
groove  down  the  center,  pitted  (Fig.  17). 
Collections:  83  (0);  representative:  T.  S. 
Brandegee  1139  (uc);  M.  Jones  511 
(pom);  C.  Baker  549  (gh,  ny,  pom.  rm, 
us);  L.  Higgins  2235,  2236,  2244,  (bry, 
WTs) ;  A.  Nelson  9812  (rm,  uc)  ;  W.  Web- 
er 9416  (rsa,  uc.  ut);  B.  Hartman  2782 
(bry.  rm). 

Habit.\t. —  Gravelly  and  sandy  soils 
and  talus  slopes  from  7,050  feet  elevation 
upward  to  tiniberline.  Commonly  in  open 
tundra  and  grassy  alpine  slopes  of  spruce, 
fir,  pine,  or  aspen  communities.  Some- 
times growing  as  a  weed  along  roadsides 
and  in  waste  field,  July  to  September. 

Distribution. —  Mostly  along  or  east 
of  the  Continental  Divide  in  central  and 
south  central  Colorado,  south  to  Colfax 
Co.,  New  Mexico  (Map  5). 


Fig.   16.     Phacelia  bakeri   (Brand)   Macbride. 
R.  Hartman  2782  (bry). 

This  taxon  was  misinterpreted  by  Brand 
(1913).  The  type  specimen  is  the  only 
material  cited  by  him  that  belongs  to 
bakeri  in  a  strict  sense.  Macbride  (1917) 
and  Voss  (1937)  present  additional  in- 
formation on  Brand's  treatment  of  this 
entity.  P.  bakeri  is  related  to  P.  gland- 
iilosa  Nutt.,  differing  in  the  lack  of 
excavations  on  the  ventral  surface  of  the 
seeds,  usually  greener  herbage,  later 
flowering  time,  and  distribution. 


148 


GREAT  BASIN  NATURALIST 


Vol.  35,  No.  2 


^: 


Fig.  17.  Dorsal  and  ventral  view  of  the 
seeds  of  P.  bakeri  (Brand)  Macbride.  C.  Baker 
549  (gh). 

8.  Pfiacelia  hombycina  Wooton  &  Slandley 
Fig.  18 

Phacelia  bombycina  Wooton  &  Standley.  Contr. 
U.S.  Natl.  Herb.  16:163.  1913.  Holotype: 
New  Mexico:  Carton  Co.:  on  gravelly  banks 
at  Mangas  Springs,  March  or  April  1880, 
H.  Rusby  276  (us!);  Isotypes  (ny,  ny,  uc, 
us).  Paratypes:  New  Mexico:  Grant  Co.: 
Bear  Mountains  near  Silver  City,  no  date, 
C.  Metcalfe  75  (ny,  pom,  uc,  us,  us). 
Phacelia  tenuipes  Wooton  &  Standley.  Contr. 
U.S.  Natl.  Herb.  16:163.  1913.  Holotype: 
Carizalillo  Spring,  17  April  1902,  E.  Mearns 
91    (us!). 

Annual  0.9-4.1  dm  tall;  stems  1 -sev- 
eral, often  branched  throughout,  erect  and 
very  brittle,  setose  to  puberulent  and 
glandular  at  least  in  the  inflorescence; 
basal  leaves  petiolate  (the  petiole  up  to 
5  cm  long),  oblong  to  nearly  orbicular, 
pinnatifid  into  oblong  to  ovate,  crenate 
lobes,  setose  and  often  glandular  on  both 
surfaces,  1.5-8  cm  long,  0.5-2.5  cm  wide, 
upper  reduced,  short  petiolate,  lobed;  in- 
florescence paniculate,  narrow,  with  a 
few  branches;  individual  cymes  with 
numerous,  crowded  flowers,  pedicels  short, 
stout,  setose  to  puberulent  and  glandular; 
sepals  oblong  to  elliptical,  up  to  3.5  mm 
long,  setose  to  glandular;  corolla  blue  to 
violet,  5-6  mm  long  and  wide,  campan- 
ulate,  lobes  pubescent;  stamens  exserted, 
anthers  yellow,  filaments  bluish  violet; 
style  exserted,  bifid  to  about  the  middle, 
pubescent  below  the  middle,  bluish  vio- 
let; capsule  2.5-2.7  mm  long  and  broad, 
globose,  pilose  and  glandular  especially 
at  the  apex;  mature  seeds  oblong  to  ellip- 
tical, cjTnbiform,  2.2-2.5  mm  long,  1-1.4 
mm    wide,    dark    brown,    ventral    surface 


Map  5.  Southern  Colorado  and  northern 
New  Mexico.  Distribution  of  P.  bakeri  (Brand) 
Macbride. 

pitted  and  divided  by  a  prominent  ridge, 
the  ridge  corrugated  on  one  side,  margins 
corrugated,  lighter  than  the  center,  dorsal 
surface  pitted  (Fig.  19).  Collections  89 
(9);  representative:  L.  Goodding  2230 
(rm,  uc);  W.  Cottam  10198  (uc) ;  D. 
Demaree  42048  (uc);  L.  Higgins  2877 
(bry,  wts);  H.  Ripley  and  R.  Barnebv 
4218  (rsa);  D.  Atwood  2195,  2241,  2256, 
2280  (bry)  ;  D.  Atwood  2250,  2253,  2255 

(bry,  CAS,  NY.  WTs). 

Habitat. —  Sandy,  gravelly,  or  lava 
sloj)es  and  mesas  from  an  elevation  of 
1,500  to  7,500  feet.  Commonly  found  in 
the  Larrea,  Prosopis,  and  other  Lower 
Sonoran  mixed  shrub  communities,  late 
March  to  late  May. 

Distribution. —  Sierra  Co.,  New  Mex- 
ico, southward  to  Chihuahua  and  Sonora. 
Mexico,  westward  through  southern  and 
central  Arizona  to  Yavapai  and  Coconino 
COS.   (Map  6). 

The  character  differences  that  Wooton 
and  Standley  (1913)  used  to  separate 
P.  tenuipes  and  P.  bombycina  var}'  de- 
pending  on    the   maturity    of   the  plants 


June  1975 


ATWOOD:    PHACELIA   CRENULATAE   GROUP 


149 


Fig.   18.     Phacelia  bombycina  Wooton  &  Standley.    D.  Atwood  2253   (bry). 


and  environmental  conditions.  Of  the  two 
taxa,  the  original  description  of  the  latter 
better  fits  the  entity  concerned.  For  these 
reasons  P.  tenuipes  is  placed  in  synonomy. 
The  holotype  data  indicate  that  P.  bom- 
bycina was  collected  in  March  and  April; 
however,  an  isotype  in  the  U.S.  herbarium 
Rives  the  exact  data  as  25  March  1880. 


This  entity  has  been  confused  with 
both  P.  intermedia  and  P.  crenulata.  It  is 
roacHly  distinguished  from  the  former  by 
the  exserted  stamens,  larger  blue  lavender 
corollas,  very  brittle  and  less  glandular 
stems,  and  smaller  seeds.  It  differs  from 
the  latter  by  its  brittle  stems  and  smaller, 
darker  seeds. 


150 


GREAT  BASIN  NATURALIST 


Vol.  35,  No.  2 


Fig.  19.  Dorsal  and  ventral  view  of  the 
seeds  of  P.  bombycina  Wooton  &  Standley.  D. 
Atwood  2255  (bry). 

9.  Phacelia  coerulea  Greene 
Fig.  20 

Phacelia  coerulea  Greene,  Bull.  Torr.  Bot.  Club 
8:122.  1881.  Lectotype:  bluffs  of  the  Gila 
(New  Mexico),  23  April  1881,  E.  Greene  s.n. 
(gh!). 

Phacelia  invenusta  Gray,  Proc.  Amer.  Acad. 
20:303.  1885.  Lectotype:  Arizona:  Pima 
Co.:  Sierra  Tucson,  12  April  1884,  C.  G 
Pringle  s.n.  (gh!);  duplicates  (us),  in  part 
Phacelia  intermedia  Wooton,  in  part.  Para 
types:  New  Mexico:  Socorro  Co.:  San  An 
tonio,  Quitman  Mountains,  14  March  1852 
C.  Wright  1579  (gh,  gh,  gh,  ny);  Texas 
El  Paso  Co.:    El  Paso,  March   1851,  Thurber 

11    (gh,  gh,  NY,  ny). 

Annual,  0.5-6  dm  tall;  stems  erect, 
branched  throughout,  reddish,  puberulent 
to  setose  and  sparsely  to  densely  stipitate- 
glandular,  leafy  throughout;  leaves  oblong 
to  ovate,  gradually  reduced  from  the  base 
upward,  upper  deeply  sinuate,  lower  pin- 
natifid,  dorsal  surface  with  setose  ap- 
pressed  hairs,  dorsal  surface  setose  to 
glandular,  0.6-8  cm  long,  0.3-2.5  cm  wide, 
petioles  from  5  cm  long  at  the  base  to 
nearly  sessile  at  apex,  margins  crenate; 
inflorescence  terminal,  commonly  loosely 
paniculate  or  cymose,  setose,  stlpitate- 
glandular  and  puberulent;  scorpioid  cymes 
compact  in  flower  but  loosening  in  fruit, 
1.5-7  cm  long;  flowers  nearly  sessile  in 
flower  to  1  mm  long  in  fruit;  corolla 
campanulate,  lobes  pale  mauve  to  blue 
(turning  white  in  fruit),  tube  yellowish, 
3-4  mm  long  and  broad,  glabrous;  sepals 
narrowly  oblanceolate  to  elliptical,  2.5-4 
mm  wide,  setose  to  brownish  stipitate- 
glandular,  3/4  as  long  as  the  corolla, 
shorter  or  sometimes  longer  than  the  cap- 


Map  6.  Southwestern  New  Mexico,  Arizona, 
and  adjacent  Sonora,  Mexico.  Distribution  of  P. 
bombycina  Wooton  and  Standley. 

sule;  stamens  mostly  included  to  slightly 
exserted,  anthers  yellow,  ovate,  filaments 
bluish;  style  included  to  slightly  exserted, 
equaling  the  stamens;  capsule  globose, 
2.5-3.5  mm  long,  1-1.8  nun  wide;  mature 
seeds  dark  brown,  ventral  surface  pitted 
and  divided  by  a  prominent  ridge,  the 
ridge  corrugated  on  one  side,  margins  usu- 
ally corrugated,  dorsal  surface  pitted,  0.3- 
0.4  mm  of  the  margin  slightly  elevated 
and  smoother  than  the  pitted  center  (Fig. 
21).  Collections:  112  (6) ;  representative: 
M.  Jones  s.n.  (pom);  W.  Wooton  s.n. 
(ny);  C.  Parry  934  (ny);  E.  Greene  s.n. 
(gh);  C.  Wright  1579  (gh,  gh,  gh,  ny)  ; 
D.  Atwood  2137,  2152,  2196a,  2573 
(bry);  D.  Atwood  2197,  2281  (bry,  gas, 
NY,  wTs);  L.  Higgins  3126,  3134,  2978, 
2999  (bry.  wts). 

Habitat. —  Gravelly  and  arid  calcar- 
eous hills  and  banks,  sandy-gravelly 
stream  beds,  and  rocky  ledges  from  2,000 
to  6.000  feet.  Commonly  associated  with 
the  paloverde  and  creosote  mixed  shrub 
communities.  Usually  locally  scattered, 
late  February  to  early  July. 


June  1975 


ATWOOD:    PHACELIA   CRENULATAE  GROUP 


151 


Fig.  20.     Phacelia  coerulea  Greene.    D.  Atwood  2196a    (dry). 


Distribution. —  Southern  Nevada  in 
Clark  Co.,  southeastern  Cahfornia  in  San 
Bernardino  Co.,  eastward  through  Mohave 
Co.    to    southern    Arizona    and    southern 


New  Mexico  from  Socorro  and  Lincoln 
COS.  to  EI  Paso,  Presidio,  and  Brewster 
COS.,  Texas,  and  adjacent  Mexico  in  the 
state  of  Chihuahua.    Onlv  one  collection 


152 


GREAT  BASIN  NATURALIST 


Vol.  35,  No.  2 


Fig.  21.  Dorsal  and  ventral  view  of  the 
seeds  of  P.  coerulea  Greene.  H.  Ripley  and  R. 
Barneby  3361    (c.as). 

is    knowTi    from    Nevada    and    California 
(Map  7). 

Greene  cited  no  material  on  which  he 
based  his  original  description  in  November 
1881.  Voss  (1937)  noted  this  fact  and  se- 
lected Greene's  collection  at  the  Gray 
Herbarium  as  the  type,  since  he  had  col- 
lected P.  coerulea  in  the  spring  (23  April 


Map  7.  Southern  Nevada,  southeastern  Cali- 
fornia, Arizona,  New  Me.xico,  western  Texas,  and 
adjacent  Chihuahua,  Mexico.  Distribution  of  P. 
coerulea  Greene. 


1881)  of  the  same  year.  However,  Howell 
(1943),  in  Sertulum  Greeneanum,  indi- 
cates that  a  part  of  the  type  collection 
chosen  by  Voss  is  present  in  the  Greene 
Herbarium  at  Notre  Dame  and  that  this 
specimen  should  be  the  type.  Since  Voss 
selected  the  specimen  at  the  Gray  Her- 
barium as  the  type,  it  should  probably 
stand  as  such  even  though  Greene's  orig- 
inal is  at  Notre  Dame. 

This  taxon  is  most  closely  related  to 
P.  bombycina  and  secondarily  to  P.  cren- 
ulata.  It  differs  from  these  species  in  its 
included  stamens  and  smaller  corollas. 
Jones  (1908)  contended  that  P.  coerulea 
and  P.  invcnusta  Gray  were  separate 
species.  The  author  believes,  as  did  Gray 
(1886)  and  Voss  (1937),  that  the  latter 
is  the  same  entity  as  the  former.  Howell 
(1941)  reported  P.  coerulea  from  San 
Bernardino  Co.,  California  (H.  Ripley  and 
R.  Barneby  3361  (cas,  rsa)  .  To  my  know- 
ledge this  is  the  only  collection  from 
California. 

No  type  was  selected  by  Gray  when  he 
described  P.  invenusta;  therefore  I  have 
selected  the  Pringle  collection  (gh)  as  the 
lectotype.  Duplicates  are  located  at  cas 
and  a  fragment  at  us. 


10.  Phacelia  congesta  Hooker 
Fig.  22 

Phacelia^  congesta  Hooker.   Bot.  Mag.  62: t.  3452. 
1835.    Holotype:   Texas:   Galveston  Co.:    Gal- 
veston Bay.  Drummond  303   (gh!). 
Phacelia    congesta    Hooker   var.    typica   Voss. 
Bull.    Torr.    Bot.    Club    64:133.      1937. 
Torr.  Bot.  Club  64:133.      1937. 
Phacelia  congesta  Hooker  var.  dissecta  Gray. 
Syn.   Fl.   11:1    SuppL,  415.     1886.   Holotype: 
Texas:   Dallas  Co.:   shaded  rocks.  Dallas,  May 
and   July   1880,  Reverchon  s.n.    (gh!);   dupli- 
cates  (C.-^S,  GH,  GH,  NY,  Ny). 

Phacelia    dissecta    (Gray)    Small.    Fl.    South- 
eastern U.S.   pp.   972,    1337.     1903. 
Phacelia  conferta  D.   Don.   Gen.   Syst.   Gard. 
4:397.      1837. 

Erect,  often  robust,  annual  plants,  1-10 
dm  tall;  stems  simple  or  diffusely 
branched  throughout,  with  midticellular 
stipitate  glands  and  puberulent  to  hispid 
unicellular  hairs;  leaves  oblong  in  out- 
line, pinnately  compound,  the  3  ter- 
minal lobes  usually  not  completely  lobed 
to  the  midrib,  often  somewhat  larger  than 
the  lower,  usually  petiolate  lobes,  1-12 
cm  long,  0.5-4  cm  wide,  strigose  and 
sometimes     glandular:     inflorescence     of 


June  1975 


ATWOOD:    PHACELIA   CRENULATAE   GROUP 


153 


J  '<    'fl§- 


terminal,  compound,  scorpioid  cymes,  the 
individual  cymes  1.5-15  cm  long,  pedicels 
2.5  mm  long  or  less,  pubescent;  sepals 
narrowly  linear,  3-4.5  mm  long,  0.5-0.7 
mm  wide,  setose  to  hispid  and  often  gland- 
ular; corolla  campanulate,  blue  (rarely 
white),  4-6  mm  long  and  wide,  the  lobes 
pubescent;  stamens  and  style  exserted, 
ca.  2-4  mm  long,  anthers  pale  yellow, 
filaments  purplish;  style  7-8  mm  long, 
bifid  3/4  its  length,  lower  1/4  pubescent; 
capsule  subglobose  to  oval,  2.3-3.6  mm 
long,  2.3-3  mm  wide,  puberulent  and 
often  glandular;  mature  seeds  usually  4, 
sometimes  1  or  2,  2.6-3.2  mm  long,  1.2- 
1.4  mm  wide,  elliptical  to  oblong,  brown, 
reticulate  to  scabrous,  ventral  surface  ex- 
cavated on  both  sides  of  the  ridge  (Fig. 
23).  Collections:  222  (21);  representa- 
tive: V.  Cory  28660  (gh);  R.  McVaugh 
7780  (uc);  E.  Palmer  33743  (ny,  us);  H. 
Ripley  and  R.  Barneby  11107  (c^s) ;  E. 
Tyler  s.n.  (us);  L.  Higgins  2671,  3162 
(bry);  D.  Atwood  2048a,  2049,  2063, 
2098,  2099,  2104-2107,  2111,  2117 
(bry). 

Habitat. —  Commonly  associated  with 
Prosopis,  Larrecu  Acacia^  and  Opuntia 
in  sandy  to  sandy  loam,  rocky  limestone, 
or  sandstone  flats  and  outcrops.  Along 
the  coast  of  southern  Texas  it  grows  on 
low  shoreline  dunes  and  is  usually  as- 
sociated with  Quercus,  February  to  Sep- 
tember, 300  to  7,000  feet. 

Distribution. —  Scattered  throughout 
most  of  central  and  southern  Texas,  west 
to  Eddv  and  Dona  Ana  cos.,  New  Mexico; 
northeastern  Mexico  and  Caddo  and  Com- 


Fig.  22.     Phacelia    congesta    Hooker. 
wood  2046  (bry). 


D.    At- 


Fig.  23.     Dorsal    and    ventral    view    of    the 
seeds    of    P.    congesta    Hooker.    D.    Atwood    2117 

( BRY ) . 


154 


GREAT  BASIN  NATURALIST 


Vol.  35,  No.  2 


manche  cos.,  Oklahoma.  Specimens  from 
Florida,  Massachusetts,  and  Sweden  are 
presumably  cultivated  (Map  8). 

Some  plants  from  Tamaulipas,  Zacate- 
cas,  and  adjacent  Nuevo  Leon,  Mexico, 
are  fall-flowering  and  differ  in  sufficient 
morphological  features  to  warrant  further 
investigation.  They  are  apparently  rhiz- 
omatous  perennials  and  possess  a  con- 
gested inflorescence  and  small,  pale  lav- 
ender flowers. 

This  taxon  varies  throughout  its  range 
in  pubescence  and  in  leaf  size  and  shape. 
The  number  of  seeds  per  capsule  was 
used  by  former  workers  as  an  important 
character  in  separating  var.  dissecta 
from  var.  congesta.  However,  the  author 
has  examined  capsules  from  the  type  ma- 
terial of  var.  dissecta  and  found  that 
they  possess  4  seeds.  Other  material  ex- 
amined varies  in  the  number  of  seeds  per 
capsule.  This  variation  probably  results 
from  environmental  conditions  and  is 
hardly  consistent  enough  to  warrant  tax- 
onomic  recognition. 

The  seeds  of  P.  congesta  are  dark  when 
immature  and  are  light  brown  upon 
reaching  maturity.  A  more  or  less  mottled 
pattern  can  be  observed  in  the  different 
stages  of  development. 


11.  Phacelia  constancei  Atwood 
Fig.  24 

Phacelia  constancei  Atwood.  Rhodora  74(800): 
451-454.  1972.  Holotype:  Arizona:  Coco- 
nino Co.:  1  mi  north  of  Fredonia,  27  Mav 
1968,    D.    Atwood,     1385a     (bry!).     Isotypes 

(aRIZ,    bry,    CAS,    NY,    UC,    Us). 

Erect  biennial  herb,  1.5-4.3  dm  tall, 
leafy  throughout;  stems  stout,  simple  or 
branched  throughout,  reddish,  from  hir- 
sutulous  to  hirsute,  and  finely  glandular; 
leaves  mostly  petiolate,  1-10  cm  long,  0.3- 
1.5  cm  wide,  revolute,  from  undulate  to 
pinnatifid,  linear  to  lanceolate,  upper  sur- 
face hirsutulous  with  scattered  glandular 
hairs;  inflorescence  of  compound  scorpioid 
cymes;  pedicels  to  1  mm  long;  sepals  el- 
liptic to  oblanceolate,  3-4  mm  long,  hir- 
sutulous to  hirsute  and  stipitate-glandular; 
corolla  tubular,  whitish,  5-6  mm  long;  sta- 
mens exserted  3-4  mm;  style  bifid,  lower 
1/3  pubescent,  exserted  3-4  mm  longer 
than  the  stamens;  capsule  subglobose, 
glandular,    and     hirsutulous     throughout. 


Map  8.  Texas,  adjacent  New  Mexico,  Okla- 
homa, and  Mexico.  Distribution  of  P.  congesta 
Hooker. 


shorter  than  the  sepals,  mature  seeds  4, 
black,  2.5-2.8  mm  long,  1-1.2  mm  wide,  el- 
liptic, the  margins  corrugated,  ventral 
surface  finely  pitted,  excavated,  and  di- 
vided by  a  prominent  ridge,  the  ridge  cor- 
rugated on  one  side,  the  dorsal  surface 
finely  pitted.  Collections:  12  (10);  rep- 
resentative: H.  Ripley  and  R.  Barneby 
4836  (cAs,  rsa);  D.  Atwood  1529  (bry, 
GH,  uc);  D.  Atwood  1785  (ariz,  bry,  cas, 
NY,  uc,  us,  uTc);  D.  Atwood  1792b  (bry, 

CAS,    GH,    NY,    POM.    RM.    US). 

Habitat. —  Alkaline  clay  bluffs  and 
flats  of  the  Moenkopi  formation,  5,500 
feet.  Late  May  to  early  August. 

Distribution. —  Mohave  Co.,  Arizona, 
and  Kane  Co.,  Utah  (Map  9). 

Taxonomically  P.  constancei  appears  to 
be  most  closely  related  to  P.  palmeri  Torr. 
ex  Wats,  but  is  distinguished  from  that 
species  by  the  smaller  growth  form,  nar- 
rower and  more  revolute  leaves,  the  red- 
dish stems  with  shorter  and  fewer  hairs, 
and  the  leafier  lateral  inflorescence 
branches. 


June  1975 


ATWOOD:    PHACELIA   CRENULATAE   GROUP 


155 


Fig.  24.     Phacelia  constancei  Atwood.  D.  Atwood   1 


835a   (bry). 


156 


GREAT  BASIN  NATURALIST 


Vol.  35,  No.  2 


Map  9.     Southern    Utali    and    northern    Ari- 
zona.   Distribution  of  P.  constancci  Atwood. 


12.  Phacelia  corrugata  A.  Nelson 
Fig.  25 

Phacelia  corrugata  A.  Nelson,  Bot.  Gaz.  34:26. 
1902.  Holotype:  Colorado:  Garfield  Co.:  Rifle, 
23  June  1900.  G.  E.  Osterhout  2129  (rm!); 
Isotypes  (ny,  rm).  Paratypes:  Colorado: 
Mesa  Co.:  Palisades,  14  May  1898,  C.  S. 
Crandall  4174  (rm,  rm,  us). 
Phacelia  crenulata  Torr.  ex  Wats.  var.  cor- 
rugata (A.  Nels.)  Brand,  Das  Pflanzenreich 
IV,  251:79.     1913. 

Phacelia  orbicularis  Rydberg,  Bull.  Torr. 
Bot.  Club  40:479.  1913.  Holotype:  Utah: 
Wayne  Co.:  Marvine  Laccolite.  22  July  1894, 
M,  Jones  4663   (us!). 

Plants  annual  or  winter  annuals,  0.5- 
4.3  dm  tall;  stems  greenish,  from  puber- 
ulent  to  finely  stipitate-glandular  and 
sometimes  with  a  few  longer  hairs  inter- 
mixed; leaves  1-10  cm  long,  setose  to  stri- 
gose  and  stipitate-glandular;  sepals  4-5.5 
mm  long;  corolla  campanulate,  deep  blue, 
6  mm  long  or  more;  stamens  and  style 
exserted  over  3  mm,  filaments  and  style 
blue;  capsule  elliptic,  3.8-4.5  mm  lon^; 
mature  seeds  oblong  to  elliptic,  light 
brown,  3.1-4  mm  long,  1.3-1.7  mm  wide, 
pitted,  the  ventral  surface  corrugated  on 
the  margins  and  one  side  of  the  ridge  (Fig. 
26).  Collections:  216  (44);  representa- 
tive: R.  Barneby  13033  (cas,  ny)  :  C.  Par- 
ry s.n.  (cas,  uc)  ;  A.  Eastwood  and  J. 
Howell  9358  (cas,  gh,  utc);  L.  Higgins 
3304,  3303,  3305  (bry,  wts);  D.  Atwood 


1314,  1489,  1856,  2539,  2523,  2583,  2581, 
2618  (bry). 

Habitat. —  This  species  grows  in  a 
large  number  of  habitats  from  dry,  grav- 
elly hillsides  and  flats,  sandy  soil,  and 
red  shaly  clay  to  heavy  clay  soils.  It 
grows  with  Atriplex,  Sarcobatus^  Ephedra, 
Coleogyne  or  grass,  from  3,500  to  7,000 
feet  elevation.  Late  April  to  mid- July. 

Distribution. —  Eastern  Nevada,  east- 
ward to  Colorado  from  Garfield  and  Gun- 


Fig.  25.     Phacelia   corrugata   A.    Nelson.    D. 
Atvvood  2457   (bry). 


1975 


ATWOOD:    PHACELIA   CRENULATAE  GROUP 


157 


^B 

.Lj 

!  Fig.  26.     Dorsal    and    ventral    viev/    of    the 

I   seeds  of  P.  corrugata  A.  Nelson.  D.  Atwood  2583 
i    (bry). 

nison  cos.,  southward  through  Ouray  and 
Montezuma  cos.  to  northwestern  New 
Mexico  and  northern  Arizona  (Map  10). 
This  species  is  related  to  P.  crenulata 
and  apparently  intergrades  with  it  in 
eastern  Nevada  and  western  Utah.  In 
general,  however,  it  is  easily  distinguished 
from  the  latter  by  its  greenish  yellow 
stems,  finer  pubescence,  and  lighter,  nar- 
rower seeds. 


13.  Phacelia  coulter i  Greenman 

Phacelia  coulteri  Greenman.  Proc.  Amer.  Acad. 
41:241.  1904.  Holotype:  Mexico:  state  of 
Hidalgo:  fields  about  Buena  Vista  Station. 
4  August  1904,  C.  Pringle  8988  (gh!);  Iso- 
types  (cAS,  MEXU,  ny,  ny,  pom,  uc,  us,  us). 
Paratypes:  Mexico:  Vera  Cruz:  Real  del 
Monte,  T.  Coulter,  no  date  921  (?). 
Phacelia  neoniexicana  Thurber  ex  Torr.  var. 
coulteri  (Greenman)  Brand.  Das  Pflanzen- 
reich  IV.  251:84.  1913, 
Phacelia  grandulosa  Hem.  Biol.  Cent.  Amer. 
Bot.  2:359.     1882. 

Stout  annual,  3.4-5.4  dm  tall;  stems 
solitary  or  branched,  brownish  to  red- 
dish, hirsute  and  provided  with  a  softer 
indument;  leaves  pinnately  compound, 
finely  dissected,  0.2-1  dm  long,  3-4.5  cm 
wide,  hispid,  with  small  dark  glands,  mid- 
rib and  some  of  the  lateral  veins  prom- 
inent ventrally  and  therefore  producing  a 
furrowlike  appearance  dorsally,  the  mar- 
gins of  the  pinnae  thick  ventrally,  lower 
leaves  petiolate,  the  pedicels  gradually  re- 
duced upward;  inflorescence  of  terminal 
compound  scorpioid  cymes,  hispid,  with 
brownish  glandular  hairs,  the  cymes  con- 
gested, up  to  7  cm  long,  pedicels  1  mm 
long  or  less;  sepals  linear  to  oblanceolate, 


Map  10.  New  Mexico, 
Utah,  and  adjacent  Nevada. 
corrugata  A.  Nelson. 


Arizona,    Colorado, 
Distribution    of    P. 


3.3-3.6  mm  long,  0.5-1.3  mm  wide,  hir- 
sute and  brownish  glandular;  corolla 
campanulate,  bluish  purple,  4-5  mm  long 
and  broad,  finely  pubescent  petals  barely 
fimbriate;  stamens  and  style  exserted  1- 
3.5  mm,  the  anthers  yellow,  filaments 
bluish;  style  ca.  6  mm  long,  bifid  1/2 
its  length,  lower  1/2  glandular  and  puber- 
ulent;  capsule  oval,  2.4-3  mm  long  and 
broad,  glandular  and  puberulent;  mature 
seeds  4,  oblong,  brown,  1.6-1.9  mm  long, 
0.9-1  mm  wide,  pitted  over  the  entire  sur- 
face, the  ridge  level  with  the  margins  and 
deeply  excavated  on  both  sides  (Fig.  27). 
Collections:  3  (0);  representative:  C. 
Pringle  8988  (cas,  gh,  mexu,  ny,  pom, 
uc,  us);  G.  Rzedowski  16995  (mexu); 
Tinsley  et  al.  8  (uc). 

Habitat. —  A  weed  of  fields  and  road- 
sides as  well  as  meadows  and  well-vege- 
tated areas  in  the  mountains,  7,350  to 
8,500  feet.  July  to  August. 

Distribution. —  Known  only  from  the 
states  of  Hidalgo  and  Zacatecas,  Mexico. 

Greenman  did  not  select  a  holotype  but 
Pringle  8988  was  cited  and  is  probably 
the  type  material. 


158 


GREAT  BASIN  NATURALIST 


Vol.  35,  No.  2 


foliis  angustioribus,  corollis  violaceis,  stam- 
inibus  et  stylis  excertis  5-6  mm,  capsulis 
subglobosis  3-4  mm  longis  2-3  mm  latis, 
seminibus  ellipticis  2-3  mm  longis,  1-1.75 
mm  latis. 

Annuals,  1.4-3.9  dm  tall;  stems  simple 
to  more  commonly  branched  from  the 
base,  finely  and  densely  stipitate-glandu- 
lar,  setose  and  puberulent,  reddish  purple 
especially  below;  leaves  1.1-9  cm  long,  0.2- 
1  cm  wide  (up  to  2.2  mm  wide),  stipitate- 
glandular  and  setose,  petiolate;  pedicels  1 
mm  long;  corolla  campanulate,  violet,  7-10 
mm  long  and  broad;  stamens  and  style 
exserted  5-6  mm;  capsule  subglobose,  3-4 

Fig.  27.  Dorsal  and  ventral  view  of  the 
seeds  of  P.  coulteri  Greenman.  C.  Pringle  8988 
(mexu). 

Phacelia  coulteri  is  related  to  P.  alba 
but  differs  from  that  taxon  in  having 
smaller  seeds  (2  mm  long)  and  larger 
campanulate  corollas  (4.5  mm  long).  The 
stamens  and  style  are  exserted  up  to  3.5 
mm,  the  corolla  lobes  are  only  shallowly 
erose,  and  the  leaves  are  more  finely  dis- 
sected. 

14.  Phacelia  crenulata  Torr.  in  Wats. 

Plants  annual,  0.25-8.3  dm  tall;  stems 
1 -several,  simple  to  much  branched,  stip- 
itate-glandular,  setose  or  puberulent,  red- 
dish purple  to  green;  leaves  0.4-1.2  dm 
long,  0.2-4.0  cm  wide,  strigose  to  setose 
and  stipitate-glandular,  sessile  to  petiolate; 
inflorescence  of  compound  scorpioid 
cymes,  setose  and  stipitate-glandular,  the 
pedicels  0.5-2  mm  long;  sepals  cleft  to  near 
the  base,  the  lobes  elliptic  to  oblanceolate, 
3-3.5  mm  long,  1-1.4  mm  wide,  glan- 
dular and  setose;  corolla  campanulate  to 
rotate-campanulate,  violet,  blue,  or  pur- 
ple, pubescent,  4.5-10  mm  long  and  broad; 
stamens  and  style  exserted  5.5-11  mm; 
style  bifid  3/4  its  length,  glandular  be- 
low; capsule  globose  to  subglobose,  2.6-4.1 
mm  long,  2-3  mm  wide,  puberulent  and 
glandular;  seeds  4,  elliptic  to  oblong,  2-3.6 
mm  long,  1-2  mm  wide,  the  dorsal  surface 
with  a  dark  center  and  lighter  margins, 
the  ventral  surface  corrugated. 

14a.  Phacelia  crenulata  vnr.  angustifolia 

At  wood,  var.  no  v. 

Fig.  28 

Phacelia    crenulatae    Torr.    in    Wats.    var.    crenu-  Fig.  28.     Phacelia   crenulata   Ton-,    in   Wats. 

latae  affinis   sed   brevioribus   et   ramosioribus.       var.   angustijolia  Atwood.   Atwood   2523    (dry). 


June  197: 


ATWOOD:    PHACELIA   CRENULATAE  GROUP 


159 


mm  long,  2-3  mm  wide;  seeds  4,  elliptic, 
2-3  mm  long,   1-1.75  mm  wide. 

Type. —  Arizona;  Coconino  Co.:  Small 
mesas  just  north  of  Wupatki  National 
Monument  Headquarters,  sandy  soil 
covered  with  volcanic  ash,  18  May  1970, 
N.D.  Atwood  2597  (Holot>ioe:  bry;  Iso- 
types:   to  be  distributed). 

Additional  materials  examined:  Ar- 
izona: Coconino  Co.:  D.  Atwood  1784, 
2602,  2600,  2606,  2604,  3650,  4559,  4555, 
2597  (bry);  L.  Higgins  5187,  5396  (bry); 
L.  Williams  5993  (bry);  L.  Cureton  45 
(bry).  Utah:  Beaver  Co.:  D.  Atwood  and 
L.  Higgins  3839  (bry).  Garfield  Co.:  D. 
Atwood  1356  (bry).  Kane  Co.:  D.  Atwood 
1532B,  3603,  3612  (bry);  L.  Higgins  and 
D.  Atwood  5247  (bry);  R.  Allen  211,  243 
(bry). 

Habitat. —  Sandy,  clay,  or  rocky 
ground  in  the  can3'ons  and  benches  be- 
low 5,000  feet  elevation,  April- June. 

Distribution. —  Coconino  Co.,  Arizona, 
north  to  Kane,  Garfield,  and  Beaver  cos., 
Utah   (Map  11). 


Map  1 1 .  Utah,  Nevada.  Arizona,  and  Cali- 
fornia. Distribution  of  P.  crenulata  Torr.  in  Wats, 
var.  crenulata  #;  var.  augustifolia  *. 


14b.  Phacelia  crenulata  var.  crenulata 

Phacelia  crenulata  Torr.  in  Wats.,  Bot.  King 
Exped.  251.  1871.  Holotype:  Nevada:  Per- 
shing Co.:  Trinity  Mountains,  May  1868, 
S.  Watson  873  (ny!);  duplicates  (gh,  us). 
Phacelia  crenulata  Torr  in  Wats.  var.  vul- 
garis Brand,  Das  Pflanzenreich  IV.  251:78. 
1913. 

Phacelia  crenulata  Torr.  in  Wats.  var. 
funerea  Voss  in  Munz.  Man.  So.  Calif.  Bot. 
409,  600.  1935.  Holotype:  California:  Mono 
Co.:  Black  Canyon,  White  Mountains,  12 
May    1930,   V.    Duran   561    (pom!);    Isotypes 

(b,  CAS,  GH,  NY,  UC,  Us). 

Plants  annual,  0.25-8.3  dm  tall;  stems 
1 -several,  simple  or  branched,  puberulent, 
pilose,  setose  and  stipitate-glandular,  red- 
dish purple  to  green;  leaves  0.4-1.2  dm 
long,  0.5-4  cm  wide,  strigose  to  setose  and 
stipitate-glandular,  sessile  to  petiolate;  in- 
florescence of  compound  scorpioid  cymes, 
puberulent  to  setose  and  stipitate-gland- 
ular, the  pedicels  0.5-2  mm  long;  sepals 
elliptic  to  oblanceolate,  3-5.3  mm  long, 
1-1.4  mm  wide,  setose  and  stipitate-gland- 
ular; corolla  campanulate  to  rotate-cam- 
panulate,  blue,  pale  purple,  or  violet,  pub- 
escent, 4.5-7  mm  long  and  broad;  stamens 
and  style  exserted  5.5-11  mm;  style  bifid 
3/4  its  length,  glandular  pubescent  below; 


capsule  globose  to  subglobose,  2.6-4.1  mm 
long,  2.3-3.2  mm  wide,  puberulent  and 
glandular;  seeds  4,  elliptic  to  oblong,  2.8- 
3.6  mm  long,  1.2-2  mm  wide,  the  dorsal 
surface  with  a  dark  center  and  light  mar- 
gins, the  ventral  surface  corrugated  (Fig. 
29).  Collections:  124  (13);  representa- 
tive: Lemmon  s.n.  (us);  C.  Purpus  5976 
(uc);  J.  Howell  26588  (cas);  C.  Purpus 
s.n.    (uc);   D.  Atwood  2597,  2600,  2623 


Fig.  29.  Dorsal  and  ventral  view  of  the 
seeds  of  P.  crenulata  Torr.  in  Wats.  var.  crenu- 
lata. D.  Atwood  2623  (bry). 


160 


GREAT  BASIN  NATURALIST 


Vol.  35,  No.  2 


(bry);   p.   Munz    14790    (cas,   pom);   A. 
Nelson  3433  (rm). 

Habitat. —  Rock  slides,  limestone  talus, 
lava  flows,  gravelly  and  sandy  soil  of  the 
foothills  and  canyons,  from  4,600  to  8,000 
feet  elevation.  Late  February  to  early 
July. 

Distribution. —  Nevada,  eastward  to 
western  and  southern  Utah  (except  Wash- 
ington Co.),  and  southward  to  Mohave 
and  Coconino  cos.,  Arizona,  and  eastern 
California  from  Nevada  Co.  southward 
to  San  Bernardino  Co.  (Map  11). 

Phacelia  crenulata  was  described  from 
immature  specimens  (lacking  seeds)  in 
the  northern  part  of  its  range  in  north- 
western Nevada.  This  has  resulted  in 
some  confusion  as  to  its  relationship  to 
such  other  closely  allied  species  as 
P.  corrugata  A.  Nels.  and  P.  ambigua 
Jones.  Examination  of  specimens  from 
Pershing  Co.,  Nevada,  as  well  as  sur- 
rounding counties,  reveals  that  P.  cren- 
ulata is  quite  distinct  from  both  P.  cor- 
rugata and  P.  ambigua.  However,  there 
is  some  overlap  in  morphological  charac- 
ters where  the  species  grow  together.  For 
the  most  part  P.  crenulata  differs  from 
P.  ambigua  in  its  reddish,  usually  simple 
and  only  slightly  setose  stems,  and  its 
distinctive  seed  characters.  P.  corrugata 
is  easily  distinguished  by  its  yellowish 
green  stems,  uniformly  fine-glandular 
pubescence,  and  distinctive  seed. 

The  type  of  var.  funera  Voss  appears 
to  be  rnorphologically  the  same  as  the 
type  of  P.  crenulata.  The  author  has  been 
unable  to  study  this  complex  in  the  field; 
when  field  observations  and  greenhouse 
studies  have  been  accomplished,  a  change 
in  taxonomic  status  may  be  necessary. 
However,  until  such  studies  are  carried 
out,  the  present  arrangement  seems  the 
most  natural. 

15.  Phacelia  denticulata  Osterhout 
Fig.  30 

Phacelia  denticulata  Osterhout.  Torrcya  16:70. 
1916.  Holotypc:  Colorado:  Larimer  Co.:  Tlie 
Glades,  Owl  Canyon,  between  Fort  Collins 
and  Livermore.  18  June  1915,  G.  Osterhout 
5233  (rm!);  Isotypes  (ny.  rm.  rm,  rm). 
Phacelia  glandiilosa  Nutt.  ssp.  eu-glandulosn 
Brand  var.  australis  Brand,  Das  Pflanzenreich 
IV.  251:82-83.  1913,  in  part.  Lectotype: 
Colorado:  El  Paso  Co.:  Manitou.  15  July 
1903.  F.E.  and  F.S.  Clements  47.1,  in  part 
(us!);   duplicate    (gh). 


Phacelia  neomexicana  Thurber  ex  Torr.  var. 
microphylla  Brand,  Das  Pflanzenreich  IV, 
251:84.  1913.  Lectotype:  Colorado:  Can- 
yon, 18  July  1878,  central  Colorado,  T. 
Brandegee  s.n.    (gh). 

Annual  plants,  0.5-5.4  dm  tall;  stems 
erect,  simple  or  sometimes  branched,  se- 
tose and  stipitate-glandular;  leaves  oblong 
to  oblanceolate  in  outhne,  pinnately  cleft 
or  divided,  1-7.5  cm  long,  0.5-4.5  cm  wide, 


Fig.  30.     Phacelia    denticulata    Osterhout.    J. 
Ewan   18154   (uc). 


June  1975 


ATWOOI):     PHACELIA   CRENULATAE   GROUP 


161 


strigose  and  stipitate-glandular;  inflores- 
cence of  terminal  scorpioid  c}Tiies,  the 
longer  cymes  becoming  10  cm  long  in 
fruit;  sepals  narrowly  linear  to  oblanceo- 
late,  2.5  mm  long  in  flower,  5-6  mm  long 
in  fruit,  0.8-0.9  mm  wide,  setose  and  stip- 
itate-glandular; corolla  tubular,  light  blue, 
3.5-4.5  mm  long,  2.3  mm  wide,  the  lobes 
short,  denticulate;  stamens  included; 
style  included.  l)ifid,  glandular  at  base; 
capsule  ovoid,  5  mm  long,  3-3.5  mm  wide, 
]:)ilose  and  glandular;  mature  seeds  4,  el- 
liptical to  oblong,  brown,  4  mm  long,  1.7 
mm  wide,  ventral  surface  slightly  exca- 
vated on  each  side  of  the  curved  ridge, 
alveolate,  dorsal  surface  alveolate  (Fig. 
31).  Collections:  61  (7);  representative: 
H.  Ripley  and  R.  Barneby  7525  (cas, 
ny);  W.^  Weber  5974  (c.\s);  C.  Shear 
3306  (ny);  a.  Nelson  1361  (rm);  D.  At- 
wood  1941,  1946,  1959,  1973   (bry). 

Habit.at. —  Gravelly,  sandy,  or  clay 
banks,  draws  and  flats  of  the  prairie  to 
higher  mountain  slopes  from  6,800  to 
9,700  feet.  Commonly  as  understor}'  of 
Quercus^  Cercocarpus.  Artemisia,  and  Fi- 
nns edulis.  Frequently  associated  with 
Populus  trcmuloidcs  and  Pseudotsuga  at 
the  higher  elevations,  .Tune  to  September. 


•  ' 

^  •  \     •  i 

;            ^\             •  i 

\    Y'  \ 

1 
1 

Map  12.     Colorado    and    southern   Wyoming. 
Distribution  of  P.  dendculata  Osterhout. 


Fig.  31.  Dorsal  and  ventral  view  of  the 
seeds  of  P.  dendculata  Osterhout.  D.  Atwood  1973 
(bry). 

Distribution. —  Albany  and  Laramie 
COS.,  Wyoming,  southward  through  cen- 
tral Colorado  (Map  12). 

Phacelia  dendculata  is  most  closely  re- 
lated to  P.  neomexicana,  from  which  it  dif- 
feres  in  having  included  stamens,  a  denser 
glandular  pubescence,  larger  seeds,  and 
a  more  robust,  erect  habit. 

The  holotype  of  P.  neomexicana  var. 
?nicrop/iylla  Brand  was  deposited  in  the 
Berlin  Herbarium  (Brand  1913).  The 
author  has  seen  the  entire  collection  of  the 
Crenulatae  group  at  Berlin.  Apparently 
most  of  their  collection  was  destroyed 
during  the  war.  This  has  made  it  neces- 
sary to  select  a  lectotype  for  var.  micro- 
phylla;  the  author  has  chosen  the  Brand- 
egee  collection  at  GH  as  the  lectotype. 
Brand  (1913)  failed  to  select  a  holotype 
for  P.  neomexicana  var.  australis.  Most 
of  the  material  cited  by  him  belongs  to 
P.  bakeri  (Pur]uis  838"  Baker  549,  and 
Clements  47.1,  in  part).  The  Clements 
collection  consists  of  several  sheets,  most 
of  which  are  P.  bakeri.  However,  the 
Clements  collection  at  the  U.S.  National 
Herbarium  and  Gra}^  Herbarium  includes 
the  only  sheets  which  represent  var. 
australis.  I  designate  the  specimen  at  the 
U.S.  National  Herbarium  as  the  lectot>T)e. 

16.  Phacelia  formosula  Osterhout 
Fig.  32 

Phacelia  formosula  Osterhout.  Bull.  Torr.  Bot. 
Club  46:54.  1919.  Holotype:  Colorado: 
Jackson  Co.:  North  Park  near  Waldren. 
along  the  road  descending  to  Michigan  Creek, 
6  August  1918,  G.  Osterhout  5794  (rm!); 
Isotypes:   (rm,  rm,  rm). 


162 


GREAT  BASIN  NATURALIST 


Vol.  35,  No.  2    f 


-'f 


Fig.  32.     Phacelia    formosula    Osterhout.    D. 
Atvvood   1971    (bry). 

Annual,  1.5-2.2  dm  high,  up  to  3  dm 
broad;  stems  single  or  branched  through- 
out (especially  at  base),  glandular  and 
hirsute,  somewhat  grayish;  leaves  lanceo- 
late or  elliptical,  3-7  cm  long  (up  to  4.5), 
1-3  cm  wide,  pinnately  divided,  strigose, 
hirsute  and  glandular;  inflorescence  of 
compound  scorpioid  cymes,  cymes  up  to 
5  cm  long  in  fruit,  and  more  densely 
glandular  than  the  stems;  sepals  3.2-3.8 
mm  long,  0.5-0.8  mm  wide,  glandular  and 
hirsute;  corolla  campanulate,  violet,  6  mm 
long,  6  mm  wide,  slightly  glandular  and 
pilose;  stamens  and  style  long  exserted; 
style  ca.  2  mm  longer  than  the  stamens 
and    puberulcnt    throughout;    capsule   ob- 


long to  oval,  3.5  mm  long,  glandular  and 
hirsute;  mature  seeds  4,  oblong,  dark 
brown,  2.5-3  mm  long,  1.2-1.4  mm  wide, 
excavated  ventrally  on  each  side  of  the 
ridge,  pitted,  dorsal  surface  pitted,  margins 
rounded  and  smooth.  Collections:  7  (2); 
representative:  D.  Keck  889  (cas,  uc); 
H.  Ripley  and  R.  Barneby  9008  (cas); 
D.  Atwood  1977,  1977a  (bry). 

Habitat. —  Loose  sandy  soil  of  sand- 
stone bluffs  at  an  elevation  of  8,300  feet. 
Associated  with  Artemisia  and  Tetrady- 
mia,  July  to  August. 

Distribution. —  Apparently  confined 
to  Jackson  Co.,  Colorado. 

This  species  appears  to  be  most  closely 
related  to  P.  glandulosa  Nutt.  but  can  be 
distinguished  from  that  species  by  its  usu- 
ally much-branched,  erect  to  spreading 
habit,  less  exserted  stamens  and  style, 
darker  seeds,  narrower  calyx  lobes,  and 
more  pubescent  style.  P.  glandulosa  is  a 
somewhat  variable  species,  and  P.  for- 
mosula may,  perhaps,  best  be  treated  as  a 
variety  of  it;  however,  additional  mate- 
rials and  field  work  are  necessary  before 
this  suggestion  can  be  confirmed. 

17.  Phacelia  glandulosa  Nutt. 
Fig.  33 

Phacelia  glandulosa   Nutt..   Journ.   Acad.   Phil.  n. 
ser.  1:160.     1848.     Holotype:  Wyoming:  Lin- 
coln Co.:   about  Ham's  Fork,  Colorado,  of  the 
West,    July,    T.    Nuttall    93    (gh?). 
Eutoca  glandulosa  (Nutt.)  Hook.,  Kew  Journ. 
Bot.    3:293.      1851.      Wyoming:     Sweetwater 
Co^:    Greenriver.    31    May    1897.    A.    Nelson 
3050   (rm!);   Isotypes   (gh,  ny,  rm). 
Phacelia   glandulosa   Nutt.    subsp.    eu-glandu- 
losa  Brand  var.  deserla  Brand. 
Das  Pflanzenreich  IV,  251.82.      1913. 

Plants  annual  or  possibly  biennial,  0.6- 
3.6  dm  tall;  stems  simple  or  branched, 
erect,  densely  stipitate-glandular  and  hir- 
sute; leaves  lanceolate  to  oblong  in  out- 
line, pinnatifid,  1-7  cm  long,  0.5-3  mm 
wide,  glandular  and  densely  hirsute,  the 
lower  petiolate,  the  upper  subsessile;  in- 
florescence of  congested  terminal  com- 
pound scorpioid  cymes,  stipitate-glan- 
dular and  hirsute,  1-1.5  mm  long,  cymes 
elongating  to  6.5  cm  in  fruit;  sepals  el- 
liptical to  oblanceolate,  3-4  mm  long,  1.2- 
1.4  mm  wide;  corolla  campanulate,  pur- 
]ile  to  bluish,  5-7  mm  long  and  broad, 
the  lobes  pubescent  and  often  more  or  less 
crenate;    stamens   and   style   exserted   5-9 


June  1975 


ATWOOD:    PHACELIA   CRENULATAE  GROUP 


163 


.::-^ 


Fig.  33.     Phacelia    glandulosa    Nutt.    Hitch- 
cock 10804  (uc). 

min,  the  style  bifid  3/4  its  length,  the 
lower  1/4  pubescent;  capsule  subglobose, 
3.5-4  mm  long,  3.2-3.3  mm  wide,  glan- 
dular and  setose;  mature  seeds  elliptic  to 
oblong,  reddish  brown,  2.4-3.3  mm  long, 
1.1-1.4  mm  wide,  pitted,  the  ventral  sur- 
face excavated  on  both  sides  of  the  ridge 
(Fig.  34).  Collections:  25  (1);  representa- 
tive: E.  Graham  9774  (cas,  us)  ;  R.  Davis 
585  (us);  H.  Ripley  and  R.  Barneby 
8826  (cas,  ny);  S.  Watson  281  (gh,  us); 
A.  Rudvalis  70  (bry);  H.  Fitch  s.n.  (cas, 
NY,  pom);  a.  Williams  s.n.  (ny). 


Fig.  34.     Dorsal    and    ventral    view    of    the 
seeds  of  P.  glandulosa  Nuttall. 

Habitat. —  Rock  slides,  sandy  talus 
slopes,  and  clay  knolls  from  5,000  to  7,050 
feet.  Mid-June  to  early  August. 

Distribution. —  West  of  the  Continen- 
tal Divide  in  Rio  Blanco  Co.,  Colorado, 
northward  to  southwestern  and  western 
Wyoming,  southwestern  Montana,  and 
central  Idaho  in  Lemhi  and  Custer  cos. 
(Map  13). 

For  a  discussion  of  this  taxon  see 
P.  formosula. 

18.  Phacelia  howcUiana  Atwood 
Fig.  35 

Phacelia  howelliana  Atwood.  Rhodora  74(800): 
456-462.  1972.  Holotype:  Utah:  San  Juan 
Co.:  ca.  0.4  mi  north  of  Bluff  on  Utah  high- 
way 163,  in  mouth  of  canyon,  13  May  1970, 
D.   Atwood  2454    (bry);   Isotypes    (ariz,  asc, 

B,    BRY,    CAS,    GH,    NY,    UC,    US,    UTc). 

Plants  annual,  0.9-2.3  dm  tall;  stems 
mostly  branched  and  leafy  toward  the 
base,  glandular  and  hirsute;  leaves 
broadly  oblong  to  oval,  2.0-6.0  cm  long, 
1.0-2.5  cm  wide,  irregularly  crenate  to 
lobed,  strigose  and  slightly  glandular,  the 
petiole  up  to  5  cm  long;  inflorescence  of 
compound  scorpioid  cymes;  pedicels  up 
to  2  mm  long;  sepals  linear  to  narrowly 
oblanceolate,  3.5-4.5  mm  long,  1.0-1.2 
mm  wide,  glandular  and  hirsute;  corolla 
5-6  mm  long,  6-7  mm  wide,  rotate  to 
funnelform,  the  lobes  light  violet  to  blue, 
the  tube  white;  stamens  and  style  ex- 
serted  3-4  mm,  style  shorter  than  the 
stamens,  bifid  3/4  its  length,  lower  1/4 
pubescent;  capsule  oblong  to  subglobose, 
glandular  and  hirsutulous,  especially  to- 
ward the  apex;  seeds  4,  brown,  3.2-4  mm 


164 


GREAT  BASIN  NATURALIST 


Vol.  35,  No.  2 


Fig.  35.     Phacelia  howelliatia  Atvvood.    D.   Atwood  2454   (bry). 


long,  1.4-1.8  mm  wide,  elliptical,  the  mar- 
gins corrugated,  involute  to  flattened, 
ventral  surface  pitted,  excavated,  and  di- 
vided by  a  ])rominent  ridge,  the  ridge 
sometimes  curved  to  one  side  and  barely 
corrugated,  dorsal  surface  reddish  brown, 
smooth  and  surrounded  by  a  lighter  mar- 
gin (Fig.  36).  Collections:  16  (3):  rep- 
resentative: S.  Welsh,  D.  Atwood  and  G. 
Moore  9957  (bry);  A.  Eastwood  s.n. 
(ny);  B.  Harrison  11244  (bry);  C.  Han- 


sen  101    (bry);  M.  .lones  s.n.    (pom);  D. 

Atwood      2511       (ARIZ.      ASC,      B.      BRY,      CAS- 
DIXTE.    Gil.    JEPS.    NY.    POM,    RM,    RSA,    UC. 

US.    ITT,    UTC.    wsc);    ,T.    Howell    24689 

(CVS). 

Habitat. —  Red  sandv,  gravellv.  or  clay 
soils  at  ca.  4,500  to  5,000  feet.  ' 

Distribution. —  Known  only  from  San 
.Tuan  and  Grand  cos.,  -Utah.  It  probabh 
OTows   in    (Colorado    near   Moab   and   also 


ATWOOD:     PHACELIA   CRENUL.\TAE   GROUP 


165 


Map  13.     Idaho,     Montana,     Wyoming,    and 
Utah.    Distribution  of  P.  glandulosa  Nuttall. 

Monument  Valley  in  Arizona,  although 
no  specimens  have  been  seen  from  either 
area   (Map  14). 

This  entity  is  related  to  P.  corrugata 
A.  Nelson  which  grows  throughout  most 
of  Utah  and  extends  to  Colorado,  northern 
New  Mexico,  and  northern  Arizona.  It 
is  distinguished  from  P.  corrugata  by  its 
low,  much-branched  growth  form  and 
smaller  corolla  with  a  white  tube.  The 
leaves  are  mostly  basal,  and  the  seeds  are 
larger  and  reddish  brown. 
19.  Phacelia  integri folia  Torr.  ex  Watson 

Plants  annual  (possibly  biennial  in 
^ar.  te.vana) ,  1.2-6  dm  tall,  stems  stout, 
green  to  reddish  brown,  simple  or 
branched,  puberulent,  stipitate-glandular 
and  hirsute;  leaves  simple,  crenate  to 
somewhat  cleft,  oblong  to  ovate  or  lanceo- 
late, strigose,  finely  glandular,  setose, 
lower  leaves  long  petiolate,  the  upper  short 
pctiolate  to  sessile;  inflorescence  of  com- 
()ound  scorpioid  cymes,  the  cymes  elon- 
ij,ating  to  as  much  as  2.1  dm  in  fruit,  ped- 
i(  els  0.5-2  mm  long;  sepals  elliptical  to 
nl)]anceolate,  2.5-6.5  mm  long,  0.9-2.8 
mm  wdde,  often  reddish,  puberulent,  hir- 
sute and  stipitate-glandular;  corolla  cam- 
panulate,    purplish    to    lavender,    4.5-6.5 


Fig.  36.  Dorsal  and  ventral  view  of  the 
seeds  of  P.  howelliana  Atwood.  J.  T.  Howell 
24687   (c.^s). 


mm  long  and  broad,  the  lobes  crenulate, 
pubescent;  stamens  and  style  exserted  4- 
6.5  mm;  style  bifid  2/3-3/4  its  length, 
pubescent  below,  the  filaments  purplish, 
the  stamens  bluish  green;  capsule  ovoid 
to  globose,  2.6-5.3  mm  long,  1.1-3.5  mm 
wide,  glandular  and  puberulent;  mature 
seeds  ovate  or  elliptic  to  oblong,  reddish 
brown  or  dark  brown  to  black,  2.4-4.5 
mm  long,  1.3-2.2  nun  wide,  the  dorsal 
surface  pitted  and  transversely  ridged, 
the  margins  more  finely  pitted  than  the 


Map  14.     San     Juan     and     Grand     counties, 
Utah.    Distribution  of  P.  howelliana  Atwood. 


166 


GREAT  BASIN  NATURALIST 


Vol.  35,  No.  2 


excavated  portions  (these  tending  to  have      surface   excavated   on    both   sides   of   the 
transversely    elongate    pits),    the    ventral      ridge,  corrugated  or  corrugations  lacking. 


la. 


lb. 


Key  to  the  varieties  of  P.  integrifoUa 

Mature  seeds  3  mm  long  or  less,  1.4  mm  wide  or  less,  ventrally  cor- 
rugated; capsule  3.1  mm  long  or  less;  plants  of  southeastern  New 
Mexico  and  adjacent  Texas var.   texana 

Mature  seeds  3.1  mm  long  or  more,  1.7  mm  wide  or  more,  ventral 
corrugations  lacking;  capsule  3.2  mm  long  or  more;  plants  wide- 
spread in  rocky  to  sandy  soil var.  integrifoUa 


19a.  var.  integrifoUa 
Fig.  37 

Phacelia  integrifoUa  Torr.  in  Wats.,  Ann.  Lye. 
New  York  2:222,  t.  3.  1826.  Holotype:  on 
the  Platte,  25  June  1820.  Dr.  James  s.n. 
(ny!).  Phacelia  arenicola  Brandegee.  Univ. 
Calif.  Pub.  Bot.  4:185.  1911.  Holotype:  Mex- 
ico: Coahuila:  El  Tore  near  Movano.  July 
1910,  C.  A.  Purpus  4458  (us!). 
Phacelia  integrifoUa  Torr.  in  Wats.  var. 
arenicola  (Brandegee)  Brand.  Das  Pflanzen- 
reich  IV,  251:82.     1913. 

Plants  annual,  1.6-6  dm  tall;  stems  pu- 
berulent,  finely  to  densely  stipitate-glan- 
dular  and  hirsute;  leaves  1-13  cm  long,  0.5- 
3  cm  wide;  cymes  elongating  to  2.1  cm  in 
fruit,  pedicels  1  mm  long;  sepals  ob- 
lanceolate  to  elliptic,  3.5-4.5  mm  long 
(4.4-6.5  mm  in  fruit),  1-1.8  mm  wide 
(1.1-2.8  mm  in  fruit);  corolla  5-6.5  mm 
long  and  broad;  stamens  and  style  ex- 
serted  5-6  mm;  capsule  ovoid  to  globose, 
3.2-5.3  mm  long,  3-3.5  mm  wide;  mature 
seeds  oblong  to  elliptic,  dark  brown  to 
black,  3.1-4.5  mm  long,  1.7-2.2  mm  wide, 
transverse  ridges  on  the  dorsal  surface 
cpiitG  distinct,  the  ventral  surface  lacking 
corrugations,  the  ridge  often  curved  to  one 
side  (Fig.  38).  Collections:  113  (23);  rep- 
resentative: L.  Higgins  3138,  3129,  3131 
(bry);  D.  Atwood  2555,  2556,  2278,  2275, 
2273,  2169  (b,  bry,  cas,  ny.  wts)  ;  D.  At- 
wood 2265,  2263,  2557a,'  2274,  2171 
(bry). 

Habit.at.—  Sandy  hills  and  flats,  rocky 
hillsides  of  Larrca,  Yucca,  Quercus,  Coleo- 
gyne.  and  grass  communities.  From  3.750 
to  7,500  feet,  late  March  to  mid-Ser)tem- 
ber. 

Distribution. —  Southeastern  Utah  in 
Kane  and  San  Juan  cos.,  southward 
through  northeastern  Arizona,  eastward 
through  much  of  Now  Mexico  to  western 
Texas  and  Chihuahua,  Mexico  (Map  15) 


Fig.  37.     Phacrlia  integrifoUa  Torr.  e.\  Wats. 
var.    integrifoUa.    L,    Higgins    3131     (bry). 


June  1975 


ATWOOD:    PHACELIA   CRENLLATAE  GROUP 


167 


Fig.  38.  Dorsal  and  ventral  view  of  the 
seeds  of  P.  integrifolia  Torr.  ex  Wats.  var.  in- 
tegrifolia.  D.  Atwood  2556  (bry). 

Considerable  confusion  has  existed  as  to 
the  relationships  of  this  taxon  to  other 
species.  This  has  probably  resulted  from 
the  inadequate  type  material  and  initial 
misunderstandings  that  have  been  per- 
petuated and  even  enlarged  upon  by  sub- 
sequent authors,  Brand  fl913)  and  Voss 
(1937).  These  misunderstandings  have, 
in  part,  come  about  through  lack  of  field 
work  and  by  the  fact  that  the  corollas  in 
most  herbarium  specimens  fade  to  white 


Map  15.  Parts  of  southwestern  United  States 
and  adjacent  Mexico.  Distribution  of  P.  integrifolia 
Torr.  ex  Wats.:  #  var.  integrifolia;  var.  O.  tex- 
ana. 


and  appear  to  be  tubular.  Var.  integrifolia 
has  a  broad  distribution  and  is  easily  dis- 
tinguished b}'  its  large,  broad,  noncor- 
rugated  seeds.  Plants  from  the  higher 
elevations  in  western  New^  Mexico  and 
eastern  Arizona  are  different  morpho- 
logically. Additional  field  work  is  nec- 
essar)'  to  decide  whether  these  differ- 
ences are  sufficient  to  warrant  taxonomic 
recognition. 

19b.  var.  texana  (Vossj  Atwood, 
new-  comb. 

Phacelia  texana  Voss.  Bull.  Torr.  Bot.  Club  64:141. 
1937.  Holotvpe:  Texas:  Hudspeth  Co.:  Fin- 
lay,  5  May  1931,  M.  E.  Jones  28500  (pom!); 
Isotypes  (rm.  uc);  photo  at  (bry.  ny.  uc, 
us). 

Plants  annual  or  possibly  biennial.  1.2- 
4.3  dm  tall:  stems  puberulent,  densely 
covered  with  short  stipitate  glands  (usu- 
ally 0.2  mm  long  or  less;  and  sometimes 
^^dth  a  few  scattered  longer,  simple  hairs; 
leaves  1-10  cm  long,  0.3-2  cm  wide,  stip- 
itate-glandular  (0.2  mm  long  or  less),  and 
puberulent;  cymes  elongating  to  1.4  dm 
in  fruit,  pedicels  0.5-1.2  mm  long;  sepals 
elliptical  to  oblanceolate,  more  or  less 
heteromorphic  ^two  narrow  and  three 
broad),  2.5-3  mm  long  (3.5-5.5  in  fruit), 
0.9-1.4  mm  wide  (1.2-2  in  fruit)  puber- 
ulent, stipitate-glandular,  and  setose;  co- 
rolla 4.5-6  mm  long  and  broad;  stamens 
and  style  exserted  4-6.5  mm;  capsule 
globose,  2.6-3.2  mm  long,  1.1-2.6  mm 
wide;  mature  seeds  ovate,  reddish  bro\Mi, 
2.4-3  mm  long,  1.1-1.4  mm  wide,  trans- 
verse ridges  on  the  dorsal  surface  only 
fairly  distinct,  the  central  surface  cor- 
rugated on  the  ridge  and  part  of  the  mar- 
gin. Collections:  38  (Oj;  representative: 
C.  Cory  37574  (uc);  H.  Wilkens  2209 
(uc);  L.  Higgins  3157  (bry,  wts)  ;  T. 
Collins  1182  (uc);  U.  Waterfall  4558 
(gh,  CAS,  NY);  R.  McVaugh  8163  (uc). 

H.ABiTAT.^ —  Gypsum,  limestone,  and  cal- 
careous soils  in  mixed  shrub  commun- 
ities. Commonly  associated  with  Larrea. 
Foquieria,  and  Acacia,  2,550  to  6,850  feet 
elevation.    Late  April  to  early  October. 

Distribution. —  Southeastern  New 
Mexico  and  adjacent  Texas   (Map  15). 

Var.  texana  differs  morphologically 
from  var.  integrifolia  in  seeds  and  size  of 
capsule.  The  corollas  of  var.  texana  ap- 
pear to  be  lavender  in  contrast  to  the  pur- 


168 


GREAT  BASIN  NATURALIST 


Vol.  35,  No.  2 


plish  corollas  of  the  latter.  Jones  (1931) 
indicated  that  he  had  collected  the  type 
at  Findlay,  correctly  spelled  Finlay.  The 
correct  collection  number  is  28500  rather 
than  285ae  as  cited  by  Voss  (1937). 

20.  Phacelia  intermedia  Wooton 
Fig.    39 

Phacelia  intermedia  Wooton.  Bull.  Torr.  Bot. 
Club  25:457.  1898.  Holotype:  New  Mex- 
ico: Dona  Ana  Co.:  mesa  west  of  the  Organ 
Mountains.  10  April  1893,  E.  O.  Wooton 
(us!);  duplicates  (uc,  us). 

Plants  annual,  0.6-3.6  dm  tall,  stems 
simple  or  branched,  often  reddish,  puber- 
ulent  with  short  stipitate  glands;  leaves 
narrowly  oblong,  elliptic  to  ovate-lanceo- 
late, sinuate  to  deeply  lobed  or  pinna- 
tifid,  0.5-8  cm  long,  strigose  and  stipitate- 
glandular  or  glandular,  petiolate  to  ses- 
sile above;  inflorescence  of  compound 
scorpioid  cymes,  terminal  on  the  main 
stem  and  lateral  branches,  the  cymes  up 
to  1.5  dm  long  in  fruit,  pedicels  subses- 
sile  to  1  mm  long  in  flower,  slightly  longer 
in  fruit;  sepals  linear  to  oblanceolate,  2.5- 
3.7  mm  long,  1.3-1.5  mm  wide,  setose 
and  glandular;  corolla  bluish  violet,  6 
mm  long  or  less,  campanulate,  pubescent, 
the  lobes  crenate  to  entire;  stamens  and 
style  exserted  3  mm  or  less,  filaments 
violet,  anthers  yellow,  style  violet,  bifid 
2/3  its  length,  the  lower  1/3  glandular 
and  puberulent;  capsule  oval,  3  mm  long 
and  broad,  glandular  and  puberulent;  ma- 
ture seeds  ovate,  2.7-2.9  mm  long,  1.5- 
1.6  mm  wide,  dark  brown,  pitted,  ventral 
surface  corrugated  on  one  side  of  the 
ridge,  both  margins  and  partly  to  com- 
pletel}^  across  the  excavations  (Fig.  40) . 
Collections:  34  (7);  representative:  E. 
Wooton  s.n.  (us);  D.  Atwood  2554,  2560, 
2570,  2565,  2572.  2557.  2170,  (bry);  L. 
Higgins  3118,  3114  (bry,  wts)  ;  H.  Bob- 
isud  149   (nm). 

Habit.at. —  Sandy  to  gravelly  or  c\i\\ 
soils  of  foothills  and  higher  mesas  from 
3,750  to  5,000  feet.  Often  associated  with 
Larrea  and  Prosopis,  March  to  May. 

Distribution. —  Central  New  Mexico, 
southward  to  western  Texas  and  northern 
Mexico  (Map  16). 

This  taxon  is  related  to  P.  corru<yat(i 
but  is  distinguished  from  it  by  the  smaller, 
darker,  and  more  distinctly  corrugated 
seeds,  smaller,  lighter  corolla,  and  shortlv 


Fig.  39.  Phacelia  intermedia  Wooton.  D. 
Atwood  2560   (bry). 

exserted  stamens.  It  has  been  confused  to 
some  degree  with  P.  bombycina  W.  &  S. 
but  is  easily  separated  from  it  by  seed, 
]iubosceiice.   and    vegetative   features. 

21.  P/iacelia  uiartiniillarcnsis  Atwood 

Phytologia  26  i d  i :  .1)7.  197) 

Fig.  41 

Holotype:  Utah:  Kane  Co.:  Tropic  Shale 
formation  ca.  6  mi  east  of  Glen  Canyon  City, 
along  road  to  Warm  Creek,  S.  Welsh  and  D. 
.Xtwcod    9809,     (Bin);    IsoKpes    (aiuz.    .\su,    bry, 

oil,    RM.    uc,    UT,    UTC). 


June  1975 


ATWOOD:    PHACELIA   CRENULATAE   GROUP 


169 


Fig.  40.     Dorsal    and    ventral    view    of    the 
seeds  of  P.  intermedia  Wooton.  D.  Atwood  2557 

(bry). 

Plants  annual,  0.9-5  dm  tall;  stems 
erect  or  sometimes  branched  below,  yel- 
lowish or  green,  densely  stipitate-glan- 
dular;  leaves  simple,  oblong  to  lanceolate, 
irregularly  crenate  to  dentate,  1-7  cm  long, 
0.5-3  cm  wide,  stipitate-glandular,  setose 
to  strigulous,  with  sessile  leaves  or  nearly 
so;  inflorescence  of  terminal  or  lateral 
compound      scorpioid      cymes,      stipitate- 


1 

• 

/■ 

Fig.  41.     Phacelia 
Atwood  2632   (bry). 


mammillarensis    Atwood. 


Map  16.     New    Me.xico    and    western   Texas. 
Distribution  of  P.  intermedia  Wooton. 


with  glandular,  puberulent,  hirsute  to  se- 
tose; sepals  elliptic  to  oblanceolate,  4-6  mm 
long,  1-2  mm  wide,  stipitate-glandular, 
and  hirsute  to  setose;  corolla  tubular  to 
funnelform,  the  lobes  pale  blue  to  white, 
5-8  mm  long;  stamens  and  style  ex- 
serted  5-10  mm,  the  anthers  lavender, 
the  style  bifid  ca.  1/2  its  length,  the  lower 
1/4  pubescent;  capsule  subglobose,  4-5 
mm  long,  pubescent;  seeds  4,  3  mm  long, 
1.5  mm  wide,  brown,  pitted  dorsally, 
ventral  surface  pitted,  excavated,  and  di- 
vided by  a  prominent  ridge,  one  side  of 


170 


GREAT  BASIN  NATURALIST 


Vol.  35,  No.  2 


the  ridge  corrugated,  margins  corrugated. 
Collections:  20  (15);  representative:  D. 
Atwood  2628.  2632,  1874,  1878,  4553, 
3743,  3835  (bry);  B.  Olsen  34  (bry). 

Habitat. —  Endemic  to  the  Tropic 
Shale-Kaiparowits  formations. 

Distribution. —  Kane  and  Garfield 
COS.,  Utah,  May- June. 

Phacelia  mammillarensis  is  related  to 
P.  corrugata  Nelson  but  differs  in  its 
larger  stature,  sessile  leaves  (at  least 
above),  light  blue  to  whitish  corolla,  and 
larger,  more  densely  stipitate-glandular 
pubescence.  The  leaves  are  not  deeply 
lobed  or  pinnate  as  is  typical  in  many 
plants  of  P.  corrugata. 

22.  Phacelia  neomeiicana  Thurber 
ex  Torr. 

Phacelia  neomexicana  Thurber  ex  Torr.  Bot. 
Mex.  Bound.  Surv.  143.  1859.  Holotype: 
New  Mexico:  Grant  Co.:  pine  woods  Santa 
Rita  Copper  Mines,  August  1851.  Thurber 
1111  (ny!);  duplicate  (gh).  Paratype:  New 
Mexico:  1851,  C.  Wright  1577  (ny.  us). 
Phacelia  glandulosa  Nutt.  var.  neomexicana 
(Thurber  ex  Torr.)  Gray.  Proc.  Anier. 
Acad.  10:319.     1875. 

Phacelia  neomexicana  Thurber  ex  Torr.  var. 
eu-neomexicana  Brand.  Das  Pflenzenreich  IV. 
251:83.     1913. 

Plants  annual,  0.8-6.8  dm  tall;  stems 
erect  to  sparsely  branched,  setose  and  with 
small  stipitate-glandular  hairs,  often  red- 
dish, leafy;  leaves  pinnate,  the  secondary 
pinnae  irregularly  incised,  3-8.5  cm  long, 
1-4.5  cm  wide,  strigose  and  stipitate- 
glandular,  petiolate,  the  petiole  1.5  cm 
long  or  less;  inflorescence  terminal  on  the 
main  stem  and  lateral  branches  (some- 
times arising  from  the  axils  of  the  upper- 
most leaves),  more  glandular  than  the 
stem,  the  individual  inflorescence  branches 
with  1-3  cymes,  the  cymes  up  to  1  dm 
long  in  fruit,  flowers  congestecl  and  short 
pedicellate  (0.5  mm  long);  sepals  linear 
to  narrowly  oblanceolate,  2.7  mm  long  in 
flower  to  4.5  mm  long  in  fruit,  setose  and 
heavily  glandular;  corolla  campanulate, 
blue,  ca.  4  mm  long,  ca.  3-3.5  mm  wide. 
the  lobes  pubescent  and  orose;  stamens  ex- 
serted  1  mm,  4.5  mm  long  bifid  3/4  its 
length,  glandular  on  the  lower  1/4;  cap- 
sule oval  to  elli[)tic,  4.5-4.7  mm  long,  3 
mm  wide,  setose  and  heavily  glandular, 
the  raphe  oblanceolate;  mature  seeds  4, 
oblong,   brown    (immature   .seeds   mottled 


with  dark  areas),  3.2-3.3  mm  long,  1.1-1.5 
mm  wide,  alveolate,  ventral  surface  ex- 
cavated on  both  sides  of  the  ridge,  alveo- 
late. Collections:  20  (0);  representative; 
II.  Ripley  and  R.  Barneby  5096  (rsa)  ; 
W.  Chapman  s.n.  (us);  E.  Greene  s.n. 
(ny);  O.  Metcalfe  1506  (cas,  gh,  ny); 
E.  Wooton  s.n.  (us):  E.  Castettes  4852 
(us);  B.  Dunn  6208  (ny). 

Habitat. —  Pine  and  oak  woods  in  can- 
yons and  on  mountain  slopes,  in  rocky  to 
sandy  soils,  from  6,800  to  9,000  feet,  late 
.luly  to  mid-October. 

Distribution. —  Apache  Co.,  Arizona, 
eastward  to  New  Mexico  in  Grand,  Otero. 
Socorro,  Lincoln,  Torrance,  Bernarillo,  and 
Taos  COS.    (Map   17). 

P.  neomexicana  is  apparently  most 
closely  related  to  P.  denticulata  but  dif- 
fers from  that  species  in  its  exserted  sta- 
mens and  style,  smaller  and  more  deeply 
excavated  seeds.  The  style  is  also  longer 
(4.5  mm)  and  the  corolla  is  campanulate. 

23.  Phacelia  pallida  Johnston 

Phacelia  pallida  Johnston.  Journ.  Arnold  Arb. 
24:98.  1943.  Holotype:  Me.xico:  Coahuila: 
gypsum    beds   on    the   escarpment   of   Canada 


Map    17.     Colorado  and  eastern  Arizona.    Dis- 
tribution  of  P.   neomexicana  Thurber  ex  Torr. 


June  1975 


ATWOOD:    PHACELIA   CRENIJLATAE   C.ROUP 


171 


Oscuro  near  Tanque  La  Luz,  26  August 
1941,  I.  Johnston  8486  (gh!);  Isotypes  (gh). 
Phacelia  petiolata  Johnston.  Journ.  Arnold 
Arb.  24:98.  1943.  Holotype:  Mexico:  Chi- 
huahua: 12  miles  south  of  Ojinaga,  10-12 
August  1941.  I.  Johnston  8040  (gh).  Para- 
types:  Mexico:  Chihuahua:  11.5  mi  south  of 
Ojinaga.  10-12  August  1941,  I.  Johnston  8036 
(gh);  Coahuila:  San  Lorenzo  de  la  Laguna, 
75  mi  northwest  of  Parras,  May  1880,  E. 
Palmer  851    (gh). 

Perennial  plants,  1.3-3.5  dm  tall;  stems 
branched,  especiall}'  at  the  base,  erect  to 
decumbent,  puberulent,  hirsute  to  setose 
with  flattened  multicellular  stipitate 
glands;  leaves  siinple,  some  of  the  lower 
with  several  small  lobes  borne  on  the 
petiole  below  the  oblong,  lanceolate  to 
broadly  elliptic  blade,  1-7  cm  long,  1-3 
mm  wide,  long  petiolate  below  to  short 
petiolate  above,  strigose,  glandular  and 
setose,  the  margins  irregularly  crenate  to 
sinuate;  inflorescence  terminal,  racemose, 
puberulent  to  hispid  (when  old)  and  stip- 
itate-glandular,  cymes  densely  flowered, 
up  to  15  cm  long  in  fruit,  pedicels  0.5-1 
mm  long;  sepals  oblanceolate  to  spatulate, 
4-5  mm  long,  1.4-1.7  mm  wide,  glandular, 
hirsute;  corolla  tubular  to  salverform, 
pale  lilac,  lavender,  or  white,  4.6  mm  long. 


Map.  18.  Brewster  County,  Texas,  Coahuila 
and  Chihuahua,  Mexico.  Distribution  of  P.  pallida 
Johnston. 


ca  4  mm  wide;  stamens  and  style  ex- 
serted  5-7  nnu,  style  bifid  3/4  its  length, 
the  lower  1/2  pubescent;  capsule  sub- 
globose,  3-3.5  mm  long,  2.4-2.6  mm  wide, 
puberulent;  mature  seeds  oblong  to  ellip- 
tic, brown  to  blackish,  pitted,  2.6-3  mm 
long,  1-1.5  mm  wide,  the  ventral  surface 
excavated  on  both  sides  of  the  corrugated 
ridge,  margins  corrugated,  dorsal  surface 
transversely  ridged.  Collections:  6  (0); 
representative:  C.  Purpus  5084  (uc);  O. 
Sperry  1694  (us);  I.  Johnston  80-10,  8036 
(gh). 

Habitat. —  Apparently  P.  pallida  is 
confined  to  gypsum  and  limestone  soil. 

Distribution. —  Brewster  Co.,  Texas, 
southward  into  Coahuila  and  Chihuahua, 
Mexico  (Map  18). 

P.  pallida  and  P.  petiolata  are  known 
only  from  the  type  collections  and  ap- 
pear to  be  the  same  entity.  Therefore  the 
author  has  placed  P.  petiolata  as  a  syno- 
nym of  the  former.  Additional  collections 
are  needed. 


24.   Phacelia  palmeri  Torr.  ex  Wats. 
Fig.  42 

Phacelia  palmeri  Torr.  ex  Wats.  Bot.  King  Ex- 
ped.  251.  1871.  Holotype:  Utah:  Washing- 
ton Co.:  southern  Utah  near  St.  George  on 
the  Rio  Virgin,  1870.  Palmer  4  (ny!);  Iso- 
types (gh,  us). 

Phacelia  joetida  Goodding.  Bot.  Gaz.  37:58. 
1904.  Holotype:  Utah:  Washington  Co.:  vol- 
canic slopes.  Diamond  Valley,  16  May 
1902,  L.  N.  Goodding  833  (rm!). 
Phacelia  palmeri  Torr.  ex  Wats.  var.  foctida 
(Goodding)  Brand.  Das  Pflanzenreich  IV, 
251:79.     1913. 

Phacelia  palmeri  Torr.   ex  Wats.  var.   typica 
Voss.  Bull.  Torr.   Bot.  Club  64:90.     1937. 
Phacelia  integrifolia  Torr.  var.  palmeri  (Torr. 
ex   Wats.)    Gray.   Proc.   Amer.  Acad.    10:318. 
1875. 

Robust  biennial,  2.9  dm  tall;  stems  stout, 
usually  solitary  (sometimes  with  few  to 
several  branches  at  the  base),  densely 
glandular,  hirsute  and  pilose,  becoming 
hispid  with  age;  leaves  oblong  to  lance- 
olate, irregularly  sinuate,  crenate,  den- 
tate, or  serrate,  2-13  cm  long,  0.5-3  cm 
wide,  lower  densely  tufted,  petiolate  and 
larger  than  the  sessile,  gradually  reduced 
cauline  leaves,  stipitate-glandular  and 
strigose;  inflorescence  a  dense  spicate  th}T- 
sus,  0.4-4.2  dm  long,  individual  scorpioid 
cymes  uji  to  14  cm  long  in  fruit,  pedicels 


172 


GREAT  BASIN  NATURALIST 


Vol.  35,  No.  2 


Fig.  42.     Phacelia  palmed  Torr.  ex  Wats.  D.  Alwood   1690    (bry). 

about  1  mm  long  in  fruit;  sepals  oblong  to  long,    pubescent:    stamens    and    style    ex- 

spatulate,  4-5  mm  long,  1-1.8  mm  wide,  serted     5-7     nnn,     style     bilid,     the     un- 

glandular  to  hirsute;   corolla  pale    (whit-  branched   portion   pubescent;   capsule  gio- 

ish,  lavender,  or  violet),  tubular,  5-7  mm  bose,    ).2  nnn  long,  glandular  to  hirsute; 


197c 


ATWOOD:    PHACELIA   CRENULATAE   GROUP 


173 


mature  seeds  4,  elliptic,  black,  2.5  mm 
long,  1.5  mm  wide,  excavated  on  both 
sides  of  the  ridge,  the  ridge  corrugated 
on  one  side,  pitted,  margins  corrugated, 
fvirrows  or  grooves  partly  corrugated,  dor- 
sal surface  longitudinally  pitted  and  trans- 
versely ridged  (Fig.  43).  Collections:  54 
(16);  representative:  D.  Atvvood  1530, 
1390,  1720,  1409  (bry);  L.  Higgins  817, 
1244  (bry);  D.  Atwood  and  L.  Higgins 
1682    (bry,  us);  D.  Atwood   1723,   1712 

(bry,  CAS,  GH.  NY,  RM,  Us). 

Habitat. —  Mostly  on  barren  to  sparse- 
ly vegetated  gypsum  flats,  washes,  and 
hillsides  but  not  uncommon  on  rocky  to 
sandy  soil.  In  Diamond  Valley  (north 
of  St.  George)  it  grows  on  volcanic  cinder 
cones,  at  2,700  to  5,000  feet  elevation,  late 
March  to  August.  Commonly  associated 
with  Larrca,  Juniperus.  Cou>ania,  Follugia. 
and  Atriplex. 

Distribution. —  Clark  Co.,  Nevada, 
eastward  to  Washington  and  Iron  cos., 
Utah,  and  Mohave  Co.,  Arizona  (Map  19). 

P.  palmeri  is  related  to  P.  constancei 
Atwood  but  differs  in  having  taller  stems, 
larger  and  less  revolute  leaves,  and 
coarser  and  longer  hair.  The  lateral 
branches  are  fewer  and  less  leafy,  and  the 
stems  lack  the  reddish  color  characteristic 
of  the  latter.  Both  species  occur  on  the 
Moenkopi  formation  but  have  different 
ranges.  A  form  from  Lake  Mead,  Arizona 
(E.  U.  Clover  6470  &  6230),  has  seeds 
that  are  not  corrugated  and  are  smaller 
and  narrower  (2.0  mm  long  and  1.2  mm 
wide),  dark  dorsally  but  brown  ventrally 
with  darker  glands.    However,  in  the  ma- 


r 


Fig.  43.  Dorsal  and  ventral  view  of  the 
seeds  of  P.  palmeri  Torr.  ex  Wats.  D.  Atwood 
1723  (bry). 


Map  19.  Southwestern  Utali,  northwestern 
Arizona,  and  southeastern  Nevada.  Distribution  of 
P.  palmeri  Torr.  ex  Wats. 

terial  from  Glendale,  Nevada,  the  seeds 
are  dark  dorsally  and  lighter  ventrally 
and  are  less  corrugated  than  in  typical 
material. 


25.  Phacelia  pcdiceUata  Gray 
Fig.  44 

Phacelia  pedicellata  Gray.  Syn.  Fl  II.  1:160. 
1878.  Holotype:  Mexicor  Baja  California: 
Lower  California,  1875,  Dr.  Streets  s.n. 
(gh!);  Isotype   (us). 

Plants  annual,  1-6  dm  tall;  stems 
branching  or  sometimes  simple,  brittle, 
villous  to  setose  with  multicellular  stalked 
glands;  leaves  suborbicular  to  oblong,  pin- 
nately  compound  with  3-9  pinnae,  0.3-1.3 
cm  long,  1.5-11.5  cm  wide,  villous  to  setose 
and  glandular;  inflorescence  of  compound 
cymes,  somewhat  dichotomously  branched; 
pedicels  filiform,  2.6  mm  long,  densely 
setose  to  hirsute;  sepals  linear  to  oblance- 
olate,  2.8-7.9  mm  long,  0.6-2  mm  wide, 
setose  to  hirsute  and  glandular;  corolla 
lobes  lavender,  violet,  or  white,  the  tube 
white,  ca.  5  mm  long  and  broad;  stamens 
and  style  exserted,  style  pubescent,  upper 


174 


GREAT  BASIN  NATURALIST 


Vol.  35,  No.  2 


Fig.  44.     Phacelia  pcdicellata  Gray.  W.  Jcp- 
son  12482  (c.\s). 


Habitat. —  Dry  gravel  and  sandy 
washes,  often  in  the  shade  of  large  boul- 
ders, limestone  cliffs,  and  as  understory  of 
larger  plants,  below  sea  level  in  Death 
Valley  to  5,000  feet  elevation.  It  has  been 
collected  in  the  middle  of  February  in 
Mexico  but  usually  flowers  from  March  to 
the  middle  of  June. 

Distribution. —  Southern  Nevada  in 
Nye  and  Clark  cos.,  southward  through 
San  Bernardino  to  central  Baja  California, 
eastward  to  Coconino^  Graham,  Gila,  Pi- 
nal, and  Pima  cos.,  Arizona  (Map  20). 

Phacelia  pedicellata  is  most  closely  re- 
lated to  P.  scariosa  h\xt  differs  in  having 
narrower,  longer,  and  more  pubescent 
calyx  lobes,  a  more  compound  and  con- 
gested inflorescence,  and  a  heavier,  glan- 
dular, villous,  and  setose  pubescence.  The 
leaves  have  3-9  pinnae,  whereas  those  of 
P.  scariosa  have  only  3-5  lobes,  and  the 
style  is  more  pubescent.  That  they  are 
related  is  indicated  by  the  similar  seeds, 
corolla,  pedicels,  and  brittle  stems. 


26.  Phacelia  popei  Torr.  &  Gray 
Fig.  46 

Phacelia  popei  Torr.   &   Gray.   Pacific   Rail.   Rep. 
Explor.    Mississippi   2:172.     1885.     Holotype: 
Te.xas:     Llano     Estacado,     no    date,     Captain 
Pope  s.n.   (gh!).  Paratype:  Te.xas:   Pecos  Co.: 
C.  Wright  1578  (gh,  ny,  us). 
Phacelia    popei    Torr.    &    Gray    var.    typica 
Voss.    Bull.    ToiT.    Hot.    Club   64:94.     1937. 
Phacelia    similis    Wooton    &    Standley.    Bull. 
Torr.    Bot.    Club    36:111.      1909.     Holotype: 
New  Me.xico:    Sierra  Co.:   on  the  plains  near 
Nutt  Station,  12  May  1905,  O.  Metcalfe  1665 
(ny!). 
Phacelia    popei    Torr.    &    Gray    var.    similis 


1/3  bifid;  capsule  globular,  3-3.4  mm 
long,  2.3-2.5  mm  wide,  pilose  to  glan- 
dular; mature  seeds  4,  elliptical,  3  mm 
long,  1.1-1.8  mm  wide,  ventral  surface  ex- 
cavated, pitted  to  tuberculate,  the  ridge 
corrugated  on  one  side,  dorsal  surface  tu- 
berculate and  pitted,  margins  corrugated 
(Fig.  45).  Collections:  155  (3);  represen- 
tative: A.  Eastwood  17400  (cas,  ny,  us); 
M.  .Tones  s.n.  (pom,  utc);  W.  Cottam 
13125  (ut);  T.  Brandegee  s.n.  (ny);  H. 
Ripley  and  R.  Barneby  2952  (rsa);  .1. 
Howell  3952  (utc);  D.  Atwood  2339 
(bry)  ;  S.  Welsh,  D.  Atwood  and  E.  Matt- 
hews 9633  (bry). 


Fi^'.  45.     Dorsal     and     \entral     view    of    the 
seeds  of  P.  pedicellata  Gray.  M.  Beal   (jeps). 


Tune  1975 

AT  WOOD:    PHACELI 

V      \ 

N 

•• 

•  •\4 

VN 

^ 

\^_t  -   ••  •  •  1 

S         c»^ 

■^"^^^ 

^ 

^ 

Map  20.  Part  of  southwestern  United  States 
and  adjacent  Baja  California,  Mexico.  Distribution 
of  P.  pedicellata  Gray. 

(Wooton  &  Standley)    Voss.   Bull.   Torr.   Hot. 

Club  64:94.     1913. 

Phacelia  glandulosa  A.   Gray   in   Brand,   Das 

Pflanzenreich     IV,     251:84.      1913,     in     syn- 

onomy. 

Phacelia    depauperata    Wooton    &    Standley. 

Contr.     U.     S.     Natl.     Herb.     16:163.     1913. 

Holotype:    New   Mexico:    Caves   Co.:    Arroyo 

Ranch  near  Roswell,   1903,  D,   Griffiths  4249 

(us!). 

branched  from  the  base,  0.5-3.6  dm  tall, 
with  simple  spreading  or  bent  hirsute  and 
somewhat  glandular  hairs  intermixed  with 
a  finer  pubescence;  leaves  narrowly  ob- 
long, pinnate  to  bipinnate,  with  linear  or 
lanceolate  divisions,  2-15  cm  long,  1-3  cm 
wide,  petiolate,  strigose  to  glandular,  ex- 
cept on  the  petioles  and  then  like  that  on 
the  stems;  inflorescence  of  compound 
scorpioid  cymes,  the  cymes  crowded,  ter- 
minal, up  to  10  cm  long  in  fruit,  glan- 
dular and  hirsute,  flowers  nearly  sessile 
(pedicels  ca.  0.5  mm  long) ;  corolla  cam- 
panulate,  blue  to  purplish,  3.5-7  mm  long 
and  broad,  pubescent;  sepals  oblanceolate 
to  spatulate,  somewhat  keeled  (at  least  at. 
the  base),  2.3-3.9  mm  long,  1-1.8  mm 
wide,    glandular,    hirsute;     stamens    and 


175 


style  exserted,  style  exserted  ca  2  mm 
longer  than  the  stamens,  bifid  2/3  its 
length,  lower  1/3  pubescent;  capsule  glo- 
bose, 2.3-2.5  mm  long,  2.4-3.1  mm  wide, 
glandular  and  pilose;  mature  seeds  4, 
cymbiform,  ovate,  brown,  1.8  mm  long, 
1.4  mm  wide,  favose  to  reticulate,  ventral 
surface  deeply  excavated  on  both  sides  of 
the  ridge,  dorsal  surface  reticulate  and 
transversely  ridged  (Fig.  47).  Collections: 
150  (11);  representative:  D.  Atwood 
2096a,  2095,  2268,  2266,  2153,  2159,  2131, 
2133,  2134  (bry);  L.  Higgins  3083,  3025 
(bry,  wts);  R.  Barneby  12593  (cas,  ny)  ; 
M.  Jones  25750  (pom);  A.  Hershey  s.n. 
(cas). 

Habitat. —  Commonly  in  sandy  or 
sandy  clay  soil  of  roadsides.  Less  com- 
monly in  limestone  or  rocky  soil  and  as- 
sociated with  Larrea,  Prosopis^  Yucca,  or 
short  grass  prairie.  Mid-February  to  late 
May. 


Fig.  46. 
Higgins  2931 


Phacelia  popei  Torrey  &  Gray.   L. 

(bry). 


176 


GREAT  BASIN  NATURALIST 


Vol.  35,  No.  2 


Fig.  47.  Dorsal  and  ventral  view  of  the 
seeds  of  P.popei  Torrey  &  Gray.  V.  Cory  13616 
(gh). 

Distribution. —  West  central  Texas 
from  Dickens  and  Lubbock  cos.  soutli- 
ward  to  southwestern  Texas,  westward  to 
New  Mexico  from  Roosevelt  Co.,  west- 
W'ard  to  Sierra  and  Luna  cos.,  Nuevo  Leon, 
Mexico  (Map  21). 

Wooton  and  Standley  (1909),  in  their 
original  description,  indicated  that  P.  si- 


milis  was  most  closely  related  to  P.  arizo- 
nica  but  failed  to  mention  P.  popei.  Com- 
parison of  the  type  material  of  the  two 
entities  leaves  little  doubt  that  they  are 
identical.  Also,  P.  arizonica  is  very  dis- 
tinct from  P.  popei.  Torrey  and  Gray 
(1885),  in  their  original  description  of 
P.  popei.,  indicated  that  the  corolla  was 
"apparently  white."  I  have  seen  this  tax- 
on  in  the  field  in  Texas  and  New  Mexico 
and  find  that  the  corolla  is  not  white  or 
only  seldom  so.  The  flowers  of  some 
specimens  turn  white  upon  drying,  and 
this  may  have  been  the  case  with  the  type 
material. 

27.  Pliacclia  rafachmsis  Atwood 
Fig.  48 

Phacelia  rafaelcnsis  Atwood.  Rhodora  74(800): 
454-457.  1972.  Holotype:  Utah:  Wayne 
Co.:  Capitol  Reef  National  Monument,  12 
June    1969,  D.  Atwood  and  L.  Higgins  1834 

(bry!). 


Map  21.  New  Mexico,  Texas,  and  Nuevo 
Leon,  Mexico.  Distribution  of  P.  popei  Torrey  & 
Gray. 


Fig.   4b.     Phacelia  rafaelcnsis  Atwood.  D.  At- 
wood  and   L.   Higgins   1834   (liin'). 


June  1975 


ATWOOD:    PHACELIA   CRENULATAE   GROUP 


177 


Erect  biennial  herb,  0.8-5.4  dm  tall; 
stems  stout,  simple  or  sometimes  branched 
at  the  base,  olive  green  to  brownish  glan- 
dular, and  hirsute;  basal  leaves  petiolate, 
dentate,  crenate  to  pinnatifid,  2-7  cm  long; 
0.5-1.5  cm  wide,  strigose  to  hirsute,  cau- 
line  leaves  sessile,  undulate  to  crenate  or 
dentate,  oblong-lanceolate,  1-10  cm  long, 
0.5-3.5  cm  wide,  strigose  to  hirsute  and 
sparsely  stipitate-glandular;  inflores- 
cence mainly  terminal,  paniculate,  some 
axillary,  flowers  nearly  sessile;  sepals 
oblanceolate  to  spatulate,  5-4  mm  long  in 
flower,  5-6  mm  long  in  fruit,  1-1.7  mm 
wide,  glandular  and  hirsute;  corolla  tub- 
ular, pale  and  grooved  with  the  lobes 
somewhat  spreading,  5-6  mm  long;  sta- 
mens and  style  exserted  only  3-5  mm, 
anthers  dull  in  color,  style  bifid  3/4  its 
length,  the  lower  half  pubescent;  capsule 
globose,  4-5  mm  long,  stipitate-glandular 
and  hirsute;  mature  seeds  4,  elliptic  to 
oblong,  3.5-4  mm  long,  1.5-2  mm  wide, 
ventral  surface  alveolate,  lighter  than  the 
dorsal  surface,  excavated  and  divided  by  a 
prominent  ridge,  the  ridge  sometimes  cor- 
rugated along  one  side,  the  margins  usu- 
ally entire,  dorsal  surface  brown  and  less 
deeply  pitted,  the  surface  often  smoothish 
(Fig.  49).  Collections:  33  (19);  repre- 
sentative: M.  Jones  s.n.  (pom);  H.  Ripley 
and  R.  Rarneby  4362  (rsa);  W.  Cottam 
13313  (ut);  D.  Atw^ood  1530,  1853,  1843, 
1855,  1847,  1703,  1417,  1698,  1860  (bry); 
S.  Welsh,  D.  Atwood,  and  G.  Moore  9846, 
9844,  9903  (bry);  D.  Atwood  1390  (bry). 

Habit.at. —  Clay  hills  of  the  Moenkopi 
formation.  May  to  June. 

Distribution. —  Emery  Co.,  Utah, 
southward  to  Kane  Co.,  Utah,  and  Mo- 
have Co.,  Arizona,  eastward  to  Washing- 
ton Co.,  Utah  (Map  22). 

Phacelia  rafaelensis  is  related  to  P.  uta- 
hensis  but  differs  in  having  a  slightly  tub- 
ular and  grooved  corolla,  with  the  lobes 
somewhat  spreading  and  the  stamens  and 
style  exserted  only  3-5  mm.  The  filaments 
and  stamens  are  dull  in  color,  and  the 
ridge  is  sometimes  corrugated. 

28.Phocelia  robusta  (Macbr.)   Johnst. 
Fig.  50 

Phacelia  robusta  (Macbr.)   Johnst.  Journ.  Arnold. 
Arb.  24:97.     1943. 

Phacelia    integrifolia    Torr.     e.x    Wats.    var. 
robusta  Macbride.   Contr.   Gray   Herb.   49:25. 


Fig.  49.  Dorsal  and  ventral  view  of  the 
seeds  of  P.  rafaelensis  Atwood.  S.  Welsh  et  al, 
9903    (bry). 


1917.     Holotype:  Texas:  Presidio  Co.:  Chinati 
Mountains,  no  date,  Harvard  250   (gh!). 

Robust,  viscid  annual  or  biennial,  4.5-12 
dm  tall;  stems  branched  at  the  base  or 
simple,  brownish,  puberulent,  pilose  and 
densely  glandular,  the  glandular  hairs 
flattened,  stipitate,  unicellular  to  multi- 
cellular; leaves  broadly  ovate  to  orbicular, 
irregularly  crenate  to  sinuate,  sometimes 
with  a  single  lobe  below  the  leaf  blade, 
2-11.5  cm  long,  1.5-9  cm  wide,  gradually 


Map  22.     Southern    Utah    and    northwestern 
Arizona.    Distribution  of  P.  rafaelensis  Atwood. 


178 


GREAT  BASIN  NATURALIST 


Vol.  35,  No.  2 


i    \f 


L^' ' 


d 


r 


Fig.  50.     Phacelia  robusta   (Macbride) 
ston.  I.  Higgins  3268  (bry). 


John- 


reduced  upwards,  puberulent,  stipitate- 
glandular,  setose  to  hirsute,  the  lower  long- 
petiolate  to  subsessile  above;  inflorescence 
of  compound  scorjnoid  cymes,  the  chines 
up  to  1.3  dm  long  in  fruit,  pubescence  the 
same  as  that  of  the  stem,  pedicels  1-1.7 
mm  long;  sepals  spatulate,  4.5-5.8  mm 
long,  1.5-2.5  mm  wide;  corolla  salver- 
form,  pale  lavender,  5-6  mm  long,  ca.  4 
mm  wide,  pubescent;  stamens  and  style 
exserted  4-6  mm.  style  bifid  3/4  its  length, 
lower  1/2  pubescent;  capsule  subglobose, 
3.9-4.1  mm  long,  2.8-3mm  wide,  puber- 
ulent to  strigose;  mature  seeds  oblong  to 
ovate,  reddish  brown,  2.9-3.7  mm  long, 
1.1-1.7  mm  wide,  ventral  surface  exca- 
vated on  both  sides  of  the  corrugated 
ridge,  pitted,  the  margins  at  least  partly 
corrugated  (Fig.  51).  Collections:  36  (0); 
representative:  C.  Pringle  255  (iic.  rsa)  : 
E.  Palmer  34077   (ny):  L.  Hincklev  829 


(gh,  ny);  U.  Waterfall  7316,  8255,  8283 
(uc);  C.  Smith  289  (uc);  G.  Stevens  1636 
(gh,  ny);  L.  Higgins  3256,  3182  (bry, 
wts). 

Habitat. —  Gravelly  sand  bars,  clay 
slopes,  and  rocky  hills  from  ca.  3,500  to 
6,000  feet  elevation,  March  to  August. 

Distribution. —  Barber  Co.,  Kansas, 
southward  through  Woods,  Blaine,  Custer, 
Washita,  and  Horman  cos.,  Oklahoma,  to 
north  central  and  southwestern  Texas  and 
Chihuahua,  Mexico  (Map  23). 

This  taxon  has  been  confused  with 
P.  inter grifolia,  but  it  is  easily  recognized 
by  the  white,  tubular  corollas,  robust 
habit,  and  larger,  less  glandular  leaves. 
In  seed  characters,  P.  robusta  is  similar  to 
P.  integrifolia  var.  texana.  There  appear 
to  be  several  distinct  entities  included 
within  this  taxon.  The  material  in  north 
central  Texas  is  disjunct  in  distribution 
from  that  in  Presidio  and  Brewster  cos., 
but  additional  material  is  needed  to  deter- 
mine if  there  are  sufficient  morphological 
characters  to  delineate  the  populations. 
Also,  specimens  from  southern  Colorado 
and  adjacent  New  Mexico  appear  to  be 
different  and  need  to  be  investigated 
further. 

29.  Phacelia  riipestris  Greene 

Phacelia  rupestris  Greene.  Leaflets  1:152.  1905. 
Holotype:  New  Me.xico:  Sierra  Co.:  south 
end  of  the  Black  Range,  1  mi  west  of  Hills- 
boro.  25  June  1904.  O.  Metcalfe  1012  (gh!); 
Isotypes  (cvs.  ny,  pom.  uc.  uc,  us,  us,  us). 
Phacelia  congesta  Hook.  var.  rupestris 
(Greene)  Macbride.  Contr.  Gray  Herb. 
49:25.     1917. 


.  '-H»^*V 


Fig.  51.  Dorsal  and  ventral  view  of  the 
seeds  of  P.  robusta  (Macbride)  Johnston.  L. 
Hinckley  829  (ny). 


ATWOOD:    PHACELIA   CRErsULATAE   C.ROUP 


179 


Map  23.  Southern  Kansas.  Texas,  and  ad- 
joining states.  Distribution  of  P.  robusta  (Mac- 
bride)   Johnston. 


Perennial  plants  (sometimes  flowering 
the  first  year),  1-6  dm  tall;  stems  1-many, 
usually  from  a  woody  caudex.  pubescence 
of  whitish  hairs,  these  hispid  to  setose  and 
finer,  sometimes  glandular  (these  not 
multicellular  glands);  leaves  pinnately 
compound,  the  terminal  part  incompletely 
3-5  lobed  and  larger  than  the  lower  pin- 
nae, 1.5-10  cm  long,  1-5  cm  wide,  setose 
to  densely  pilose;  inflorescence  of  ter- 
minal compound  scorpioid  c}Tnes,  individ- 
ual oTtnes  1.5-4.5  cm  long,  pedicels  1.5-2.5 
mm  long;  sepals  linear  to  oblanceolate, 
3-4.8  mm  long,  0.5-0.8  mm  w^ide,  setose 
to  hirsute;  corolla  campanulate,  white,  2-4 
mm  long  and  broad,  lobes  pubescent;  sta- 
mens and  style  exserted  up  to  ca.  2  mm, 
anthers  dull  blue;  style  5-6  mm  long, 
bifid  over  1/2  its  length,  the  lower  un- 
branched  portion  pubescent;  capsule  oval 
to  ovate,  2.6-3  mm  long,  2.4-2.5  mm  wide, 
finely  pubescent  with  a  few  long  hairs 
near  the  apex;  mature  seeds  4,  brovsn,  2.1- 
2.7  mm  long,  1-1.1  mm  wide,  elliptical 
to  oblong,  reticulate  scabrous,  ventral  sur- 
face excavated  on  both  sides  of  the  ridge 
(Fig.    52).     Collections:     78    (1);    repre- 


sentative: L.  Goodding  2330  (gh,  rm, 
uc):  H.  Rusby  s.n.  (cas.  us);  B.  Maguire 
11204  (bry);'  a.  Nelson  1248  (ny);  C. 
Pringle  162  (mexu,  ny,  us);  E.  Wooton 
s.n.  (ny);  D.  Dunn  8525  (uc). 

Habit.\t. —  Growing  on  coarse,  sandy 
soil  and  gravel  bars,  moist  shady  crevices 
of  limestone  cliffs,  and  ledges  in  canyons 
and  arroyos,  at  2,100  to  6,500  feet  ele- 
vation. Flowering  commonly  occurs 
from  late  June  to  late  August  but  some- 
times as  early  as  mid-March  and  as  late 
as  the  last  of  October.  Usually  associated 
with  Larrea,  Acacia.  Lippia.  Fallugia. 
Chilopsis,  and  Opuntia  at  lower  elevations 
and  with  Qucrcus.  Cercocarpus.  Juniperus, 
Pinus,  and  Fra.rinus  at  higher  elevations. 

Distribution. —  Southern  New  Mex- 
ico from  Socorro  Co.  westward  to  Pinal, 
Pima,  and  Cochise  cos..  Arizona,  and 
southward  to  southwestern  Texas  and  ad- 
jacent Mexico  (Map  24). 

This  entity  is  closely  related  to  P.  con- 
gesta  but  differs  in  having  smaller,  white 
corollas,  shorter  and  fewer  flowered 
cymes,  a  perennial  habit,  and  the  ab- 
scence  of  multicellular  glands.  At  times 
it  apparently  flowers  in  the  first  year,  at 
least  in  the  more  southern  parts  of  its 
range. 


30.  Pliacelia  scariosa  T.  S.  Brandegee 
Fig.  53 

Phacelia  scariosa  T.  S.  Brandegee.  Proc.  Calif. 
Acad.  Sci.  2:185.  1889.  Holotype:  Mexico: 
Baja  California:  Lower  California,  12  Janu- 
ary 1889,  Brandegee  s.n.  (uc!);  duplicates 
(ny,  us). 


► 


Fig.  52.     Dorsal  and  ventral  view  of  the  seeds 
of  P.  rupestris  Greene.  W.  Eggleston  16341   (us). 


180 


GREAT  BASIN  NATURALIST 


Vol.  35,  No.  2 


• 

I 

^ 

Map  24.  Southeastern  Arizona,  New  Mexi- 
co, southwestern  Texas,  and  adjoining  Mexico. 
Distribution  of  P.  rupestris  Greene. 

Plants  annual,  0.5-4  dm  tall;  steins 
branching  from  the  base  (sometimes 
dichotomously),  finely  glandular  and  pi- 
lose; leaves  ovate  to  oblong,  deeply  cleft 
to  more  often  pinnately  divided  with  3-5 
pinnae,  terminal  leaflet  usually  trilobed 
and  larger  than  the  other  leaflets,  1.5-8 
cm  long,  1-5.5  cm  wide,  strigose  and  some- 
what glandular;  inflorescence  paniculate 
to  racemose,  glandular  to  villous,  cymes 
elongate,  open,  up  to  11  cm  long,  pedicels 
filiform,  3  mm  long  in  flower,  up  to  8 
mm  long  in  fruit;  corolla  broadly  cam- 
panulate,  bluish  to  lavender,  with  white 
throat  and  tube,  4  mm  long,  4-5  mm  wide, 
pubescent;  sepals  obovate,  2-3  mm  long 
in  flower,  becoming  broadly  obovate  and 
conspicuously  enlarged  and  scarious  in 
fruit,  4.5-7.6  mm  long,  2.3-3.7  mm  wide, 
glandular  to  villous;  stamens  and  style 
exserted  1-2  mm,  style  cleft  1/3  its  length, 
lower  1/3  pubescent;  ca])sule  globular, 
3.3-3.7  mm  long,  2.6-3.5  mm  wide,  glan- 
dular to  pilose;  mature  seeds  4,  brown,  el- 
liptical, 2.5-3  mm  long,  1.2-1.3  mm  wide, 
ventral  surface  di\ided  by  a  prominent 
ridge,  pitted,  margins  corrugated,  the  ridge 


corrugated  on  one  side,  dorsal  surface 
curved,  pitted  (Fig.  54).  Collections:  42 
(0);  representative:  C.  Orcutt  13  (cas, 
NY,  us);  M.  Jones  24069  (pom,  rm);  T. 
Brandegee  s.n.  (uc);  R.  Moran  3890 
(uc);  D.  Porter  236  (cas,  mexu);  F. 
Shreve7023  (us). 

Habitat. —  Sandy,  gravelly  washes, 
rocky  hillsides,  and  lava  flows  from  200  to 
5,000  feet  elevation.  Apparently  a  winter 
annual,  flowering  from  late  October  to 
mid-June. 

Distribution. —  Lower  California  and 
adjacent  Sonora,  Mexico  (Map  25). 

Phacelia  scariosa  is  apparently  related  to 
P.  pedicellata  and  is  discussed  under  that 
species. 


Fif,'.  53.     Phacelia  scariosa  T.    S.   Brandegee. 
Wiggins  7887   (us). 


June  1975 


ATWOOD:    PHACELIA  CRENULATAE  GROUP 


181 


1884,  Lemmon  and  wife  (uc,  uc.  us);  Sun- 
set Mountain,  Flagstaff.  21  August  1915,  H. 
Rusby  s.n.   (ny). 

Phacelia  macdougalii  Heller  in  Brand.  Das 
Pflanzenreich  IV.  251:80.     1913.  as  synonym. 

Annual  plants,  1-3.4  dm  tall;  stems 
erect,  simple  or  branched  at  the  base,  with 
simple    setose,    hirsute    and   multicellular 


Fig.  54.     Dorsal    and    ventral    view    of    the 
seeds  of  P.  scariosa  Brandegee.  I.  Johnston  3884 

(c.^s). 


*^^ 


:Sj^ 


Map  25.     Baja  California  and  Sonora.  Mexi- 
co.   Distribution  of  P.  scariosa  Brandegee. 

31. Phacelia  serrata  Voss 
Fig.  55 

Phacelia  serrata  Voss.  Bull.  Torr.  Bot.  Club 
64:88-89.  1937.  Holotype:  Arizona:  Coco- 
nino Co.:  San  Francisco  Mountains,  May- 
October  1900,  C.  A.  Purpus  8064  (pom!); 
Isotypes  (uc,  us).  Paratypes:  Arizona:  Coco- 
nino Co.:  vicinity  of  Flagstaff,  15  July  1898. 
Macdougal  288  (ny,  rm,  us,  us);  volcanic 
scoria,    San   Francisco   Mountains,   September 


Fig.  55.     Phacelia  serrata  Voss.  J.  T.  Howell 
and  G.  True  45184  (c.\s). 


182 


GREAT  BASIN  NATURALIST 


Vol.  35,  No.  2 


glandular  hairs;  leaves  lanceolate,  serrate, 
dentate  to  shallowiy  lobed,  1.4  cm  long, 
0.5-2  cm  wide,  with  a  basal  leaf  cluster, 
especially  when  young,  gradually  reduced 
upward,  the  upper  sessile  or  nearly  so,  the 
lower  with  a  petiole  up  to  1.5  cm  long, 
setose,  hirsute  and  glandular;  inflores- 
cence of  compound  scorpioid  cymes,  se- 
tose, puberulent  with  multicellular  stip- 
itate  glands,  pedicels  up  to  1  mm  long; 
sepals  elliptical  to  oblanceolate,  more  or 
less  keeled  at  the  base,  3.5-6.5  mm  long, 
1.5-2  mm  wide,  setose,  puberulent  and 
stipitate-glandular;  corolla  rotate  (appear- 
ing tubular  in  some  pressed  specimens), 
blue  to  light  violet,  3-4  mm  long  and 
broad,  pubescent;  stamens  and  style  ex- 
serted,  style  bifid  3/4  its  length,  lower 
1/4  pubescent;  capsule  subglobose,  2.8- 
3.5  mm  long,  2-2.5  mm  wide,  glandular 
and  puberulent;  mature  seeds  4,  elliptical 
to  oblong  (sometimes  unequally  so  when 
one  of  the  margins  is  involute),  dark 
brown,  3-3.2  mm  long,  1-1.3  mm  wide, 
excavated  and  divided  by  a  prominent 
ridge,  the  ridge  corrugated  on  one  side, 
the  margins  corrugated,  pitted  and  often 
one  or  both  involute,  dorsal  surface  smooth 
and  shiny  to  somewhat  dull,  sometimes 
faintly  pitted,  the  tip  and  margins  darker 
for  part  of  their  length  (or  at  least  dif- 
ferent in  appearance  from  the  rest  of  the 
dorsal  surface).  Fig.  56.  Collections:  15 
(2);  representative:  .1.  Hill  s.n.  (us);  L. 
Goodding  1526  (uc);  D.  Atwood  2586 
(bry);  D.  Dunn  12644  (rsa);  H.  Hansen 
615  (rm);  J.  Howell  and  G.  True  (bry. 
cas)  . 

Habitat. —  Confined  to  volcanic  scoria 
slopes  of  open  yellow  pine  forest  and  ju- 
niper flats  and  hills.  Flowering  from  late 
June  to  mid-September.  5,900  to  7,150 
feet. 

Distribution. —  In  and  around  Sunset 
Crater  National  Monument  and  San 
Francisco  Mountains  north  of  Flagstaff. 
Coconino   Co.,  Arizona. 

The  relationships  of  this  entity  to  other 
taxa  in  this  group  are  questionable  at  the 
present  time.  Part  of  the  paratype  ma- 
terial cited  by  Voss  (1937)  belongs  to 
P.  palmeri.  The  specimen  in  question  is 
Palmer  335  (gh,  ny)  and  was  probably 
collected  in  southern  Utah  or  the  extreme 
northern  part  of  Arizona  in  Mohave  Co. 
The  label  bears  the  data  southern  Utah- 
northern    Arizona.    Brand     (19H)     inad- 


vertently listed  P.  macdougalii  as  a  syno- 
nym of  P.  integrifoUa  Torr.  even  though 
it  was  only  a  manuscript  name  and  had 
never  been  published.  I  have  seen  the 
specimens  in  question  and  conclude  that 
they  are  referable  to  P.  serrata  Voss.  Voss 
(1937)  described  P.  serrata  but  failed  to 
mention  P.  macdougalii. 

32.  Phacelia  splendens  Eastwood 
Fig.  57 

Phacelia  splendens  Eastwood.  Zoe  4:9.  1893. 
Holotype:  Colorado:  Mesa  Co.:  Grand  Junc- 
tion, 19  May  1892.  A.  Eastwood  s.n.  (cas?); 
Isotypes  (uc.  uc,  us.  sketch  at  ny). 
Phacelia  glandulosa  Nutt,  subsp.  splendens 
(Eastwood)  Brand.  Das  Pflanzenreich  IV, 
251:83.     1913. 

Plants  annual,  0.5-2.7  dm  tall;  stems 
erect,  simple  or  branched  leafy,  puber- 
ulent, with  scattered  stipitate-glandular 
hairs;  leaves  pinnatifid,  2-7.5  cm  long,  0.7- 
4  cm  wide,  petiolate,  leaf  blade  essen- 
tially glabrous  (pubescent  only  on  the  pet- 
iole and  rachis  or  lower  portion  of  the 
pinnae);  inflorescence  terminal  on  each 
branch  and  the  main  stem,  cymes  compact 
and  densely  flowered,  pedicels  short  but 
lengthening  to  as  much  as  1.7  mm  in  fruit, 
slightly  more  pubescent  than  the  stem; 
sepals  linear  to  narrowly  oblanceolate,  2.5- 
3  mm  long  in  flower,  4-4.4  mm  long  in 
fruit,  0.6-1  mm  wide,  hirsute  and  with  a 
few  scattered  glandular  hairs;  corolla  cam- 
panulate,  the  lobes  bright  blue,  the  tube 
yellowish,  4-8  mm  long  and  broad,  glab- 
rous to  sparsely  pubescent;  stamens  and 
style  exserted  7-11  mm,  the  filaments 
blue,  anthers  yellow,  style  bifid  ca.  2/3 


Fig.  56.     Dorsal  and  ventral  view  of  the  seeds 
of    P.    serrata    Voss.    J.    T. -Howell    and    G.    True 

45184   (bky). 


ATWOOD:    PHACELIA   CRENULATAE   CROUP 


183 


Fig.  57.     Phacelia     splendens     Eastwood     R. 
Barneby  12797   (cas). 


feet  elevation.  Commonly  associated  with 
Atriplci\  but  in  Mesa  Verde  National 
Monument  it  was  collected  in  pinyon- 
juniper.   Mid-May  to  mid-July. 

Distribution. —  Known  only  from 
western  and  southwestern  Colorado  and 
northwestern  New  Mexico  (Map  26). 

This  species  is  related  to  P.  corrugata 
and  P.  utahensis  and  may  be  a  link  be- 
tween the  two  complexes.  It  can  be  dis- 
tinguished from  the  former  by  its  nearly 
glabrous  and  more-divided  leaves,  yellow- 
ish corolla  tube,  and  different-textured 
and  less-corrugated  seeds.  From  the  latter, 
it  differs  in  having  a  shorter  and  less- 
glandular  indument,  a  less-robust  and  less- 
branched  habit,  and  different  seeds.  The 
seeds  were  reported  by  both  Eastwood 
(1893')  and  Voss  (1937)  as  lacking  cor- 
rugations. Observations  of  mature  seeds 
of  P.  splendens  demonstrate  that  there 
definitely  are  evident  corrugations  on  one 
side  of  the  ridge  and  sometimes  on  one  of 
the  incurved  margins. 

33.  Phacelia  utahensis  Voss 

Fig.  59 

Phacelia  utahensis  Voss.  Bull.  Torr.  Bot.  Club 
64:135.  1937.  Holotype:  Utah  Sanpete  Co.: 
Gunnison.  7  ,Iune  1910,  M.  Jones  s.n.   (pom). 

Plants  stout,  erect  annuals,  0.8-5.8  dm 
tall;  stems  usually  simple,  sometimes 
branched  at  the  base,  brow'nish  to  yellow^- 
ish,  densely  glandular  and  finely  pubes- 
cent; leaves  linear  to  narrowly  lanceolate, 
strigose  to  ciliate  on  the  margins,  wdth 
scattered  glands  (especially  the  upper), 
1.5-12  cm  long,  0.5-1.5  cm  wide,  the  mar- 


its  length,  the  undivided  portions  puber- 
ulent  and  glandular;  capsule  subglobose, 
4-4.5  mm  long,  3-3.5  mm  long,  1.5  mm 
wide,  finely  favose,  the  ventral  surface  ex- 
cavated on  both  sides  of  the  ridge,  the 
ridge  with  evident  corrugations  on  one 
side,  the  margins  more  or  less  revolute 
(Fig.  58).  Collections:  23  (4);  representa- 
tive: W.  Weber  7509  (cas,  rm,  rsa,  uc); 
D.  Atwood  2532  (bry)  ;  D.  Atwood  and 
L.  Higgins  1814  (bry);  L.  Iliggins  3302 
(bry,  wts);  S.  Welsh  756  (bry);  R.  Bar- 
nebv  12743  (cas,  ny,  rsa);  E.  Payson  671 
(gh). 

Habitat. —  Apparently  confined  to  the 
Mancos   Shale  formation,  4,500  to  6,000 


Fig.  58.  Dorsal  and  ventral  view  of  the 
seeds  of  P.  splendens  Eastwood.  D.  Atwood  and 
L.  Higgins  1814  (bry). 


184 


GREAT  BASIN  NATURALIST 


Vol.  35,  No.  2 


1 — ' 

N 

1^ 

,...../-'  r- 

/ 

— ^ -'            1 

I 

i 

1 

1*   .--f--. 

1 

<'        1 

V--^. 

—     •« 

y 

■*--\y*-i 

/' 

/^  -|           r- 

/ 

"v 

•/         r--^- 

•  / 

/' 

•  / 

•• 

Map  26.     Southwestern  Colorado  and  adjoin- 
ing states.    Disti'ibution  of  P.  splendens  Eastwood. 

gins  often  revolute,  crenate,  undulate  to 
irregularly  dentate,  basal  ones  petiolate 
and  dense,  the  upper  sessile,  aiu-iculate  to 
cordate;  inflorescence  thyrsoid,  u])  to  3.4 
dm  long,  often  with  a  few  lateral,  leafy 
inflorescence  branches  below,  stipitate- 
glandular  and  finely  pubescent,  cymes 
mostly  in  pairs,  (or  1-3),  up  to  4  dm 
long  in  fruit,  densely  flowered,  the  ped- 
icels, 1-1.5  mm  long;  sepals  oblanceolate, 
3-4  mm  long,  0.8-1.1  mm  wide,  glandular 
and  hirsute;  corolla  rotate  to  campan- 
ulate,  the  lobes  bluish  to  violet,  the  tube 
yellowish,  ca.  3-4  mm  long,  ca.  6  mm 
broad,  glabrous;  stamens  exserted  9-10 
mm,  filaments  violet,  anthers  yellow;  style 
exserted  ca.  10  mm,  bifid  3/4  its  length, 
the  lower  1/4  setose  and  glandular;  cap- 
sule globose  to  subglobose,  3.5-4.1  mm 
long,  2.6-3.5  mm  wide,  glandular  and 
setose;  mature  seeds  4,  elliptical,  dark 
(reddish),  the  dorsal  surface  faintly  pitted. 
the  ventral  surface  excavated  on  both 
sides  of  the  ridge,  often  lighter  than  the 
dorsal  surface,  pitied  with  the  markings 
in  the  excavations  longer  (transversely) 
than  those  of  the  ridge  or  margins,  the 
ridge  .sometimes  faintlv  corrugated  on  one 


'\!b/="  Y 


J/'*^. 


Fig.  59.     PhaccUa    utahcnsis    Voss.     D. 
wood  and  L.  Higgins  1624  (bry). 


At- 


side.  Collections:  22  (13);  representative: 
.1.  Howell  and  G.  True  44640  (bry,  cas)  ; 
L.  Higgins  1624  (bry);  D.  Atwood  1520, 
1893,  1835,  1684,  1892,  1894,  1895,  1526, 
1519,  1518,  1525.  1528,  1527  (bry). 

Habitat. —  Endemic  to  the  Arapian 
Shale  formation. 

Distribution. —  Sevier  and  Sanpete 
COS.,  Utah,  from  5,500  to  5,700  feet,  April 
to  June  (Map  27). 

34.  PhaccUa  vossii  Atwood 
Fig.  60 

PhaccUa  vossii  Atwood.  Rhodora  74(800)  :462- 
465.  1972.  Sierra  Madre  Oriental,  calcite 
and  limestone  hills  bej'ond  Pablillo  toward 
Santa  Clara,  ca.  15  mi  southwest  of  Galeana, 
18  July  1934,  C.  H.  Mueller  1075  (gh!); 
Isotype  (mexu).  Paratype:  Mexico:  Nuevo 
Leon:  Hacienda  Pablillo,  Galeana.  13  August 
1936.   M.   Taylor   156    (ny,   uc). 

Peremiial  ]>lants,  2.1-7.^  chn  high;  stems 
erect  from  a  woody  caudex.  with  hirsute  to 
setose  and  stipilatf^-ghuukdar  hairs;  leaves 


June  1975 


ATWOOD:    PHACELIA  CRENULATAE  GROUP 


185 


N 

\ 

Map  27.     Central  Utah  in  Sanpete  and  Sevier 
counties.    Distribution  of  P.  utahensis  Voss. 


linear  to  lanceolate,  2-11  cm  long,  0.5-2.2 
cm  wide,  revolute,  ventral  surface  heavily 
glandular,  dorsal  surface  hirsute,  with  scat- 
tered stipitate-glandular  hairs,  margins 
dentate  to  irregularly  toothed,  petiolate, 
the  petiole  up  to  1.5  cm  long;  inflorescence 
axillary  to  terminal,  scattered  along  the 
stems  for  as  much  as  1/2  its  length  or 
less,  consisting  of  simple  to  compound 
scorpioid  cymes,  individual  cymes  up  to 
7.5  cm  long  in  fruit,  pedicels  up  to  2  mm 
long  in  fruit,  glandular  and  hirsute;  co- 
rolla tubular  to  short  campanulate,  pale 
lavender  (rarely  white),  5.5  mm  long; 
sepals  oblanceolate  to  spatulate,  4.6  mm 
long,  1.5-2  mm  wide,  glandular  and  hir- 
sute; stamens  exserted,  filaments  purple, 
anthers  greenish  blue;  style  exserted  ca. 
2  nun  longer  than  the  stamens  ca.  9  mm 
long,  bifid  for  3/4  its  length,  lower  1/4 
pubescent;  capsule  ovate,  3.7  min  long, 
2.5-2.7  mm  wide,  glandular  and  strigose; 
inmiature  seeds  4,  elliptical  to  oblong, 
brown,  2.5-3.1  mm  long,  1-1.4  mm  wide. 
Collections:  4  (1);  representative:  known 
only  from  the  type  collections  and  D.  At- 
wood  and  ,1.  Reveal  5985,  25  September 
1973   (bry). 


Fig.  60.     Phacelia  vossii  Atwood.  G.  Mueller 
1075    (me.xu). 


186 


GREAT  BASIN  NATURALIST 


Vol.  35,  No.  2 


Fig.  61.     Phaceiia  welshii  Atwood.    D.  Atwood  2605   (bry). 


June  1975 


ATWOOD:    PIIACF.LIA   CRENULATAE  (IROUP 


187 


1  Iabitat. —  Apparently  endeiriic  to  cal- 
(itc  and  limestone  soils. 

Distribution. —  Known  only  from  the 
type  locality  near  Galeana  in  the  state 
of  Nuevo  Leon,  Mexico. 

The  corollas  of  the  type  specimens  have 
faded  to  white,  which  is  a  common  oc- 
currence in  several  species  of  this  group. 

This  taxon  appears  to  be  related  to 
/•*.  pinnatifida  Griseb.,  which,  according  to 
Brand  (1913),  occurs  m  the  Andes  of 
South  America  in  Peru,  Bolivia,  and  Ar- 
gentina. 

35.  Phacelia  ivelshii  At  wood 
Fig.  61 

Phacelia  welshii  Atwood.  Rliodora  74(800) :465- 
468.  1972.  Holotj'pe:  Arizona:  Coconino 
Co.:  along  highway  89  iust  north  of  Gray 
Mountain,  19  May  1970.  D.  Atwood  2608 
(bry!);  Isotypes  (.ariz,  asc,  b,  bry,  c.-\s,  colo, 

DIXIE,   GH.   NY,    POM,   RM,   RS.\,    US,    US,    UT,    UTC, 
WSC,    WTS). 

Annual,  1-5.5  dm  tall;  stems  more  or 
less  yellowish  to  green,  simple  or  branched, 
leafy,  hirsute  and  densely  co\ered  with 
multicellular  stipitate  glands;  leaves  ob- 
long to  lanceolate,  1.5-8  cm  long,  0.5-2.7 
cm  wide,  hirsute  and  densely  glandular, 
the  margins  often  revolute,  undulate  and 
dentate,  the  basal  leaves  clustered,  petio- 
late,  the  petiole  2  cm  long  or  less,  cauline 
leaves  sessile  or  nearly  so,  often  cordate 
at  the  base;  inflorescence  of  compound 
scorpioid  cymes,  these  terminal  at  the 
ends  of  the  main  stem  and  lateral 
branches,  densely  glandular  and  hirsute, 
the  individual  cymes  congested,  but  loos- 
ening in  fruit,  up  to  10  cm  long,  pedicels 
up  to  1.5  mm  long;  sepals  spatulate  to 
oblanceolate,  3.5-4.5  mm  long,  0.7-1.7  mm 
wide,  hirsute  and  stipitate-glandular;  co- 
rolla campanulate,  purplish  to  blue,  5-6 
mm  long  and  broad,  pubescent;  stamens 
exserted  ca.  8-10  mm,  filaments  the  same 
color  as  the  corolla,  anthers  yellow;  style 
exserted  ca.  8  mm,  bifid  3/4  its  length, 
the  lower  1/4  setose  and  glandular;  cajj- 
sule  oval,  3-3.2  mm  long,  2.9-3.1  mm  wide, 
hirsute  and  glandular;  mature  seeds  4. 
oblong,  brown  ventrallv  to  reddish  dor- 
sally,  pitted,  2.8-3.4  mm  long,  1.3-1.5  mm 
wide,  the  ventral  surface  lighter  than  the 
dorsal  surface,  the  ridge  corrugated  on  one 
side,  the  margins  corrugated  and  more  or 
less  revolute.    Collections:    15    (6);  repre- 


sentative:   D.    Atwood   2591    (bry.   cas); 
J.  Howell  24397   (c..\s) ;  D.  Atwood  2598 

(aRIZ,   B,    bry,    CAS,    COLO,    DIXIE,    GH,    NY)  ; 

D.  Demaree  43982  (uc);  D.  Atwood  2601 
(asc.  bry.  c.\s.  lts,  wsc). 

Habitat. —  Red  shale  formation. 

Distribution. —  Coconino  Co.,  Arizona. 

Phacelia  welshii  is  probably  most  closely 
related  to  P.  utahensis  and  P.  corrugata. 
It  can  be  distinguished  from  the  former  by 
its  broader,  dark  brown  seeds,  long  (up 
to  1.3  mm  long),  flattened,  multicellular, 
stipitate  glands,  and  broadly  lanceolate 
leaves.  The  more  open  inflorescence,  cor- 
rugated seeds,  and  broader,  shorter, 
densely  glandular  leaves  easily  separate 
P.  welshii  from  P.  corrugata. 


Appendix  I 

Synonyms 

The  following  is  a  list  of  synonyms  in  the 
Phacelia  Crenulatae  group.  The  names  in  the  left 
column  are  the  synonyms,  and  the  number  to  the 
right  is  the  reference  to  the  numbered  taxa  in 
the  present  treatment. 

EUTOCA 

E.  glandulosa  Hook 17 

PHACELIA 

P.  arenicola    Brandegee    19a 

P.  conferta    D.     Don    .■ 10 

P.  congesta  yar.   dissecta  Gray  10 

P.  congesta  yar.  rupestris  (Greene)  Macbride  29 

P.  congesta  yar.   typica  Voss  10 

P.  dissecta  (Gray)    Small  10 

P.  crenulata  yar.  amhigua   (Jones)    Macbride  3a 

P.  crenulata  var.  bakeri  Biand  7 

P.  crenulata  yar.  corrugata  (Nels.)   Bi-and  ....  12 

P.  crenulata  yar.   funerea  Voss   in  Munz   ....  14b 

P.  crenulata  yar.  minutiflora  (Voss)   Jeps 3b 

P.  crenulata  yar.   vulgaris  Brand  14b 

P.  depauperata  W.   &   S  26 

P.  deserta   Nels 17 

P.  foetida  Goodding  24 

P.  glandulosa  Gray  in  Brand,  pro  syn 26 

P.  glandulosa    Hemsley    13 

P.  glandulosa    ssp.    eu-glandulosa   Brand   var. 

australis  Brand,  in  part  15 

P.  glandulosa   ssp.    eu-glandulosa   Brand   yar. 

australis  Brand,   in  part  7 

P.  glandulosa    ssp.    eu-glandulosa    Barnd   yar. 

deserta   Brand    17 

P.  glandulosa   ssp.    eu-glandulosa   1 

P.  glandulosa      ssp.      splendens      (Eastwood) 

Brand 32 

P.  glandulosa  yar.  neornexicana  (Thurber  ex 

Torr.)    Gray    22 

P.  intcgrifolia     var.     arenicola     (Brandegee) 

Brand  ..._ 19a 

P.  intcgrifolia    var.    palmeri     (Torr.    ex 

/Wats.)    Gray   24 

P.   integrifolia   var.    rohusta   Macbr 28 

P.  intermedia  Wooton,  in  part  9 

P.  invenusta  Gray   9 


188 


GREAT  BASIN  NATURALIST 


Vol.  35,  No.  2 


P.  macdougalli  Heller  in  Brand,  pro.  syn 31 

P.  neomezicana  var.  alba  (Rydberg)  Brand  ..  1 
P.  neomezicana     var.     coulteri     (Greenman) 

Brand    -  ■-  13 

P.  neomezicana  var.    coulteri   subvar.   folisis- 

sima    Brand   1 

P.  neomezicana   var.    eu-neomezicana    Brand  22 

P.  neomezicana  var.  microphylla  Bi-and  15 

P.  palmeri   var.    typica   Voss    24 

P.  petiolata  Johnston  23 

P.  popei  var.   arizonica    (Gray)    Voss   6 

P.  popei  var.  similis  (W.  &  S.)  Voss  26 

P.  popei  var.  typica  Voss  26 

P.  similis  W.  &  W 26 

P.  tenuipes   W.    &   S 8 

P.  tezana  Voss  19b 


Appendix  II 

Glossary 

Alveolate.  Honeycombed;  pits  in  the  surface  of 
the  seed. 

Auriculate.   With  earlike  appendages. 

Corrugated.    Wrinkled  or  folded. 

Cymbiform.    Boat  shaped. 

Cyme.  A  detenninate  flov^er  cluster  in  which  the 
first  flower  is  terminal  on  the  main  axis  and 
the  central  flowers  open  first. 

Denticulate.    Slightly  and  finely  toothed. 

Favose.  Honeycombed;  pits  in  the  surface  of  the 
seeds. 

Fimbriate.  Fringed  with  elongate,  slender  pro- 
cesses or  lobes  on  the  margins  of  the  corolla 
lobes. 

Geminate.   In  pairs,   as  regarding  the  seeds. 

Glandular.  A  globose-secreting  stinicture  borne 
on  the  surface  and  estipitate. 

Gypsiferous.    Containing  gypsum. 

Hirsute.   Pubescent  with  stiff,  coarse  hairs. 

Hispid.  Pubescent  with  long,  very  stiff  hairs, 
these  able  to  penetrate  the  skin. 

Pilose.  Pubescent  with  soft,  slender  hairs  point- 
ing  the   same   direction  as   if   combed. 

Pitted.    Having  little  depressions  or  pits. 

Puberulent.  Pubescent  with  very  short  hairs, 
not  stiff. 

Reticulate.    Net-veined. 

Revolute.  Rolled  backward  from  both  margins, 
toward  the  inside. 

Scabrous.  Rough  to  the  touch  owing  to  the  pres- 
ence of  short  stiff  hairs. 

Scarious.    Thin,  dry.  and  membranous,  not  green. 

Scorpioid.  A  unilateral  inflorescence  circinately 
coiled  in  bud  and  anthesis. 

Setose.  Pubescent  with  short,  rather  stiff  hairs, 
these   not   able   to   penetrate   the   skin. 

Stipitate-glandular.  A  globose,  stipitate,  secre- 
tory structure  borne  on  the  surface  of  vegeta- 
tive parts. 

Strigose.  Pubescent  with  short,  straight  appressed 
hairs. 

Tuberculate.    Having   small    knoblike    projections. 

Villous.  Pubescent  with  long  and  weak,  tangled, 
but  not  matted,  hairs. 


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RODENT  POPULATIONS,  BIOMASS,  AND  COMMUNITY 

RELATIONSHIPS  IN  ARTEMISIA  TRIDENT  AT  A. 

RUSH  VALLEY,  UTAH 

D.  W.  Nichols\  H.  D.  Smith"',  and  M.  F.  Baker' 

Abstract. —  Three  desert  Artemisia  tridentata  communities  in  Rush  Valley.  Utah,  were  trapped 
for  small  rodents  during  the  summer  of  1970,  and  population  densities  were  estimated  for  each  popula- 
tion category  using  Lincoln's  index.  Animals  were  weighed  and  rodent  biomass  calculated  by  species 
throughout  the  summer.  Population,  biomass.  and  other  data  were  then  analyzed  to  gain  an  under- 
standing of  the  community  relationships  of  the  three  study  areas  to  each  other  as  well  as  to  the 
A.  Iridentata  community  types  of  the  Great  Basin. 

Peromyscus  maniculatus.  Eutamias  minimus,  and  Reithrodontomys  megalolis  were  common  to 
area  1,  whereas  P.  maniculatus.  E.  minimus,  and  Perognathus  parvus  were  common  to  areas  2 
and  3.  The  peak  estimated  standing  crops  were  182.8  (74.0).  143.1  (57.9),  and  129.7  g/acre  (52.5 
g/ha)  for  areas  2,  1,  and  3  respectively.  The  population  and  biomass  of  area  2  peaked  in  midsum- 
mer,  area    1    early   summer,   and   area   3   late   summer. 


Introduction 

Big  sagebrush  {Artemisia  tridentata 
Nutt.)  covers  an  estimated  total  area  of 
226,364  square  miles  (586,283  km-)  in 
the  Great  Basin  and  associated  areas  of 
the  western  United  States  (Beetle,  1960) 
and  is  the  most  abundant  plant  species 
over  much  of  this  area.  According  to  Hiro- 
naka  (1963),  ''A.  tridentata  has  the  widest 
distribution  of  all  the  sagebrushes  and  oc- 
curs across  the  entire  moisture  gradient 
of  the  sagebrush  zone."  Passey  and  Hugie 
(1962)  found  A.  tridentata  occupying  a 
greater  number  of  soil  types  than  any 
other  sagebrush  species. 

A.  tridentata  is  ecologically  significant 
in  that  it  provides  not  only  food  and  cover 
for  some  species  but  competes  against 
other  desirable  food  and  cover  species. 
For  economic  reasons,  however,  many  peo- 
ple consider  sagebrush  to  be  a  highly  un- 
desirable plant.  As  a  result,  much  re- 
search done  in  the  sagebrush  community 
has  been  directly  concerned  with  control- 
ling its  spread  and  decreasing  its  abun- 
dance. Treatment  resulting  from  such  re- 
search has  sometimes  been  temporarily 
effective.  Some  treated  areas  after  14 
years  may  have  more  sagebrush  on  them 
than  adjacent  untreated  areas  (Johnson, 
1969).  The  mean  useful  life  cycle  of 
spraying  projects  throughout  Wyoming 
has  been  estimated  to  be  about  15  years 
(Kearl,  1965). 

The  economic  importance  and  ecolog- 
ical impact  of  such  control  measures 
make  research  leading  to  an  understand- 


ing of  the  A.  tridentata  community  im- 
perative. The  objective  of  this  study  is  to 
establish  baseline  data  by  estimating  the 
comparative  small  rodent  density  and  bio- 
mass of  A.  tridentata  communities  in  low- 
intermediate-  and  high-altitude  desert 
areas  of  the  Great  Basin  and  using  this 
estimation  to  compare  the  three  commun- 
ities. 

Rush  Valley,  Utah,  was  chosen  for  the 
area  of  research  because  (1)  it  is  a  desig- 
nated grazing  research  area  of  the  Inter- 
mountain  Forest  and  Range  Experimental 
Station,  which  funded  the  project;  (2)  a 
large  part  of  the  valley  is  covered  by 
A.  tridentata;  (3)  the  data  gathered  will 
augment  that  of  current  and  past  research 
in  the  valley;  and  (4)  the  data  collected 
will  aid  future  studies  and  management 
of  the  valley. 

No  literature  relating  rodent  density 
with  biomass  or  energy  flow  has  been 
published  concerning  sagebrush  commun- 
ities. There  are  works,  however,  that  have 
been  reported  for  other  terrestrial  com- 
munities. Densities  of  rodents  have  been 
studied  in  Rush  Valley,  Utah,  primarily 
in  piny  on- juniper  and  reseeded  areas 
(Baker,  1969).  Woodbury  (1955)  re- 
ported on  the  small  mammal  distribution 
in  Cedar  Valley  which  borders  Rush 
Valley  on  the  east.  Vest  (1962)  reported 
on  the  small  mammal  distribution  in  Dug- 
way  Valley  which  borders  Rush  Valley 
on  the  west.  Rodents  of  •sagebrush  com- 
munities in   both  valleys  were  discussed. 

Although  literature  concerning  biomass 
in    A.    tridentata   communities    is    sparse, 


^Department  of  Biology,  Northland  Pioneer  College,  Show  Low,  Arizona. 

^Department  of  Zoology,  Brigham  Young  University,  Provo,  Utah. 

^Wildlife  biologist,  Intemiountain  Forest  and  Range  Experiment  Station,  Provn,  Utah. 


191 


192 


GREAT  BASIN  NATURALIST 


Vol.  35,  No.  2 


much  descriptive  material  is  available  on 
the  plant  and  its  community  type.  A  mor- 
phological life  history  of  A.  tridentata  was 
written  by  Diettert  (1938),  and  Beetle 
(1960)  published  a  taxonomic  and  distrib- 
utional study  of  all  the  north  American 
sagebrush  taxa.  Other  references  concern- 
ing A.  tridentata  communities  in  Utah 
may  be  found  in  Christensen  (1967). 

Materials  and  Methods 
Study  Areas 

Three  14.5  acre  (5.87  ha)  study  areas 
located  in  the  southern  part  of  Rush 
Valley,  Tooele  County,  Utah,  were  selec- 
ted and  are  described  in  Table  1.  Area 
1,  elevation  5,100  feet  (1,554  m),  is  lo- 
cated in  the  southwest  quarter  of  Section 
10,  Township  7  south.  Range  5  west, 
Tooele  County,  Utah.  It  contains  irreg- 
ularly scattered  A.  tridentata  interspersed 
with  Chrysothamnus  puberulus,  the  grass 
Distichlis  striata,  and  bare  ground.  Dis- 
tichlis  striata,  an  indicator  of  alkaline  soil 
conditions,  is  abundant  in  a  number  of 
small  areas  with  poor  drainage  as  are 
numerous  A.  tridentata  plants  1-3  inches 
(2.5-7.6  cm)  tall.  This  short  sagebrush  is 
possibly  stunted  by  an  accumulation  of 
soil  salts.  The  area  has  a  wash  1-2  feet 
(0.3-0.6  m)  deep  and  five  feet  (1.5  m) 
wide  that  runs  across  the  west  side. 

Area  2,  elevation  5,700  feet  (1,737  m) 
is  located  in  the  northwest  quarter  of  Sec- 
tion 35,  Township  8  south,  Range  6  west, 
Tooele  County,  Utah,  and  is  characterized 
by  scattered  A.  tridentata  with  some  Sar- 
cobatus     vermiculatus     and     much     bare 


ground.  The  west  boundary  of  this  area 
is  a  dirt  road,  beyond  which  there  is  an 
extensive  stand  of  S.  vermiculatus  mixed 
with  the  grass  Agropyron  cirstatuni. 

Area  3,  elevation  6,500  feet,  (1,981  m) 
is  located  in  the  west  half  of  Section  4, 
Township  9,  Range  5  south,  Tooele 
County,  Utah,  and  is  covered  with  irreg- 
ularly scattered  A.  tridentata  with  a  dense 
understory  of  a  perennial  lupine  and  var- 
ious grasses,  principally  Agropyron  dasy- 
stachyuni  and  Sitanion  Jiystrix.  There  is 
little  bare  ground  except  on  the  east  and 
west  borders  which  were  exposed  when 
adjacent  land  was  cleared  of  A.  tridentata 
and  some  Juniperus  osteosperma  in  the 
fall  of  1969. 

Vegetation  sampling  on  all  areas  con- 
sisted of  measuring  (1)  absolute  ground 
cover,  (2)  percent  species  cover  compo- 
sition, (3)  frequency  and  density  of  pe- 
rennial species  other  than  grasses,  (4)  fre- 
quency of  all  grasses,  both  annuals  and 
perennials,  lumped  together,  and  (5)  fre- 
quency of  all  annuals,  except  grasses, 
lumped  together.  A  modified  line-point 
method  of  sampling  was  used  to  determine 
cover,  and  small  quadrats  were  used  for 
frequency  and  density  data  collection 
(Cain  and  Castro,  1959).  These  data  are 
available  in  Nicholes    (1972). 

Trapping 

Each  of  the  quadrat  study  areas,  14.5 
acres  (5.87  ha),  was  equally  divided  into 
25  squares  with  a  trap  station  located  in 
the  center  of  each  square  where  three 
Sherman      aluminum     live     traps     were 


Table  1.     General  comparisons  of  the  study  areas. 


Characteristics 


Area  1 


Area  2 


Area  3 


^Soil,    TYPE 


Slope 

Water  runoff 

Erosion 
^Climate 

Mean  annual  temp. 

Mean  annual  pupc. 
Principai.  cover 


Principal  rodents 


Elev.\tion 


deep  silt-clay  alkali 
soils  of  the  arid  &  semi- 
arid  valley  bottoms 
0-2% 
slow 
slight  to  moderate 

51   F 

8-10  inches 
Artemisia  tridentata 
Chrysothamnus  puberulus 
Distichlis  stricta 
Peromyscus  maniculatus 
Rutamias  minimus 
Reithrodontomys  megalotis 
5.100  ft.   (1,554  m) 


deep  silt-loam  soils 

of  the  semiarid 

valley  bottoms 

1-5% 

slow  to  medium 

high 


deep,  loamy,  dry  soils  of 

the  dry  subhumid  alluvial 

fans 

1-25%,  most  less  tlian  10% 

slow  to  medium 

moderate 


48  F  45-47  F 

8-12  inches  12-15  inches 

Artemisia  tridentata  Artemisia  tridentata 

Sarcobatus  vermiculatus  Lupine  sp. 

various  grasses  various  grasses 

Peromyscus  maniculatus  Peromyscus  maniculatus 

Rutamias  minimus  Rutamias  minimus 

Perognathus  parvus  Perognathus  parvus 

5,700  ft.  (1.737  m)  6.500   ft    (1.981    m) 


^(Harvey  and  Woodward,   1969) 


June  1975 


NICHOLS   ET    AL.:    RODENT   POPULATIONS 


193 


placed.  The  trapping  stations  were  160 
feet  (48.8  ni)  apart.  Rolled  oats  were 
used  as  bait.  The  traps  were  set  in  the 
afternoon,  checked  each  morning,  and 
closed  until  the  afternoon  resetting.  Each 
trapping  period  covered  five  consecutive 
nights,  every  other  week  from  2  June  to 
14  August  1970  (Table  2).  The  three 
areas  were  trapped  simultaneously  for 
six  trapping  periods.  To  offset  bias  prior 
to  each  trapping  period,  the  sequence  in 
which  each  of  the  three  areas  would  be 
checked  and  reset  was  determined  ran- 
domly. This  sequence  was  maintained 
throughout  a  trap  }:)eriod. 

At  the  end  of  six  trapping  periods  each 
area  was  "kill  trapped"  to  compare  with 
live  trapping  success  during  the  previous 
periods.  A  "kill  trapping"  consisted  of  one 
night  of  live  trapping  with  one  live  trap 
set  at  each  of  the  regular  trapping  sta- 
tions, one  live  trap  ])laced  at  the  corners 
of  each  square,  and  one  live  trap  placed 
at  the  middle  of  each  side  of  the  squares, 
for  a  total  of  121  traps.  Before  the  second 
trap  night  each  live  trap  was  replaced  by 
two  museum  special  snap  traps  for  a  total 
of  242  traps  per  quadrat.  This  trapping 
pattern  continued  three  to  four  nights 
until  the  number  of  previously  marked 
animals  caught  was  reduced  to  none  or 
nearly  none.  The  "kill  trapping"  did  not 
occur  simultaneously  for  each  area  be- 
cause of  the  large  number  of  traps  in- 
volved (Table  2). 

Animals  were  toe  clipped  for  identifica- 
tion. Data  recorded  for  each  individual 
animal  handled  during  the  study  included 
(1)  species,  (2)  sex,  (3)  age,  (4)  weight, 
(5)  trapping  station,  and  (6)  notes  con- 
cerning   the    animal's    general    condition, 

Table  2.     Schedule  of  trapping  periods. 


Areas 

'Dates 

1 

2 

3 

Live  Trapping 

1.     2-6  June 

X 

X 

X 

2.     15-19  June 

X 

X 

X 

3.     29  June-3  July 

X 

X 

X 

4.     13-17  July 

X 

X 

X 

5.     27-31  July 

X 

X 

X 

6.     10-14  Aug. 

X 

X 

X 

"Kill  Trapping" 

7.     24-27  Aug. 

X 

8.     31  Aug.-4  Sept. 

X 

9.     6-9  Sept. 

X 

^Dates   extend    from    the   first  mnming   traps   were  rhocked 
to  the  last  morning  traps  were  checked  during  a  trap  period. 


such  as  pregnancy,  parasitism,  injuries, 
and  others.  Age  classes  of  juvenile,  sub- 
adult,  and  adult  were  determined  primar- 
ily by  pelage  color  and  molt  patterns;  but 
the  appearance  of  genitals,  behavior  of 
animals,  and,  in  cases  where  age  is  ex- 
tremely difficult  to  determine,  i.e.,  chip- 
munks, the  weights  of  the  animals  were 
considered.  Animals  were  weighed  using 
a  spring-operated  scale  accurate  to  the 
nearest  0.5  g. 

Population  and  Biomass  Estimation 

For  each  species  caught  and  recaptured 
in  sufficient  numbers,  population  esti- 
mates were  made  at  the  end  of  each  trap- 
ping period  using  Lincoln's,  Hayne's,  and 
Jolly's  indices  (Giles,  1969).  Estimates 
were  made  for  ( 1 )  the  total  species  pop- 
ulation, (2)  the  population  of  each  age 
class  within  the  species,  and  (3)  the  pop- 
ulation of  each  sex  within  the  species. 
Nichols  (1972)  presents  this  data.  Com- 
parison of  the  three  estimators  showed 
Lincoln's  and  Hayne's  to  be  similar,  but 
Lincoln's  estimates  were  used  in  the  bio- 
mass calculations.  Population  estimates 
for  "kill  trapping"  periods  were  made 
after  the  first  two  nights  of  trapping, 
since  two  nights  of  "kill  trapping"  may 
have  caused  abnormal  immigration  into 
the  areas.  All  other  population  estimates 
were  calculated  using  five  days  of  live 
trapping  data. 

Species  biomass  was  calculated  by  mul- 
tiplying the  mean  species  weight  by  the 
estimated  population  number  of  that 
species  for  each  trapping  period.  Previous 
experience  had  shown  that  animals  re- 
peatedly caught  during  a  trapping  period 
tended  to  lose  weight,  likely  due  to  trap- 
ping stress.  To  compensate  for  this,  the 
mean  weights  were  calculated  two  differ- 
ent ways:  (1)  using  only  the  weight  of 
an  animal  taken  the  first  day  during  a 
given  trapping  period  and  (2)  using  the 
weight  of  an  animal  taken  every  day 
during  its  captivity  for  a  given  trapping 
period.  The  greatest  of  these  mean 
weights  for  any  population  category  was 
used  in  this  study.  In  most  instances 
method  one  was  used. 

Results 
Four  species  of  rodents  were  recaptured 
in  sufficient  numbers  to  be  considered  in 


194 


GREAT  BASIN  NATURALIST 


Vol.  35,  No.  2 


detail,  but  only  three  were  prevalent  in 
any  given  area.    These  species  were — 

Area  1    Area  2    Area  3 
Peromyscus   maniculatus         XXX 
Pcrognathus  parvus  X  X 

Reithrodontomys  megalotis     X 
Eutamias  minimus  XXX 

The  Lincoln  population  estimate  for 
each  species  caught  during  each  trapping 
period  and  study  area  is  given  in  Table 

3.  The  combined  total  number  of  rodents 
estimated,  of  all  species  considered,  for 
each  area  is  graphically  shown  in  Figure 
1.  The  following  is  a  general  breakdown 
of  each  area's  trapping  according  to  spe- 
cies. Detailed  charts  and  tables  of  data 
are  given  in  Nichols   (1972). 

Area  1 

Peromyscus  maniculatus.  The  greatest 
number  of  P.  maniculatus,  47  (Lincoln's 
estimate  50),  appeared  during  trap  period 
3  with  a  male-female  ratio  of  nearly  2:1. 
There  were  16  juveniles,  30  subadults, 
and  1  adult.  The  least  number,  24  (Lin- 
coln's estimate  25),  composed  of  3  ju- 
veniles, 17  subadults,  and  4  adults  with  a 
male-female  ratio  of  5:3,  appeared  during 
period  1.  The  greatest  mean  species 
weight,  19.6  g,  occurred  during  period  1 
and  the  least,  15.5  g,  during  period  5. 

Reithrodontomys  megalotis.  In  area  1 
R.  megalotis  was  captured  least  often  of 
the  main  species.  The  greatest  number, 
14  (Lincoln's  estimate  18),  appeared 
during  period  2  with  a  male-female  ratio 
of  3:4.  There  were  no  juveniles,  2  sub- 
adults,   and    12    adults.     During    periods 

4,  5,  and  6  no  mice  of  this  species  were 
caught.  The  greatest  mean  species  weight, 
13.9  g,  occurred  in  period  3  and  the  least, 
11.1  g,  in  period  2. 

Eutamias  minimus.  The  greatest  num- 
ber of  E.  minimus,  33  (Lincoln's  estimate 
33),  appeared  during  period  1  with  a 
male-female  ratio  of  approximately  3:2. 
There  were  16  juveniles,  14  subadults, 
and  3  adults.  The  least  number,  10  (Lin- 
coln's estimate  8),  com})osed  of  0  juv- 
eniles, 9  subadults,  and  1  adult  with  a 
male-female  ratio  of  nearly  1:1,  appeared 
during  period  6.  The  greatest  mean  spe- 
cies weight,  30.5  g,  occurred  during  period 
5  and  the  least,  28.4  g,  during  period  4. 


Area  2 

Peromyscus  maniculatus.  The  greatest 
number  of  P.  maniculatus,  49  (Lincoln's 
estimate  43),  appeared  during  trap  period 
6  with  a  male-female  ratio  of  nearly  3:2. 
There  were  5  juveniles,  38  subadults,  and 
6  adults.  The  least  number,  13  (Lin- 
coln's estimate  12),  composed  of  6  ju- 
veniles, 5  subadults,  and  2  adults  with  a 
male-female  ratio  of  5:8,  appeared  during 
period  2.  The  greatest  mean  species 
weight,  18.2  g,  occurred  during  period  5 
and  the  least,  15.3  g,  during  period  1. 

Perognathus  parvus.  In  area  2  P.  par- 
vus was  captured  least  of  the  three  main 
species.  Both  periods  5  and  6  yielded  the 
greatest  number,  10  (Lincoln's  estimates 
9  and  10),  with  male-female  ratios  of  7:3 
and  3:1.  The  respective  age  distributions 
were  0  and  2  juveniles,  9  and  7  subadults, 
1  and  1  adults,  llie  period  of  least  cap- 
ture was  period  1  when  one  adult  female 
was  captured.  The  greatest  mean  species 
weight,  18.7  g,  occurred  during  period  2 
and  the  least,  14.5  g,  during  period  3. 

Eutamias  minimus.  The  greatest  num- 
ber of  E.  minimus,  43  (Lincoln's  estimate 
46),  appeared  during  period  2  with  a 
male-female  ratio  of  approximately  4:3. 
There  were  20  juveniles,  16  subadults, 
and  7  adults.  The  least  number  of  chip- 
munks, 14  (Lincoln's  estimate  13),  com- 
posed of  1  juvenile,  12  subadults,  and  1 
adult  with  a  male-female  ratio  of  nearly 
4:1,  appeared  during  period  3.  The 
greatest  mean  species  weight,  31.1  g,  oc- 
curred during  period  5  and  the  least,  27.7 
g,  during  period  1. 

Area  3 

Peromyscys  maniculatus.  The  greatest 
nimiber  of  P.  maniculatus,  55  (Lincoln's 
estimate  49),  appeared  during  trap  period 
3  with  a  male-female  ratio  of  approx- 
imately 3:5.  There  were  16  juveniles,  31 
subadults,  and  8  adults.  The  least  num- 
ber, 29  (Lincoln's  estimate  26),  composed 
of  2  juveniles,  18  subadults,  and  9  adults 
with  a  male-female  ratio  of  3:4,  appeared 
during  period  1.  The  greatest  mean  spe- 
cies weight,  19.9  g,  occurred  during  period 
1   and   the  least,   17.5  g,  during  period  6. 

Perognathus  parvus.  The  greatest  num- 
ber of  P.  parvus,  47  (Lincohi's  estimate 
54),    appeared    during    period    6    with    a 


June  1975  nichols  et  al.:  rodent  populations 

Table   3.     Estimated   population   numbers   and    biomass   for   the   three   study    areas. 


195 


Trap  Period 


Lincoln's  N  no./ 14.5   acres 
(5.86  ha) 


Estimated   biomass   (g/14.5  acres) 
(5.86  ha) 


Species       Area  1 


Area  2 


Area  3        Area  1 


P.M. 
E.M. 
P.P. 
R.M. 

25 
33 

58 

P.M. 
E.M. 
P.P. 
R.M. 

44 
39 

18 

101 

P.M. 
E.M. 
P.P. 
R.M. 

50 
31 

1 

P.M. 
E.M. 
P.P. 
R.M. 

82 
48 
23 

71 

P.M. 
E.M. 
P.P. 
R.M. 

41 
5 

46 

P.M. 
E.M. 
P.P. 
R.M. 

35 

8 

43 

P.M. 
E.M. 

33 

12 

45 

P.M. 
E.M. 
P.P. 

P.M. 
P.P. 

112 


67 


101 


490.0 
943.8 


833.9 
488.4 
337.2 

825.6 


Area  2 


275.4 
941.8 


2650.4 


1067.5 

85.2 


1152.7 


Area  3 


517.4 
530.0 


51 

1433.8 

1217.2 

1047.4 

36 

752.4 

199.2 

698.4 

1123.2 

1311.0 

40 

199.8 

18.7 

836.0 

76 

2075.8 

1528.9 

1534.4 

49 

875.0 

283.2 

886.9 

914.5 

374.4 

29 

13.9 

29.0 

559.7 

78 

1803.4 

686.6 

1446.6 

55 

811.2 

434.2 

1078.0 

653.2 

848.4 

36 

148.0 

655.2 

91 

1464.4 

1430.6 

1733.2 

58 

635.5 

491.4 

1044.0 

2 

152.5 

746.4 

68.4 

38 

162.9 

767.6 

98 

788.0 

1400.7 

1880.0 

47 

591.5 

•  722.4 

822.5 

242.4 

1770.0 

54 

158.0 

1015.2 

1837.7 


705.2 

91.5 

794.3 

1591.0 


Key:     P.M. — Peromyscus    maniculatus;    E.iNI. — Eutamias    minim.i 


\\_y\—lirillnn,l.., 


male-female  ratio  of  approximately  7:5. 
There  were  4  juveniles,  21  subadults,  and 
22  adults.  The  least  number,  19  (Lin- 
coln's estimate  25),  composed  of  0  ju- 
veniles, 3  subadults,  and  16  adults  with  a 
male-female  ratio  of  nearly  2: 1  appeared 
during  period  1.  The  greatest  mean  spe- 
cies weight,  21.2  g,  occurred  during  period 
1  and  the  least,  18.2  g,  during  ]:)eriod  4. 


Eutamias  minimus.  In  area  3  E.  mini- 
mus was  captured  the  least  of  the  three 
main  species.  The  greatest  number,  6 
(Lincoln's  estimate  0),  appeared  during 
period  6  with  a  male-female  ratio  of  1:2. 
There  were  0  juveniles,  3  subadults,  and 
3  adults.  No  chipmunks  were  caught 
during  ])eriod  3.  The  weight,  39.0  g,  of 
one  adult  female,  the  only  animal  caught 


196 


GREAT  BASIN  NATURALIST 


Vol.  35,  No.  2 


during  period  1,  represents  the  greatest 
mean  species  weight.  The  least  mean 
species  weight  was  29.5  g  and  occurred 
during  period  3. 

Estimated  Small  Rodent  Biomass 

The  estimated  biomass  of  each  species 
for  every  period  and  study  area,  along 
with  total  biomass  per  area,  is  given  in 
Table  3.  In  addition  Table  3  lists  the 
Lincoln  population  estimates  used  in  the 
biomass  computations.  Kill-trap  data 
were  not  included  in  calculating  the 
greatest,  least,  and  mean  area  biomass  as 
given  in  the  following  description  of  ro- 
dent biomass  by  areas,  but  they  are  includ- 
ed in  Table  3.  The  kill-trap  biomass  for 
any  of  the  three  areas  fell  within  the 
limits  of  the  estimates  for  that  particular 
area  from  the  six  previous  trapping  pe- 
riods. 

Area  1.  The  greatest  estimated  small 
rodent  biomass,  143.1  g/acre  (57.0  g/ha), 
for  area  1  occurred  during  period  2.  The 
least,  54.3  g/acre  (22.0  g/ha),  occurred 
during  period  5.  The  mean  biomass  of 
area  1  over  the  six  trapping  ])eriods  was 
96.5  g/acre  (39.1  g/ha).  The  mean  was 
approached  during  periods  1  and  4.  Pe- 
riods 5  and  6  were  well  below  the  mean  as 
was  kill-trap  period  7. 

Area  2.  The  greatest  biomass,  182.8 
g/acre  (74.0  g/ha),  for  area  2  occurred 
during  period  6.  The  least,  47.4  g/acre 
(19.2  g/ha),  occurred  during  period  3. 
The  mean  biomass  of  area  2  over  the  six 
trapping  periods  was  102.5  g/acre  (41.5 
g/ha).  The  mean  was  approached  during 
periods  2,  4,  and  5.  Area  2  had  the 
greatest  and  the  lowest  biomass  of  all 
three  areas  during  the  study. 

Area  3.  The  greatest  biomass,  129.7 
g/acre  (52.5  g/ha),  for  area  3  occurred 
during  period  5  and  the  least,  72.2  g/acre 
(29.2  g/ha),  occurred  during  period  1. 
The  mean  biomass  of  area  3  over  the  six 
trapping  periods  was  109.9  g/acre  (44.5 
g/ha).  The  mean  was  approached  during 
period  2  and  kill-trai)  period  8. 

Discussion 

To  understand  an  A.  tridentata  com- 
munity a  knowledge  of  the  associated 
vertebrates  and  vegetation  must  be  ob- 
tained.   An  analysis  of  each  A.  tridentata 


study  area  is  thus  important  in  a  discus- 
sion relating  the  rodent  population  and 
biomass  of  the  individual  areas  to  each 
other  to  establish  the  picture  for  the  com- 
munity type. 

Peromyscus  majiiculatus  and  E.  mini- 
mus were  two  of  the  three  predominant 
rodent  species  in  each  area.  This  could 
be  exj)ected  for  P.  maniculatus  because  of 
its  geographic  range  over  most  North 
American  biomes  (Burt  and  Grossenhei- 
der,  1964;  King,  1968),  including  sage- 
brush regions  of  the  Great  Basin.  Eutam- 
ias  minimus  also  has  a  broad  geograph- 
ical range,  including  the  Great  Basin  and 
much  of  Canada.  Its  appearance  on  all 
study  areas  should  also  be  expected  since 
it  is  characteristic  of  sagebrush  commun- 
ities (Gordon,  1943;  Burt  and  Grossen- 
heider,  1964).  Out  of  43  vertebrate  spe- 
cies noted  during  the  study,  15  were  com- 
mon to  all  three  areas,  an  indication  that 
the  study  areas  were  similar.  Table  1, 
however,  indicates  that  notable  differ- 
ences existed  between  them.  These  dif- 
ferences resulted  from  the  distribution  of 
the  28  vertebrates  ^vhich  were  not  com- 
mon to  the  three  areas  (Nichols,  1972). 
Area  1  had  4,  area  2  none,  and  area  3 
12  unique  species  of  vertebrates  present. 
The  large  number  of  unique  species  for 
area  3  can  be  explained  by  the  presence 
of  a  more  diverse  habitat  than  in  the 
other  areas.  This  di^'ersity  was  likely  due 
to  more  favorable  climatic  and  edaphic 
factors  (Table  1).  Area  2  was  interme- 
diate in  soil,  slope,  temperature,  and  pre- 
cipitation (Table  1).  The  lack  of  verte- 
brate species  unique  to  area  2  also  indi- 
cates that  it  was  an  intermediate  area. 

Area  1  exhibited  the  least  vertebrate 
diversity,  area  2  was  transitional  but 
closer  to  area  1,  and  area  3  was  the  most 
diverse.  Most  of  the  rodent  biomass  for 
areas  1  and  2  came  from  E.  minimus, 
while  their  (ontribution  in  area  3  was 
small  (Table  3).  This  supports  the  ap- 
parent (loser  relationship  of  area  2  to 
area  1 .  It  should  also  be  kept  in  mind  that 
areas  1  and  3  are  the  farthest  apart  geo- 
grajihically  and  altitudinally,  with  area  1 
(elev.  5,100  ft.;  1,554  m)  near  the  valley 
floor,  area  3  (elev.  6,500  ft.;  1.981  m)  at 
the  base  of  the  valley-forming  mountains, 
and  area  2  (elev.  5,700  ft.;  1,737  m)  on 
the  benchland  between. them. 

The  ground  cover  of  each  area  was 
doniin;il(Hl  h\  A.  tridentata.  although  each 


June  1975 


NICHOLS   ET   AL.:    RODENT   POPULATIONS 


197 


area  had  a  different  predominant  under- 
story    species    (Nichols,    1972).     The    di- 
versity pattern  reflected  was  one  of  low 
plant  species  diversity  for  areas   1   and  2 
and  high  diversity  for  area  3.   Area  1  had 
14,  2  had  15,  and  3  had  30  plant  species 
present.     This   was   similar  to  the  verte- 
brate diversity  pattern  for  the  respective 
j  areas.    This  similarity  in  vertebrate  and 
j   plant    diversity    ])atterns    was    likely    due 
j   to  the  edaphic  and  climatic  factors  affec- 
j   ting  the  plants  which  in  turn  affected  the 
i  vertebrates.    The  lack  of  complete  simi- 
I   larity  between  vertebrate  and  plant  pat- 
terns, however,  may  have  been  due  to  the 
I   physiognomic  differences  of  the  cover  spe- 
cies present  in  the  areas  rather  than  the 
amount  of  diversity. 

It  is  suggested  that  the  amount,  distri- 
bution, and  physiognomy  of  dominant 
vegetative  cover  in  A.  tridentata  com- 
munities may  have  a  greater  influence 
than  the  diversity  of  cover  species  in  de- 
termining vertebrate  presence.  Turner 
(1950)  supports  this  idea,  especially  for 
Peromyscus  distribution,  in  his  study  of 
10  vegetative  types,  including  3  having 
A.  tridentata  as  the  dominant  or  co- 
dominant  plant.  Rosenzweig  and  Wina- 
kur  (1969)  have  hypothesized  from  stud- 
ies in  the  lower-Sonoran  desert  scrub 
vegetation  that  "the  spatial  variations  in 
density  of  some  species  [rodent]  are  re- 
sponses to  spatial  characteristics  of  their 
environment.  Important  among  these  en- 
vironmental characteristics  tend  to  be  mea- 
sures of  the  presence  and/or  absence  of 
vegetation  of  various  physiognomies." 

The  absolute  cover  of  A.  tridentata  in 
area  1  was  17.6  percent;  2  was  15.5  per- 
cent; and  3  was  23.2  percent.  Percent 
cover  composition  was  44.2  percent  (area 
1),  48.0  percent  (area  2),  and  38. .0  per- 
cent (area  3).  Area  2  had  the  least  ab- 
solute cover  of  A.  tridentata  with  the  most 
bushes  less  than  2  feet  (0.6  m)  high  but 
had  the  greatest  percent  cover  composi- 
tion of  A.  tridentata  for  the  three  areas. 
This  cover  pattern  may  have  been  why 
area  2  had  no  unique  vertebrates,  thus 
functioning  as  a  limiting  factor  to  diver- 
sity. Area  1  had  four  species  but  was  no 
more  diverse  than  area  2  in  terms  of 
plant  species,  indicating  that  in  this  case 
cover  type  had  a  greater  effect  than  did 
plant  diversity.  Area  3,  in  terms  of 
A.  tridentata  cover,  was  opposite  area  2. 

i 


Area  3  had  the  greatest  absolute  coverage 
of  A.  tridentata  with  most  bushes  greater 
than  2  feet  (0.6  m)  high  but  had  the 
least  A.  tridentata  cover  composition  for 
the  areas.  This  showed  the  opposite  ef- 
fect on  vertebrate  presence  than  the  pat- 
tern in  area  2.  Instead  of  having  no 
unique  vertebrate  species  as  in  area  2, 
there  were  12,  including  2  rodents.  The 
relationship  for  area  3,  however,  was  not 
as  pronounced  because  that  area  had 
nearly  twice  the  plant  species  diversity 
of  areas  1  and  2;  and  the  greater  plant 
species  diversity  may  be  responsible  for 
the  greater  vertebrate  diversity.  If  this  is 
true,  it  is  a  direct  reversal  of  the  results 
obtained  by  Rosenzweig  and  Winakur 
(1969)  in  the  lower-Sonoran  desert  scrub 
vegetation.  They  found  that  "the  varia- 
tion in  plant  s])ecies  diversity  failed  to  ex- 
plain the  \ariation  in  animal  species  di- 
versity and  that  some  of  the  most  faunal- 
ly  diverse  areas  had  the  fewest  species  of 
plants."  The  three  areas  reported  in  this 
study,  however,  are  in  the  Great  Basin 
cokr  desert,  which  varies  considerably 
from  the  Sonoran  hot  desert.  It  is  possible 
that  the  amount  and  distribution  of 
A.  tridentata  cover  is  only  important  in 
determining  vertebrate  distribution  in 
areas  with  low  total  absolute  cover  as  in 
areas  1  and  2  but  not  in  area  3. 

According  to  Pearson  (1965a,  1965b) 
and  Beatley  (1969),  primary  productivity 
of  A.  tridentata  communities  peaks  in  late 
si)ring  and  early  summer.  Peaks  in  rodent 
populations  in  A.  tridentata  communities 
occur  during  the  early  summer  to  fall 
period  (Turner,  1950;  Sullivan,  1961). 
Trojan  (1970)  has  shown  that  in  a  Polish 
grassland  the  rodent  biomass  increase 
during  the  summer  is  four  times  as  great 
as  during  the  winter  and  two  times  as 
great  as  during  the  spring.  Summer  and 
autumn  increases  accounted  for  89  per- 
cent of  the  annual  increase.  He  stated 
that  "winter  increases  are  of  almost  no 
importance  to  assessment  of  energy  flow 
(3.2  percent)."  His  results  may  be  appli- 
cable to  the  Great  Basin  sagebrush  zone 
because  its  increased  elevation  could  par- 
tially compensate  for  the  higher  latitude 
and  much  lower  elevation  of  Poland.  The 
Polish  study  was  done  in  a  grassland,  but 
the  areas  studied  in  Rush  Valley,  Utah, 
were  probably  grassland  before  the 
valley's  settlement  (Christensen  and 
Hutchinson,    1965).     Winter    production 


198 


GREAT  BASIN  NATURALIST 


Vol.  35,  No.  2 


may  be  important  in  some  sagebrush 
areas,  but  no  winter  data  were  taken  in 
this  study  due  to  inaccessibility. 

If  (1)  the  annual  peak  primary  pro- 
duction of  A.  tridentata  communities  was 
late  spring  and  early  summer,  (2)  ro- 
dent populations  in  A.  tridentata  com- 
munities peaked  in  the  early  summer  to 
fall,  and  (3)  there  was  little  increase 
in  rodent  biomass  during  the  previous 
winter  months,  then  the  population,  mean 
species  weights,  and  biomass  data  collec- 
ted during  this  study  should  illustrate  the 
dynamic  relationships  of  the  three  stud}' 
areas. 

Estimated  Populations 

The  total  population  estimates  of  all 
species  are  shown  in  Table  3  and  Figure 
1.  Area  1  had  a  definite  early  summer 
population  peak  followed  by  a  steady  de- 
cline and  a  leveling  off  in  the  fall,  but 
area  3  had  three  different  population 
peaks  during  the  summer  with  the  high- 
est population  occurring  in  late  summer 
(approx.  August  13).  Area  3  had  a  grad- 
ual increase  in  rodent  numbers,  peaking 
in  late  sunmier  (approx.  August  13)  and 
declining  by  September  3. 

Mean  Species  Weights 

Mean  species  weight  is  not  only  im- 
portant in  calculating  the  estimated  rodent 
biomass,  but  Walkowa  (1970)  pointed 
out  the  importance  of  species  weights  as 
an  exploitation  compensation  mechanism 
in  rodent  j)opulations.  He  found  that  re- 
production operates  as  a  compensating 
mechanism  only  if  exploitation  exceeds 
31  percent.  When  exploitation  was  0-30 
percent,  an  increase  in  exploitation  caused 
an  increase  in  the  production  of  biomass. 
In  this  study  there  appeared  to  be  no  un- 
usual predatory  or  disease  exploitation  of 
populations  above  30  percent;  thus  the 
species  were  likely  reacting  to  exploita- 
tion by  increasing  biomass  without  in- 
creasing reprodiutioii.  This  was  further 
evidenced  by  the  hu  k  of  high-poi)ulation 
densities.  The  greatest  estimated  rodent 
density  was  eight  rodents  per  acre  (3.24 
/ha)  in  area  2  during  its  summer  peak. 

In  addition  to  rodent  weights  varying 
with  population  exploitation,  they  also 
vary  with  the  animal's  dailv  activities. 
Tevis   (1955)   showed  that  the  £?ross  bodv 


Are 

a  2 

n 

|-| 

Trapping  Periods 


Fig.    1.     Number  of  rodents  estimated  in  all 
areas. 

weight  increase  in  chipmunks  going  from 
an  empty  stomach  to  a  full  stomach 
averaged  6-8  percent,  whereas  Evans 
(1949)  observed  that  voles  increased  20 
percent  in  body  weight  within  five  min- 
utes after  water  consumption.  It  is  evi- 
dent, therefore,  that  an  accurate  biomass 
estimation  requires  use  of  weights  from 
the  particular  time  and  population  being 
considered.  Mean  species  weights  used 
in  this  study  were  calculated  from  what 
was  believed  to  be  the  most  accurate 
weight  according  to  the  above  criteria, 
but  errors  may  have  occurred. 

Area  1.  The  mean  species  weight  of 
P.  manicuhttus  was  greatest  in  the  early 
spring  and  then  oscillated  between  lower 
weights  throughout  the  study.  The  high 
weight  during  period  1  reflected  the  pop- 
ulation structure  at  that  time.  There 
were  few  juvenile  and  subadult  animals 
compared  with  subsequent  periods.  After 
[)(>rio(l  1  there  were  increased  numbers 
of  juNcniles  emerging  from  the  nests, 
causing  a  sharp  decrease  in  mean  species 
weight  during  trap  period  2.  For  the  re- 
maiiidof  of  the  slud\-  fe])rocluction  caused 


June  1975 


NICHOLS   ET   AL.:    RODENT   POPULATIONS 


199 


oscillations  in  the  mean  species  weights 
between  trapping  periods.  These  oscil- 
lations were  expected  since  P.  maniculatus 
is  polyestrous  (Asdell,  1964).  The  E.  min- 
imus mean  species  weights  generally  ap- 
peared to  increase  throughout  the  study  to 
a  peak  near  the  end  of  the  summer  during 
trap  period  5.  This  increase  paralleled 
a  gradual  decrease  in  population  numbers 
throughout  the  summer.  The  E.  minimus 
reproductive  pattern,  one  litter  in  the 
spring  and  subsequent  growth  of  the 
young  during  the  summer  (Asdell,  1964), 
was  responsible  for  the  inverse  relation- 
ship. No  pattern  was  evident  for  R.  mega- 
lotis  because  of  insufficient  captures. 

Area  2.  The  mean  species  weight  pattern 
of  P.  maniculatus  in  area  2  oscillated  for 
the  same  reason  as  their  observed  oscilla- 
tions on  area  1.  but  the  greatest  mean 
species  weight  occurred  in  period  5.  There 
was  an  apparent  slow  period  in  repro- 
ductive activity  during  June  and  July 
compared  wdth  area  1.  This  undoubtedly 
allowed  the  summer  mean  species  weight 
of  the  population  to  peak  later  than  in 
area  1.  The  mean  species  weights  of 
E.  minimus  gradually  increased  dm-ing 
the  summer  because  of  their  reproductive 
pattern.  Too  few  P.  parvus  were  caught 
in  this  area  to  show  any  definite  patterns, 
but  the  same  slow  reproductive  activity 
as  noted  in  P.  maniculatus  on  this  area 
was  also  noted  for  P.  parvus. 

Area  3.  The  mean  species  weight  pat- 
tern of  P.  maniculatus  showed  reproduc- 
tion occurring  throughout  the  summer. 
The  P.  parvus  pattern  in  area  3  showed 
increased  population  numbers  associated 
with  decreased  mean  species  weights 
throughout  the  study.  This  was  exj^ected 
since  P.  parvus  is  polyestrous  (Asdell, 
1964).  Too  few  E.  minimus  were  caught 
on  this  area  to  show  any  definite  pattern. 


Biomass 

The  total  estimated  seasonal  biomass  in 
each  area  (Table  3  and  Fig.  2)  was  similar 
to  the  population  estimate  for  the  area 
(Fig.  1).  In  terms  of  biomass,  however, 
there  was  clearly  a  closer  relationship  be- 
tween areas  1  and  3  than  had  previously 
been  proposed.  Biomass  in  area  2  showed 
that  secondary  production  was  rather  un- 
stable compared  with  areas  1   and   3. 


Area   2 



2600 

Area 

A 

2400 

/  \ 
/    \ 

2200  . 

/      \ 

2000. 

/\ 

-r\ 

1800  . 

/ 

/       \ 

\    ^, 

1600- 

/ 

/ 

\/' 

^\ 

/ 

A~- 

---''    \- 

\                                \ 

UOO 

/ 

'   \ 

Ar 

1200- 

/ 

\ 

/  ^ 

/ 

\ 

/ 

1000- 

\    / 
\  / 

\ 
\ 

800 

V 

600 

- 

400 

200 

- 

Fig.  2.     Estimated  total  small  rodent  biomass 
by  trapping  periods. 

Area  1.  An  early  summer  increase  of 
50  percent  in  rodent  biomass  was  evi- 
denced in  area  1  during  a  13-day  interval 
between  trap  periods  1  and  2.  which  made 
trap  period  2  aj^pear  as  the  peak  period 
of  summer  rodent  biomass  production  for 
area  1  (Fig  2).  There  was  then  a  steady 
drop  in  biomass  over  a  42-day  period  fol- 
lowed by  a  leveling  off.  The  data  indi- 
cates, however,  that  the  number  of 
E.  minimus  and  R.  megalotis  ma}'  have 
been  overestimated  during  period  2. 
There  were  39  E.  minimus  estimated 
during  period  2,  but  only  29  were  handled. 
There  were  also  4  more  R.  megalotis  esti- 
mated than  were  actually  handled.  A 
high  number  of  unmarked  animals  being 
caught  at  the  end  of  a  trap  period  would 
cause  a  high  unmarked-to-marked  animal 
ratio  to  occur.  This  high  ratio  would  in 
turn  cause  an  overestimation  of  the  popu- 
lation. This  high  ratio  may  be  caused  by 
immigration  of  animals  into  the  area,  by 
new  animals  emerging  from  their  nests, 
or  by  various  other  factors.  In  this  in- 
stance the  possible  overestimation  may 
have  been  caused  by  new  juvenile  male 
chipmunks  becoming  available  to  the 
traps. 

If  the  actual  number  of  rodents  handled 


200 


GREAT  BASIN  NATURALIST 


Vol.  35,  No.  2 


during  period  2  is  used  in  the  biomass 
calculation,  then  the  summer  peak  on 
area  1  did  not  occur  during  trap  ])eriod  2 
but  during  trap  period  3.  The  possible  cor- 
rection is  shown  in  Figure  3.  The  total 
pattern  for  the  summer,  however,  is  still 
one  of  an  early  peak  in  rodent  biomass 
followed  by  a  steady  drop  and  then  a 
leveling  off  for  the  remainder  of  the  study. 
This  is  what  could  be  expected  for  a  com- 
munit}-  in  poor  soil  with  low  plant  spe- 
cies diversity,  as  area  1  was  earlier  shown 
to  be.  Secondary  production  peaked  early 
in  the  summer,  after  the  observed  peak 
primary  production,  thus  placing  greater 
demands  upon  the  primary  production  of 
the  community  to  support  the  increased 
secondary  production.  The  plant  com- 
munity, beyond  its  peak  production  and 
with  very  little  diversification,  likely 
could  not  produce  more  food  for  the  iii- 
creased  rodent  population,  so  the  rodent 
population  rapidly  declined  to  a  level  re- 
flecting the  probable  carrying  capacity 
of  the  community  and  then  remained 
there  through  the  end  of  the  summer.  Be- 
cause of  poor  soil  and  other  factors,  the 
carrying  capacity  was  low,  approximately 
55  g  of  rodent  biomass  per  acre    (22.27 


1400 
1200 
1000 


Trapping  Periods 


Fig.  3.  Estimated  total  small  rodent  bio- 
mass by  trapping  periods  using  possible  correc- 
tions. 


g/ha).  This  low  carrying  capacity  could 
liaAe  supported  no  more  than  one  E.  min- 
imus  and    one   P.    mcmicidatus   per   acre 

(.24/ha). 

Area  3.  The  rodent  biomass  of  area  3 
showed  an  entirely  different  pattern  than 
area  2  throughout  the  summer  (Fig.  2). 
Secondary  rodent  productivity  increased 
rapidly  after  spring  reproduction  and  was 
then  followed  by  a  short  period  of  reduc- 
tion before  rising  again.  This  reduction 
may  have  been  due  to  a  number  of  fac- 
tors but  was  most  likely  caused  by  cold 
temperatures.  The  P.  parvus  estimate  for 
the  period  of  biomass  drop  (trap  period  3) 
was  considerably  less  than  the  previous 
estimate.  This  rodent,  as  a  protective  be- 
havioral ada])tation,  regularly  goes  into 
a  state  of  torpor  when  it  encounters  cold 
(Bartholemew  and  Cade,  1957;  Beer, 
1961;  Morrison  and  Ryser,  1962;  Chew 
et  al.,  1965,  1967;  Tucker,  1962,  1963, 
1965a,  1965b,  1966).  During  the  second 
and  third  nights  of  trap  period  3  the 
coldest  summer  temperatures  were  re- 
corded for  area  3,  and  four  P.  parvus  that 
j)robably  would  ha^'e  been  caught  had  the 
nights  been  warmer  were  not  recap- 
tured. This  could  have  led  to  a  low  pop- 
ulation estimate  and  thus  a  low  biomass 
estimate  for  the  period.  If  this  were  the 
case,  then  the  biomass  of  area  3  would 
have  shown  an  increase  from  trap  period 
1  to  its  peak  in  trap  period  5.  The  pos- 
sible correction  is  shown  in  Figure  3.  At 
the  end  of  the  summer  peak  biomass  grad- 
ually declined.  Because  of  the  diversity 
of  the  }ilant  community  in  area  3,  fa- 
vorable climatic  conditions  and  primary 
production  were  jirobably  sufficient  to 
su])ply  food  for  the  gradual  summer  in- 
crease in   secondary  production. 

Area  2.  The  graphing  of  rodent  biomass 
for  area  2  showed  no  pattern  similar  to 
areas  1  or  3.  It  did,  however,  show  an 
earl}'  sininner  decline  in  standing  crop,  a 
rapid  reco^•ery  with  leveling  off  for  a  trap 
period,  a  midsummer  biomass  increase  to 
a  peak  higher  than  in  either  areas  1  or 
^,  and  a  late  summer  decline  in  standing 
crop  of  the  same  rate  as  that  which  oc- 
curred in  the  early  summer. 

Close  examination  of  the  data,  how- 
ov(n-.  for  tli(>  1\.  minituus  popidation 
showed  a  large  trap  mortality  during 
[x'riod   f).     This  c  auscd  T.iiu  oin's  index  to 


June  1975 


NICHOLS   ET   AL.:    RODENT   POPULATIONS 


201 


overestimate  the  population.  It  estimated 
20  more  E.  minimus  than  were  actually 
handled,  which  would  represent  an  over- 
estimation  of  1,060.5  g  in  hiomass.  Thus, 
at  traj)  jieriod  6  the  high  peak  shown  in 
Figure  2  would  probably  be  much  lower. 
The  possible  correction  is  shown  in  Fig- 
ure 3.  With  this  correction  the  biomass 
was  below  that  of  area  3  for  the  same 
period. 

Closer  examination  of  the  data  re- 
vealed another  possible  error.  In  trap 
period  9  there  were  29  different  E.  min- 
imus handled,  but  because  this  period  was 
a  kill-trap  period,  the  diurnal  chipmunks 
were  not  caught  during  the  one  live  trap 
night.  As  a  result,  none  were  marked, 
and  no  population  estimates  were  made  on 
day  2  of  the  kill-trap  period.  There  were 
17  different  chipmunks  handled  on  days 
1  and  2  of  this  period.  If  the  biomass  of 
these  rodents  were  added  to  the  total  bio- 
mass of  this  period,  it  would  raise  the 
total  b}'  879.7  g.  This  possible  correction 
is  also  shown  in  Figure  3. 

The  above  corrections  change  consider- 
ably the  estimated  summer  biomass  pat- 
tern of  area  2.  The  pattern  now  becomes 
one  of  an  early  summer  decline  in  bio- 
mass, a  recovery  period,  a  period  of  no 
increase  or  decrease  in  biomass,  and  a 
steady  rise  in  biomass  that  may  not  have 
peaked  before  the  study  ended.  In  terms 
of  stability  the  community  of  area  2  ap- 
peared to  be  less  stable  than  those  of 
areas  1  and  3.  The  corrected  pattern 
showed  that  in  terms  of  biomass  area  2 
was  more  similar  to  area  3  than  area  1 
as  had  been  previously  proposed. 

Conclusions 

It  appeared  that  when  the  characteris- 
tics of  ( 1 )  vertebrate  presence  and  dis- 
tribution, (2)  plant  presence  and  distri- 
bution, (3)  comparative  rodent  popula- 
tion numbers,  and  (4)  comparative  rodent 
biomass  of  the  areas  were  considered,  the 
three  areas  were  in  three  different  cli- 
matic or  edaphically  induced  successional 
stages  or  conditions.  Area  2  had  the  least 
diverse  vertebrate  presence.  This  may 
have  resulted  from  the  A.  tridcntata  cover 
being  composed  of  small  bushes  represent- 
ing the  greatest  percent  cover  composi- 
tion of  the  three  areas.  Area  2  was  the 
least  stable  in  terms  of  the  annual  cycle 


for  rodent  populations  and  biomass  in- 
crease, probably  a  result  of  the  poor  plant 
species  diversity.  Area  3  had  the  most  di- 
verse vertebrate  presence  and  was  the 
most  stable  in  terms  of  the  annual  cycle 
for  rodent  population  and  biomass  in- 
crease. This  may  have  been  due  to  the 
A.  tridentata  cover  pattern,  which  was  op- 
])Osite  that  found  in  area  2,  and/or  to  the 
increased  plant  species  diversity  found  in 
area  3.  Area  1  appeared  to  be  interme- 
diate, in  its  vertebrate  presence  and  A.  tri- 
dentata cover  patterns,  to  areas  2  and  3. 
Its  annual  cycle  in  terms  of  rodent  pop- 
ulation and  biomass  increase  showed  the 
earliest  peak  of  the  three  areas.  This  ma}' 
have  been  due  to  poorer  soil  and  climatic 
conditions  causing  an  early  peak  in  pri- 
mary productivity,  a  subsequent  early 
peak  in  rodent  productivity,  and  an  ex- 
tended period  through  the  rest  of  the  sum- 
mer when  conditions  did  not  favor  either 
primary  or  secondary  production. 

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Baker,  M.  F.  1969.  Unpublished  data  on  rodent 
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Be.\tley,  J.  C.  1969.  Biomass  of  desert  winter 
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Beer,  J.  R.  1961.  Hibernation  in  Perognathus 
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Beetle,  A.  A.  1960.  A  study  of  sagebioish,  the 
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Burt,  W.  H.,  and  R.  P.  Grossenheider.  1964. 
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Cain,  S.  A.,  and  G.  M.  de  Oliveire  Castro. 
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Chew,  R.  M.,  R.  G.  Lindbert,  and  P.  Hayden. 
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little  pocket  mouse,  Perognathus  longimem- 
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Christensen,  E.  M.  1967.  Bibliography  of 
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Christensen,  E.  M..  and  M.  A.  Hutchinson. 
1965.  Historical  obsers'ations  on  the  ecology 
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DiETTERT,  R.  A.  1938.  The  morphology  of 
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Evans,  R.  C.  1949.  A  population  study  of  house 
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GoLLEY,  F.  B.  1960.  Energy  dynamics  of  an  old 
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Gordon,  K.  1943.  The  natural  history  and  be^ 
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H.ARVEY,  J.  L.,  .AND  L.  Woodward.  1969.  Spe- 
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HiRONAKA,  M.  1963.  Plant-environment  rela- 
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Johnson,  W.  M.  1969.  Life  expectancy  of  a 
sagebrush  (Artemisia)  control  project  in  cen- 
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Kearl,  W.  G.  1965.  A  survey  of  big  sage- 
brush control  in  Wyoming:  1952-1964.  Wyo- 
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King.  J.  A.  (ed.)  1968.  Biology  of  Peromys- 
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The  Amer.  Soc.  of  Mammalogists. 

Morrison,  P.,  and  F.  A.  Ryser.  1962.  Hypo- 
thermic behavior  in  the  hispid  pocket  mouse. 
J.  Mammal.  43:529-533. 

Nicholes,  D.  W.  1972.  Small  rodent  popula- 
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thesis.    Brigham   Young   Univ..    Provo.    Utah. 

Odum,  E.  p.  1959.  Fundamentals  of  ecologv. 
W.    B.   Saunders   Co.,   Philadelphia. 

Passey.  H.  B.,  and  V.  K.  Hugie.  1962.  Sage- 
brush on  relict  ranges  in  the  Snake  River 
Plains  and  Northern  Great  Basin.  J.  Range 
Manage.   15:273-278. 

Pearson,  L.  C.  1965a.  Primary  productivitv  in 
a  northern  desert  area.  OIKOS   15:211-228. 

.      1965b.     Primarj'    production    in    grazed 

and  ungrazed  desert  communities  of  eastern 
Idaho.  Ecology  46:278-285. 

ROSENZWEIG,      M.      L.,      AND      J.      WiNAKUR.        1969. 

Population    ecology    of    desert    rodent    com- 


munities: habitats  and  environmental  com- 
plexity. Ecology  50:558-572. 

Shelford,  V.  E.  1963.  The  ecology  of  North 
America.   Univ.   Illinois  Press,  Urbana. 

Sullivan,  John  O.  1961.  Population  structure 
of  Peromyscus  maniculatus  in  two  areas  in 
Green  Canyon,  Cache  County,  Utah.  M.S. 
thesis.    Utah    St.    Univ..   Logan. 

Tevis,  L.,  Jr.  1955.  Observations  on  chipmunks 
and  mantled  squirrels  in  northeastern  Cali- 
fornia. Amer.  Midi.  Nat.  53:71-78. 

Trojan,  P.  1970.  Energy  flow  through  a  popu- 
lation of  Microtus  arvalis  in  an  agrocenosis 
during  a  period  of  mass  occurrence.  Pages 
267-279  in  Energy  flow  through  small  mam- 
mal populations.  Warszawa. 

Tucker,  V.  A.  1962.  Diurnal  torpidity  in  the 
California  pocket  mouse.  Science  136:380-381. 

.      1963.     The     energetics     of     the     torpor 

cycle  in  the  California  pocket  mouse,  Pe- 
rognathus  californicus.  Ph.D.  dissertation. 
Univ.   California   Los  Angeles.    (Diss.   Abstr.) 

.     1965a.     Oxygen     consumption,     thermal 

conductance,  and  torpor  in  the  California 
pocket  mouse,  Perognathus  californicus.  J. 
Ce.    Comp.    Physiol.    65:393-403. 

.  1965b.  The  relation  between  the  tor- 
por cycle  and  heat  exchange  in  the  Cali- 
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cus.    J.     Cell.     Comp      Physiol.     65:405-414. 

.      1966.     Diurnal  torpor  and  its  relation  to 

food  consumption  and  weight  changes  in  the 
California  pocket  mouse,  Perognathus  cali- 
fornicus. Ecology  47:245-252. 

Turner,  G.  C,  Jr.  1950.  Peromyscus  popula- 
tions as  related  to  seasons  and  vegetative 
types  at  the  hardware  ranch.  Cache  County, 
Utah.    M.S.    thesis.    Utah    St.    Univ.,   Logan. 

Vest,  E.  D.  1962.  The  plant  communities  and 
associated  fauna  of  Dug^vay  Valley  in  west- 
ern Utah.  Ph.D.  dissertation.  Univ.  of  Utah, 
Salt  Lake  City. 

Walkowa,  W.  1970.  Operation  of  compensation 
mechanisms  in  exploited  populations  of  white 
mice.  Pages  247-253  in  Energy  flow  through 
small    mammal    populations.    Warszawa. 

Woodbury,  L.  1955.  An  ecological  and  distri- 
butional study  of  small  mammals  of  Cedar 
Valley,  Utah.  M.S.  thesis.  Brigham  Young 
Univ.,   Provo,   LTtah. 


COMPUTERIZED  REDUCTION  OF  METEOROLOGIC 

MEASUREMENTS  FROM  IRRKiATED  AND  NONIRRIGATED 

PLOTS  IN  CENTRAL  UTAH' 

Ferron  L.  Andersen"  and  Paul  R.  Roper'' 

Abstract. —  Two  Fortran  iv  computer  programs  were  developed  to  facilitate  reduction  of  mete- 
orologic  data  from  iri-igated  and  nonirrigated  plots  at  Provo,  Utah.  The  first  program  compiles 
and  tabulates  daily,  monthly,  and  jearly  summaries  of  precipitation  as  rain  and/or  snow,  snowfall, 
total  snow  cover,  soil  moisture,  dew,  lelative  humidity,  potential  evaporation,  cloud  cover,  and 
wind.  Temperature  values  are  tabulated  for  measurements  taken  in  a  standard  weather  shelter.  5 
cm    beneath    soil    surface    under    grass   cover,    at    soil  surface  under  grass  cover,  and  on  bare  ground. 

The  second  program  enables  complete  computerized  (Calcomp)  construction,  labeling,  and  graph- 
ing  of    10    different   meteorologic    measurements    and  3  calculated  comparisons  of  temperature  means. 

Advantages  of  the  first  computer  program  relate  generally  to  that  obviously  noticeable  with 
any  computerized  tabulation.  Those  of  the  second  j)rogram  relate  more  specifically  to  the  greatly 
reduced  cost  of  computerized  graphs  compared  with  those  produced  manually,  as  well  as  to  the 
marked  reduction  of  eirors  compared  with  the  number  frequently  associated  with  the  usual  tedious 
and  laborious  plotting  of  voluminous  weather  data. 

Meteorologic  data  collected  for  the  year  demonstrated  the  beneficial  effect  of  irrigation  in  the 
creation    of    microenvironmenfs    for    living    organisms. 


Introduction 

A  comparison  of  meteorologic  measure- 
ments fromi  irrigated  and  nonirrigated 
plots  in  Provo,  Utah,  for  1970  through 
1972  was  published  recently  by  Andersen, 
Wright,  and  Fox  (1974).  Included  in  their 
report  was  a  detailed  description  of  the 
study  area,  instrumentation  employed, 
method  of  handling  meteorologic  clata, 
and  a  series  of  manually  rej^roduced 
graphs  depicting  the  results  for  those  three 
years.  The  current  report  is  designed  as 
a  companion  article  to  the  one  referred 
to  above.  It  extends  the  meteorologic 
measurements  through  1973  for  an  ad- 
jacently located  plot  watered  via  sprink- 
ling as  opposed  to  flood  irrigation  used 
in  the  previous  project.  Furthermore,  it 
emphasizes  those  changes  that  have  been 
incorporated  to  enable  the  graphing  of  10 
different  meteorologic  measurements  and 
3  comparisons  of  calculated  means  by  a 
Calcomp  plotter. 

The  importance  of  irrigation  in  creating 
optimum  microenvironments  for  biolog- 
ical organisms  has  been  substantiated  for 
such  invertebrates  as  mosquitoes  (Rainy 
and  Hess,  1967;  Reeves  and  Hammon, 
1962),  snails  (World  Health  Organization, 
1950),  and  nematode  larvae  (Furman, 
1944;  Honess  and  Bergstrom,  1966;  Fox, 
Andersen,  and  Hoopes,  1970;  and  Wright 
and  Andersen,  1972).  The  importance 
was  elaborated  upon  further  by  Andersen 

^This  project  was  supported  in  part  Ijy  Public  ticallli  Seri 
^Department  of  Zoology,  Brigham  Yoiuig  l^niversity.  I'n 
^Eyring   Researcli    Institute,    Provo,    Utali. 


et  al.    (1974)    and,  thus,  will  not  be  dis- 
cussed here. 

Description  of  Experimental  Plot 

The  experimental  plot  measured  10  x 
30  m  and  was  located  at  the  Brigham 
Young  LTniversity  Animal  Science  Farm, 
Provo,  Utah,  adjacent  to  the  plot  used  in 
the  study  by  Andersen  et  al.  (1974).  The 
coordinates  of  the  station  site,  elevation, 
description  of  vegetative  cover,  soil  type 
and  drainage,  instrumentation,  and  meth- 
od of  collecting  data  were  the  same  as  re- 
ported earlier.  The  plot  was  divided  into 
an  irrigated  and  a  nonirrigated  section 
separated  by  an  elevated  dike.  Addition- 
al diking,  a])roximately  8  in  (20  cm) 
high,  was  also  placed  around  the  upper 
border  and  sides  of  the  plot  to  prevent  any 
flood  irrigation  waters  from  entering  the 
experimental  area.  Beginning  in  May 
the  irrigated  portion  was  sprinkled  each 
Monday  with  a  total  of  1  in  (2.5  cm)  of 
water  as  measured  in  the  rain  gauge.  This 
was  applied  at  an  approximate  rate  of  0.5 
in/hr.  In  June  the  amount  was  in- 
creased to  1.5  in  (3.25  cm)  and  in  July 
to  2  in  (5.0  cm).  The  amounts  were  re- 
versed for  August,  September,  and  Oc- 
tober, after  which  time  sprinkling  ceased. 
This  regimen  for  1973  added  a  total  of  39 
in  (97.5  cm)  of  water  to  the  irrigated  plot 
over  the  six  month  period. 


203 


204 


GREAT  BASIN  NATURALIST 


Vol.  .55,  No.  2 


Descriptions    of    Programs 

Two  FORTRAN  IV  programs,  one  for 
data  tabulation  and  a  second  for  Calcomp 
graphing,  were  developed  for  this  project. 
Figure  1  outlines  the  collation  and  hand- 
ling of  meteorological  data  and  Figures  2 
and  3  illustrate  flow  charts  for  the  data 
tabulation  and  Calcomp  graphing  pro- 
grams respectively.  All  meteorologic  data 
for  the  year  were  entered  on  specially  de- 
signed worksheets  (Figs.  4  and  6),  key- 
punched on  80-column  IBM  cards,  and 
handled  as  diagrammed.  All  mensural 
data  not  already  in  the  metric  system 
were   so   transposed    by   appropriate   con- 


ifii 


WEATHERSTATION 


version  formulae.  Also,  the  printing  of 
all  negative  or  zero  values  was  suppressed 
whenever  such  data  were  not  meaningful. 

The  first  program  compiles  and  tab- 
idates  daily,  monthly,  and  yearly  sum- 
maries of  all  meteorologic  data  collected. 
Tables  1,  2,  and  3  are  sample  printouts  of 
one  month's  data  (May  1973),  and  Tables 
4  and  5  are  the  two-page  annual  sum- 
mary sheets. 

The  second  program  developed  for  the 
project    enables     complete    computerized 


PRINT  MONTHLY 
HEADING 

. 

>' 

READ  DATA 
FOR  ONE  DAY 

, 

CALCULATIONS 

a 

CONVERSIONS 

^/^C 

0F\. 

NO 

VONTHLY  SUMS, 

AVERAGES  6 

EXTREMES 


PRNT  MONTHLY 
D(\TA(3PAGES) 


STORE  MONTHLY 

TOTALS 

FOR  YEARLY 

SUMMARY 


Pig.    1     Diiigrarn    showing    how    metoorologir 
data  were  collated  and  handled. 


Fig.  2.     Flow    chart    for    the    data    tabulation 
program. 


June  1975 


ANDERSEN,  ROPER:  METEOROLOGIC  DATA 


205 


plotting  of  10  different  meteorolocric 
measurements  and  3  calculated  mean  tem- 
perature comparisons.  Each  deck  of  ]iro- 
gram  data  for  any  one  year  must  be  ])re- 
ceded  by  a  control  card  identifying  the 
year  and  number  of  days  in  that  year. 
Also  included  on  the  control  card  for  the 
Calcomp  plotting  program  is  a  list  of 
identifying  nmubers  which  permit  selec- 
tion of  desired  gra])hs  for  that  year.  These 
graphs  are  then  comjileted  sequentially  as 
selected. 

Figures  6  through  18  depict  comj5uter- 
ized  reproduction  of  10  different  metero- 
logic    measurements    taken    during    1973 


and  3  calculated  comparisons  of  means 
as  indicated  on  each  individual  legend. 
Table  6  gives  information  for  the  1973 
data  regarding  the  a]:)proximate  run  times 
and  current  costs  for  the  IBM  360/65  in- 
stallation  at  Brigham  Young  University. 

The  complete  printout  of  all  daily, 
monthly,  and  yearly  data  is  available 
uj)on  request  for  the  cost  of  reproduction. 
Both  com])uter  j)rograms  are  printed  here- 
in (Appendix)  for  researchers  who  might 
find  sections  or  subroutines  applicable  to 
their  specific  needs  in  meteorolog\-  or 
biology. 

Regarfling  specific  handling  of  meteoro- 


(        BEGIN         j 


READ  CONTROL 

CARDS 

POSITION  TO 

YEAR'S  DATA 


READ  YEARLY 
DATA 


BRANCH  TO 

DESIRED  GRAPH 

NUMBER 


INCREMENT  TO 

NEXT  GRAPH 

NUMBER 


>rl3 


SCALE 
DATA 


DRAW  AXES  8 

LABEL 

MOfvTTHS 


DRAW  AXES  8 

LABEL 

MONTHS 


LABEL  LEGENDS 
a  UNITS 


LABEL  LEGENDS 
8  UNITS 


PLOT  DATA 


I   i    i    i   i 


PLOT  DATA 


Fig.  3.     Flow     chart     for     the     Calcomp     plotting    program. 


206 


GREAT  BASIN  NATURALIST 


Vol.  35,  No.  2 


Table    1.     Sample  monthly  summary  of  measurements    on    precipitation,    soil    moisture,    relative 
umiditv.    potential    evapoiation,    cloud    cover,    and  wind — May    1071 


PRECIPITATION  (f 


SOIL  MOISTURE 


0.3  0.3 


PERCENT    DEfc 


TOTALS 

MEANS 

EXTREMES 
HIGH 


L28.6      23 


TE    Cf    IRRIGATION    (MAY    THROUGH    OCTOBER! 


Table  2.     Sample   monthly    summary    of    temperatures  measured  from  a  standard  weather  shelter 
and   at   5    cm   beneath   soil   surface   under   grass   cover — May   1973. 


•EATHEP     SHfl 


THERMOMETEf 


FEMPEBATUReS 


HYGPOTHERMOGRAPH 


L     SURFACE 

NON-IRRIGATfD 
MAX.  MIN.  M 


12.0 
17. T 


11.  5 

ll.O 

35.0 

12.0 

26.0 

13.3 

27.  ) 

15.1 

MEANS 
EXTREMES 


EMPERATURES  IN  DEGREES  CELSIUS 


June  1975 


ANDERSEN,  ROPER:  METEOROLOGIC  DATA 


207 


Table  3.     Sample  monthly  summary  of  temperatures  measured  from  a  standard  weather  shelter, 
at   soil   surface  under  grass  cover,   and   on  bare  soil  surface — May  1973. 


SOIL    SURF4CE    UNDER    GRASS    COVER 


8«RE    SOIL    SURFS 


THERMOMETER 


IRRIGATED 


NON-IRRIGATED 


X.           HI 

N. 

MEAN 

.3         -3 

5.0 

7 

14 

14 

12 

12 

12 

10 

K 

13 

15 

17 

15 

15 

17 

17 

'o         10 

20 

.8         K 

.1          10 

18 

U 

H 

.7         IC 

18 

8 

6 

13 

15 

15 

17 

5.8  14.3 


25.0 
25.0 
32.0 


.0 
.0 

11. 

13. 

.0 

16. 

.0 

18. 

.0 

15. 

.0 

16. 

.0 

17. 

.0 

.0 

I7I 

17. 

!o 

21. 

.0 

24. 

.0 

24. 

.0 

26. 

.0 

25. 

.0 

27. 

.0 

27. 

28. 

!o 

31. 

.0 

30. 

.0 

24. 

.0 

26. 

.0 

2<>. 

.0 

30. 

.0 

It. 

.0 

21. 

.0 

22. 

.0 

21. 

.0 

25. 

.0 

25. 

.0 

26. 

32.0 

9.0 

20.5 

17.0 

10.0 

13.5 

24.0 

9.0 

16.5 

24.0 

11. 0 

17.5 

26.0 

12.0 

19.0 

22.0 

8.0 

15.0 

28.0 

10. 0 

19.0 

30.0 

10.  0 

20.0 

30.0 

8.0 

19.0 

35.0 

9.0 

22.0 

35.0 

9.0 

22.0 

36.0 

8.0 

22.0 

35-0 

8.0 

21.5 

36.0 

9.0 

22.5 

39.0 

10.0 

24.5 

41.0 

16.0 

28.5 

43.0 

12.0 

27.5 

37.0 

13.0 

25.0 

38.0 

9.0 

23.5 

43.0 

9.0 

26.0 

40.0 

13.0 

26.5 

45.0 

14.0 

29.5 

IT.O 

7.0 

12.0 

27.0 

7.0 

17.0 

31.0 

8.0 

19.5 

36.0 

10.0 

23.0 

39.0 

10.0 

24.5 

39.0 

12.0 

25.5 

38.0 

14.0 

26.0 

.0 

0 

0 

14.0 
18.3 

.0 

5 

0 

24.0 

.0 

6 

0 

25.5 

.0 

5 

23.0 

.0 

6 

0 

24.0 

.0 

6 

0 

26.5 

.0 

2 

0 

20.0 

.0 

0 

24.5 

.0 

3 

0 

27.5 

.0 

8 

0 

23.5 

.0 

8 

0 

29.0 

.0 

0 

30.5 

.0 

8 

0 

31.0 

.0 

7 

0 

31.0 

.0 

9 

0 

32.5 

.0 

10 

0 

34.5 

.0 

15 

0 

37.5 

.0 

12 

0 

36.5 

.0 

13 

0 

32.5 

.0 

9 

0 

20.5 

.0 

9 

0 

27.0 

.0 

14 

9 

32.0 

.0 

9 

0 

30.5 

.0 

7 

0 

12.0 

.0 

6 

0 

18.5 

.0 

6 

0 

21.5 

.0 

9 

0 

21.5 

.0 

9 

0 

23.5 

.0 

11 

0 

27.0 

.0 

14 

0 

30.0 

25.5 
27.5 
24.0 


ALL    TEMPERATURES    IN    DEGREES    CELSIUS 


Table  4.     Yearly  summary  of  measurements  on    precipitation,    soil    moisture,    relative    humidity, 
potential  evaporation,  cloud  cover,  and  wind — 1973. 


MONTH 

TOTAL 
PREC 

PRECI 

PAIN 

pREr 

PITATTON 

SNCW 
PREC 

(MM) 

SNOW 
FALL 

SNOW 
COVER 

SOIL    MOISTURE 
(PERCENT! 

IRRG       NON-IRRG 

RELATI 

HUMIDI 

JANUARY 

TOTAL 
MEAN 

29.7 
1.0 

0.0 
0.0 

2';. 7 

1.0 

297.2 

3302.0 
106.5 

NO 

NO 

98 

FEBRUARY 

TOTAL 
MEAN 

33.8 
1.2 

31.7 
l.l 

2.0 
Q.l 

2T.3 
0.7 

363.2 
13.0 

NO 

NO 

98 

MARCH 

TOTAL 
MEAN 

86.6 
2.8 

58.7 
1.9 

27.9 

0.9 

279.4 

2S7.0 
9.3 

UD 

NO 

98 

APRIL 

TOTAL 
KEAN 

1.5 

33. U 
1.1 

I  1.4 

0.4 

114.3 
3.8 

114.3 

J.  8 

NO 

NO 

98 

IS    OF  POT 


AUGUST 

SEPTEMBFP 

CCTOBER 

NOVEMBER 

CECEMPER 


OTAL 
EAN 

34.0 
1.  I 

34.0 
1.  L 

0.0 
0.0 

OTAL 

18.3 
0.6 

18.3 
0.6 

0.0 
0.0 

OTAL 
EAN 

19.8 
0.6 

19.8 
0.6 

0.0 
0.0 

OTAL 

51.8 

51.8 

T.O 

tOTAL 

"FAN 

lOTAL 


13.2 

0.4 

17.0 
0.0 


0.0 
0.0 

0.0 

0.0 
0.0 

9. 
8. 

0.0 
0.0 

0.0 

0.0 

<,'. 

0.0 

0.0 

0.0 
0.0 

11. 

0.0 
0.0 

oio 

12. 

0.0 
0.0 

0.0 
0.0 

25.7 

10. 

CLOUD 

COVER         WINO 

(0-101        (KM) 

1183 

6  38 

1144 
5  40 

2012 

fc  64 

1901 
4  63 


<EARL 


317.0  179.6 

0.9  0.5 


b568.4 
17.9 


208 


GREAT  BASIN  NATURALIST 


Vol.  35,  No.  2 


MONTH. 


YEAR 


METEOROLOGIC  DATA 


PARASITOLOGY 


Table  5 
beneath  soil 
1973. 


Yearly  summary  of  temperatures  measured    i 
surface  under  grass  cover,   at  soil  surface  undt 


HELTEP  TEMPEPATURES 

HYGPCTHEBMOGRAPh 


JANUARV 

FEBRUARY 

PARCH 

APRIL 

JUNE 

JULY 

AUGUST 

SEPTEHBER 

OCTOBER 

NOVEMBER 

DECEMBER 

a    standan 
errass  covei 


weather    shelter,    at    5    cm 
and  on  bai-e  soil   surface — 


X.             MI 

N. 

MEA 

.? 

.8            -■ 

.^ 

.0            -  1 

.?. 

.9 

.5            11 

.8 

16. 

.1              19 

.2 
.1 

23. 

.5             U 

22. 

.  1             12 

.A 

15. 

.q 

n. 

.■) 

.7            -0 

.7 

SURFACE    UNDER    GRASS    COVER 


JANUARY 

-1.9 

-2.5 

_2. 

FEBRUARY 

1.0 

-2.* 

-C. 

MARCH 

-0.3 

APRIL 

12.^ 

2.6 

KAY 

33.7 

10.  <. 

JUNE 

41.2 

JULY 

35.7 

19.8 

AUGUST 

32.6 

SEPTEMBER 

2'.. 8 

13. <. 

OCTOBER 

13.0 

9.3 

NOVEMBER 

4.  3 

CECEMBER 

1.3 

NON- I  SB  I  GAT  ED 


.0 

Ij 

.0 

4 

8 

21 

13 

28. 

16 

33. 

15 

30 

20. 

14. 

-1 

0. 

CRANO    MEAN 


ALL    TEfFERATURES    IN    CEGRfES    CELSIUS 

Table   6.      1975   data   for   time   and   costs   for  weather    data    tabulation    and    Calcomp    plotting    i)ro- 
grams    using    the    IBM   360/65    installation   at   Brigham  Young  University. 


Program 

Tu 

rn  arou 
time 

nd 

Compiler 
costs 

Run 
costs 

Plotter 
costs. 

Total 

Weather  data 

tabulation 
Calcomp  graphics 

0.5  hr 
2.0  In- 

$6.75 
$7.00 

$  7.50 
$13.00 

$5.00 

$14.25 
$25.00 

$39.25 

June  1975 


ANDERSEN,  ROPER:  METEOROLOGIC  DATA 


209 


PRECIPITATION  and  SOIL  MOISTURE 

PRDVD,  UTOH,  1973 


I A    LlfaUhLJ, 


jUU 


15  1  :5 


H  /I  t  I  I -Mil  li 


15        1         !?>        1        15        a 


FEpruflffT         npwcn     .       prr^iL  rwr  juwe     ,       jult  wjcust        sEnEweEw,     octopew  ,  wovEr^pew      pecEWBEr 


SNOW   COVER 

PROVO.   LiTBH.    1973 


250- 
225-] 

cco-j 


© 


^..  infi 


6    ^ 


!H 


lA 


M 


IN3 


JHNUflflY       FEBRURRY  HflRCH      ^     flFRlL    . ^»Y  jUnF      .         JULY       ,       PJJGOST      ^    SFPTEMBCR      DCTQBEfl   .    NOVEMBER   .    DFCgUBFR 


I  RELATIVE  HUMIDITY  IN  WEATHER  SHELTER 


JHNUfWY        FEBRUflRY    ,         MARCH  fiPRlL 


JUNE      .         JULY       ,       BUGUST      ,    SEPTEMBER.      JHOBER   .    NOVEMBER   ,    DECEMBER 


210 


GREAT  BASIN  NATURALIST 


Vol.  35,  No.  2 


jUNf      ,         Mir       ,       AUGUST      .    SLPieMPeR.      OCTOBgB   .    NPYEM6FH  .    DCCEMeew 


ffwusT    ,   scrTEfipEn, ocToeE^  ,   wovEnoEff  ,   oEcempeh 


TOTAL  WIND   1   METER  ffiOVE  GROUND 

PSDVO.    U7PH,    1913 


E  160. 

i 


JBNUWY     .rCBBUOSy   .        MflBCH      .         fyWIL    .         HftY         .         jOnE      .         JULY       .      RUGUST      ,    SCPIfrBER.      OCTDBER   .    NOVFWER   .    oecCMBfK   . 


June  1975 


ANDERSEN,   ROPER:    METEORO LOGIC  DATA 


211 


TEMPERATURE  IN  WEATHER  SHELTER 

PRDVD.    UTftH.    1973 


15         1  15         1  15  1  15         1  15         1  15         1  IS         1  15         )  15         1  15         1  IS         )  IS 

JHNUPRV     .FfBBI^RY   .        MftRCH      ,         WRU    .         mf         .         JUNE      .         JUIV       .      flUCDST      .   5fP1FM6fR.      OCTQefB  .    NO»fWBfH  .    DfCCM8fR 


COMPARISON  OF  MAXIMUM  AND  MINIMUM  TEMPERflTLIR&S'l  , 

5  CM  OEfP   In  soil  L'NDfR   10  O  CRASS  COVER  ,  fyj  ^ ',   ii      ^_ 


ON    IRRIGfiTED   ftNO   NON-lflRICfllfO   PLOTS 
PROVO%   UTflHi    1973 


MAX  NON-IRR 

MIN  NON-IRR 

HftX   IRRIGWED 

HIN  IRRIGATED 


JflNUtWY       EEBflUflRY    ^       MfWCri       ^     ftPRIL     . WiY 


flOGbST      .    SEPTfMeER.      OCTOBER  .    NDVEMeER   .    DECEneER 


COMPRRISON  OF  DFllLY  MEAN  TEMPERATURES 

S   Cfl   DEEP    IN   SOIL    UNPEB    id   CM   DPflSS   UIVFR 
ON    IRHIOPTFD  AND   NDN-IBR lOflTED   TLOTS 


rrOVO.    UTRH.    1973 


212 


GREAT  BASIN  NATURALIST 


Vol.  35,  No.  2 


COMPARISON  OF  MAXIMUM  AND  MINIMUM  TEMPfRRTUP£:,S!:   r 

_..^: 

WW  NON-IRR 

50- 

ON   IRBJCflTfO  AND  N0N-IRP1&H7CD  PLOTS                                                       .  . 
PflOVO,    UTAH,    1973                                                                   .             ^    :,■        jj^ 

ii^lji': 

MIN  IRRIGOTED 

MO- 
"  30- 

M 

Wi 

%ij 

1  /: 

h 

L,¥fts:i 

^^n^ 

t\'' 

10- 

ri>^Y^  V 

■^\W%v 

temll  A 

0 
ID 

Am^i  a' JMi./W"^  ■ 

f 

•■     ■.  ^1  i  L'  v  )?(/f\/|;;;^!5t[(c^ 

^^^^^^^^^^^^^^(V^jV^I  •» 

'■■"■■■"'■■ 

I's       i       is      ')       ')5 

I          15         1          15 

1        is 

1        is       'i 

is        i         I'S        1 

15         i          15         i          I'S 

QPMUflBT     . FtBBUflRT 


RUCUST     .    SFFTFfiBFR.      0CT06CB   .    NOVCMSfH   .    DECfUBCH 


COMPfiRISDN  OF  OfllLY  MCflN  TEMPERATURES 
m  sou  suRFHCf  UNDfn  lo  i>i  Offlss 

ON  IRfilGRTfO  fiND  NON-lRRIGfllfO  PtmS 
PROVO,   UTBH,    1973 


JfiNUPRT        f-fBRUW 


SFPlffWfR       OClOefR       NDVEwerR   .    OCCfllBCR 


COMPARISON  OF  MAXIMUM   AND   MINIMUM  TEMPERATURES 
OT    SOIL   SURFRCC  ON  BflRf   GROUND   ON 

IRRlGflTCD  AND  NON- IRRIGATED  PLOTS  j 

PROVO.   UTAH.    1973  «    ■"/ 


HflX  NON-]Rfl 
WIN  NON-IRfl 
NftX  IRfllGRTEO 
(•IN  IRRIGfllfO 


I'S        i       Ts 


flocusT    .  sfPTfnefR.     ocToefR  .  NOVC^eCR  .  occfnBffl  . 


.hiiie  1975 


ANDERSEN,   ROPER:    METEOROLOGIC   DATA 


213 


logic  iiiforniatioii  from  the  i)asture  plots 
as  well  as  certain  calculations  and  con- 
versions b}'  the  computer,  the  following 
clarifications  may  be  warranted.  All 
measurements  of  precipitation  as  rain  are 
measured  and  entered  in  fractions  of 
inches  and  then  converted  before  tabu- 
lation into  millimeter  units.  New  snow 
and  total  snow  cover  are  similarly  han- 
dled. Precipitation  as  snow  is  calculated 
as  one-tenth  that  of  total  snowfall  for  any 
one  day.  Soil  moisture  samples  were  de- 
termined as  described  previously  by  An- 
dersen et  al.  (1974),  with  the  exception 
that  two  samples  were  taken  each  week 
from  the  irrigated  plot — one  just  prior 
to  sprinkling  of  the  plot  and  the  second 
24  hr  later.  Only  one  weekly  sample 
was  taken  from  the  nonirrigated  plot.  The 
weight  in  grams  of  each  sample  was  en- 
tered as  an  original  wet  weight  and  then 
as  a  dry  weight  determined  after  24  hr 
storage  at  105  C. 

Programmed  formulae  calculated  the 
percent  soil  moisture  by  dividing  the  dif- 
ference in  the  wet  and  dr}-  weight  for  each 
sample  by  the  dry  weight  value.  This  per- 
cent was  then  printed  on  the  first  print- 
out page  for  each  of  the  six  months  that 
irrigation  was  used.  The  presence  or  ab- 
sence of  dew  on  each  plot  was  noted  for 
each  morning  during  those  six  months 
and  recorded  as  a  "  +  "  when  present. 
Irrigation  by  sprinkling  was  performed 
each  Monday  during  the  six-month  period 
and  also  indicated  with  a  "  +  "  on  the  ap- 
propriate dates. 

Relative  humidity  (RH)  maxima  and 
minima,  as  well  as  the  total  number  of 
hours  each  day  at  which  98-100  percent 
RH   occurred,   were   entered   and   printed 


out  directly.  The  e\aj)orating  pan  on  the 
instnnnent  that  measured  potential  evap- 
oration was  filled  each  morning,  and  the 
daily  water  loss  noted  for  the  following 
24  hr  was  entered  in  mm  and  also  printed 
out  directly.  Since  evaporation  could  not 
be  read  in  freezing  weather,  the  measure- 
ment was  taken  only  during  May  through 
October,  those  same  months  during  which 
the  (^ne  plot  was  irrigated.  A  difficulty 
ensued  whenever  rain  occurred  because 
the  e\aporating  pan  held  only  20  nun  of 
water  and  refilled  partially  or  completely 
on  any  day  during  which  rain  fell.  Thus, 
any  potential  evaporation  which  may  ha^e 
occurred  on  such  days  was  invariably  ne- 
gated to  some  degree  by  the  rain  that  col- 
lected in  the  evaporating  pan.  Neverthe- 
less, the  recording  evaporimeter  used  in 
the  study  gave  a  much  more  accurate  re- 
flection of  the  potential  evaporation  with 
its  shallow  pan  (20  mm)  than  do  the 
large  evaporation  tanks  used  by  the  U.S. 
Weather  Bureau.  In  those  tanks  the 
water  level  is  frequently  4-6  in  (10-15 
cm)  below  the  ujiper  rim  of  the  pan  and 
hence  protected  markedly  from  the  evapo- 
rating effect  of  wind  currents. 

Any  evaluation  of  daily  cloud  cover  was 
determined  visually  each  morning  and 
recorded  in  tenths.  The  evaluation  indi- 
cated the  approximate  portion  of  the  sky 
that  was  covered  sufficiently  with  clouds 
to  cast  a  shadow  at  the  time  the  instru- 
ments were  read.  It  was  thus  the  most 
subjective  of  all  measurements  taken  but, 
nevertheless,  provided  some  estimation  of 
cloud  cover  in  this  region.  Wind  totals 
were  entered  in  mile  units  read  from  the 
anemometer  dial  each  day,  calculated  as 
the  difference  from  the  value  of  the  pre- 


214 


GREAT  BASIN  NATURALIST 


Vol.  35,  No.  2 


ceding  day,  and  converted  and  printed 
out  as  kilometers.  The  final  daily  wind 
total  for  any  preceding  year  is  listed  as  a 
starting  value  and  included  on  the  control 
card  for  the  main  program. 

Maximum  and  minimimi  temperatures 
monitored  from  a  standard  weather  shel- 
ter, from  5  cm  beneath  soil  surface  under 
grass  cover,  from  soil  surface  under  grass 
cover,  or  from  bare  ground  were  recorded, 
converted  to  Celsius  if  not  already  in  those 
units,  and  ])rinted  onto  the  second  and 
third  sheets  for  each  month's  data.  Means 
and  extremes  for  all  values  were  stored 
for  eventual  calculation  of  totals  and  grand 
means  for  the  annual  smnmary  pages. 

Daily  maximum  and  minimimi  tem- 
peratures were  measured  in  the  weather 
shelter  by  mercury-  and  alcohol-filled 
thermometers  as  well  as  by  a  standard 
hygrothermograph.  Differences  noted  in 
the  recorded  temperatures  relate  mainly 
to  the  longer  time  lag  required  by  the  bi- 
metallic sensor  within  the  thermograph 
unit. 

Discussion 

The  main  objective  of  this  paper  was 
to  present  the  computer  programs  devel- 
oped for  our  research  on  the  effect  of 
irrigation  on  pasture  microenvironments. 
These  programs  have  proved  extremely 
satisfactory  to  us,  and  hopefully  some 
sections  or  subroutines  will  be  of  value 
to  others  engaged  in  related  research  pro- 
jects. Mitchell  and  Andersen  (1969)  re- 
ported on  a  computer  program,  deAoloped 
at  the  Uni\ersity  of  Illinois  at  Urbana, 
for  handling  meteorologic  data  collected 
from  grass  plots.  Certain  similarities  exist 
between  that  program  and  the  one  re- 
ported here,  since  the  choice  of  meteoro- 
logic instruments  and  the  overall  research 
projects  at  the  two  institutions  were 
closely  correlated.  The  program  at  Illi- 
nois, however,  was  designed  to  handle 
some  additional  measurements  not  taken 
in  the  current  study,  such  as  solar  radi- 
ation. The  Illinois  study  also  gave  em- 
phasis to  conversion  data  for  a  series  of 
soil-moisture  and  soil-temperature  mea- 
surements obtained  through  moisture-cell 
leads  (wafers)  and  built-in  thermistor 
units.  The  programs  developed  at  BYU 
use  data  on  soil  moisture  only  from  simj)- 
lified  gravimetric  measurements  but  in- 
clude the  techniques  designed  for  Cal- 
comp   graphing   as   well.     The   advantage 


of  this  plotting  program  is  not  only  in  the 
funds  saved  through  not  having  to  man- 
ually plot,  trace,  and  label  all  such  graphs 
but  more  particularly  in  the  marked  re- 
duction of  errors  that  invariably  accom- 
l)an3"  the  tedious  and  laborious  tasks  en- 
countered in  manually  plotting  daily 
weather  data. 

The  impact  of  irrigation  on  the  moisture 
and  temperature  profiles  in  central  Utah 
during  1973  was  essentially  the  same  as 
that  reported  for  1970-72  by  Andersen 
et  al.  (1974).  Grand  mean  temperatures 
for  5  cm  beneath  soil  surface  under  grass 
cover,  at  soil  surface  under  grass  cover, 
or  on  bare  soil  surface  were  consistently 
lower  on  the  irrigated  plot  than  on  the 
nonirrigated  area.  These  temperature  dif- 
ferences were  most  apparent  during  the 
warm  summer  months  when  irrigation  is 
connnonly  employed  throughout  the  re- 
gion. In  July,  for  example,  the  month 
during  v.hich  most  yearly  maxima  were 
recorded,  the  average  monthly  maximum 
temperatures  measured  5  cm  beneath 
soil  surface  under  grass  cover  on  irrigated 
and  nonirrigated  plots  differed  by  6.4  C, 
by  15.2  C  for  those  measured  at  soil  sur- 
face under  grass  cover,  and  by  17.9  C  on 
bare  ground.  During  the  six  months  when 
no  irrigation  occurred,  the  differences 
were  not  so  apparent;  hence  the  grand 
means  for  the  year  do  not  give  an  ac- 
curate reflection  of  these  temperature 
ranges  for  the  irrigation  season. 

The  contrast  between  soil  moisture 
measurements  in  the  two  plots  was  read- 
ily apparent  for  the  six  months  during  | 
which  soil  samples  were  gravimetrically 
analyzed  for  moisture  content.  The  grand 
mean  for  the  percent  soil  moisture  for  that 
period  was  22.7  percent  for  the  samples 
removed  from  the  irrigated  section  and  9.4 
percent  for  those  from  the  nonirrigated 
area. 

Other  meteorologic  measurements  which 
were  taken,  such  as  the  precipitation  pat- 
tern for  rain  or  snow,  relative  humidity, 
]iotential  evaporation,  and  wind,  were 
monitored  for  both  sections  combined,  ' 
since  the  instruments  used  could  not  de- 
tect flifferences  from  microenvironments. 
The  general  pattern  of  these  values  was 
similar  to  that  recorded  for  pre\'ious  years. 
The  collective  data  for  1973  emphasize 
the  marked  beneficial  impact  of  irrigation 
in  creating  favorable  microenvironments 
for  Ha  inij;  organisms. 


June  1975 


ANDERSEN,  ROPER:    METEOROLOGIC  DATA 


215 


0002 


0005 
0006 


0007 
0008 
0OO9 
OOLO 
OOll 
0012 
0013 


0014 
0015 
0016 


0017 
0018 
0019 


0020 
0021 
0022 
0023 
0024 
0025 
0026 
0027 
0028 
0029 
0030 
0031 
0032 
0033 
0034 
0035 
0036 
0037 


Appendix 

*  weather  data  analysis,   part  i 

♦ 

*  DEVELOPED  BY  FERRON  ANDERSON,  BYU 

*  PROGRAMMED  BY  PAUL  ROSS  ROPER,  ER I 
4< 

^L'^^if^m************************  *************************  *^L^:t  ****** 


INTEGER 


NDAYS,    DAY, 
CLCV,    WIND, 


OYR,     OEWl,     IRRG,     DEW2,    HHUM,     LHUM,     T 
OLOW,     S2(26,5),     M0N(12),     HOLD,    MONTH 


REAL 


•MARC  , 
•OCTO' , 
'L  ', 

•0    ME* , 
SNOW, 


•APR  I 
•NOVF 
3*« 
8*« 

SNCV,    MOSI,    MOSN,    PEVP, 
Sl(26,8),    S3(13,24), 


•MAY  ' 
•DECE' 
•ST  ' 
•R         " 


'JUNE' 
'GRAN' 
•EMBE' 
3*'       • 


CCMMON 


THRS(35) 


'JANU',     'FEBR 

•AUGU*.     'SEPT 

•UARY',     'H 

•MBER'f     'MBER 

TPRE,    RAIN,    PRSN 

IRDW,    NIWW,    NIDW,     T(34,16) 

PAGE2(34,12).     PAGE3(34,15) 

NOAYS,  0AY(3l»,  DYR(3l),  TPRE(35),  RAIN(35),  PRS 
k  SN0W(35),  SNCV(35>,  M0SI(35),  0EW1(32),  IRRG(32) 

)  M0SN(351,  DEW2I32),  HHUM(35),  LHUM(35) 

:  PEVP(35),  CLCV(35),  WIN0(35) 

DATA  S1,S2,S3  /208*0.0,  130*0,  312*0.0/ 
TAPE  ID 

DATA  ITAPE  /5/ 
CENT(X)  =  (X-32.0)*5. 0/9.0 

REWIND  ITAPE 


LOOK  FOR  CORRECT  YEAR 

1  REAO<5,200,END=999»     NYEAR 

2  READ( ITAPE, 200, END=999I     lYEAR, 
IFdYEAR.EQ.NYEARI     GO    TO    4 

DO    3     I=l,IOAYS 

READ! ITAPE,200,END=999) 

3  CONTINUE 
GO  TO  2 


HRS, 
(13,31/ 

•JULY'  , 
•ARY  ', 
'BER  ', 
'AN   •/ 

IRWW, 


IDAYS,  MON,  OLDW 


PROCESS  DATA  PAGE  BY  PAGE 

4  DO  900  11=1,12 

CALL  HEAD  ( I YE AR , II ,  1 , MONTH) 
NOAYS  =  MON( II  I 

WRITE  PAGE  1  HEADING 

WRITE(6,120I 
DO  10  1=1, NOAYS 

READ(  ITAPE, 20H    DAYd),    DYR  (  I  )  ,     RAIN(I),     SNOW(I),     SNCVM),     IRWW, 
A  IRDW,    OEWKI),     IRRG(I),    NIWW,     NIDW,    0EW2<1). 

B  HHUMdJ,    LHUM(I),     THRS(I),     PEVP(I),    CLCV(I), 

C  WINO(I),     (T( I ,J),J=1, 16) 

PRSN(  I  )    =     SNOWd  )     ♦    0.  I 
TPRE(  I)    =    RAIN( I)     ♦    PRSN( I ) 
IF(OLOW.GT.WINDd))     OLDW    =    OLDW    -     1000 
HOLD    =    WINDd  ) 

WIND(U    =     (WIND(I)    -    OLDW)    *     1.6093    ♦    0.5 
OLDW    =    HOLD 
MOSId)    =    0.0 
MOSN( I)    =    0.0 

IFdRWW.NE.0.0)  MOSKI)  =  dPWW  -  IRDW)  *  100  /  IRDW 
IF(NIWW.NE.O.O)  MOSN(I)  =  (NIWW  -  NIDW)  ♦  100  /  NIOW 
CALL    PRINTL     (1,11) 

=    CENT(T(I,l)) 

=    CENT(T(I,2) ) 

=    CFNT(Td,3)  ) 

=    CENTITd  ,4)  ) 

=    T( 1,5) 

=    T( 1,6) 

=    T(  I,  ID 


PAGE2d  ,1) 
PAGE2( 1,2) 
PAGE2( 1,4) 
PAGE2d,5) 
PAGE2d,7) 
PAGE2( 1,8) 
PAGE2( I. 10) 


216 


GREAT  BASIN  NATURALIST 


Vol.  35,  No.  2 


0038 
0039 
0040 
0041 
0042 
0043 
0044 
0045 
0046 
0047 
0048 
0049 


0050 
005L 
0052 
0053 
0054 
0055 
0056 
0057 
0058 
0059 
0060 
0061 
0062 
0063 
0064 
0065 


0066 
0067 


0069 
0070 


0071 


)072 

0C73 
0C74 
0075 
0C76 
0077 


0C78 
0079 
0080 
0081 
0082 
0083 


PAGE2(I.in    =    T(I 

12) 

PAGE3(I,I> 

=    PAGE2(  I, I) 

PAGE3(I,2) 

=    PAGE2(I,2) 

PAGE3(It4) 

=    T(I 

7) 

PAGE3( 1,5) 

=    T(I 

8) 

PAGE3( 1,7) 

=    T(I 

13) 

PAGE3(I,8) 

=    T(  I 

14) 

PAGE3( 1,10)    =    T(I 

9) 

PAGE3(I,ll)    =    T(I 

10) 

PAGE3( 1,13)    =    T(  I 

15) 

PAGE3( 1,14)    =    T(  I 

16) 

c 

10    CONTINUE 

c 

CALL 

SUMR 

(TPRE) 

CALL 

SUMR 

(RAIN) 

CALL 

SUMR 

(PRSN) 

CALL 

SUMR 

(SNOW) 

CALL 

SUMR 

(SNCV) 

CALL 

SUMR 

(PEVPI 

CALL 

SUMZ 

(MOSI) 

CALL 

SUMZ 

(MOSN) 

CALL 

SUM! 

(HHUM) 

CALL 

SUM! 

(LHUM) 

CALL 

SUMI 

(THRS) 

CALL 

SUM! 

(CLCV) 

CALL 

SUMI 

(WIND) 

CALL 

SUMC 

( IRRG) 

CALL 

SUMC 

(OEWL) 

c 
c 
c 

CALL 

SUMC 

(DEW2) 

PAGE 

1    OF 

WEATHER 

DATA 

IFdl  .LT.5.GR.II.GT.10)    GOTO    20 


WRITE(6,l2l)  TPRE(32), 

A  M0SI(32), 

B  PEVP(32), 

WPITE(6,122)  (TPRE(I), 

A  MOSI(I), 


PRSN(32),     SN0W(32),    SNCV(32), 
IRRG(32),    M0SN(32),    0EW2(32), 


RAIN(32) 

0EW1(32) 

WIND(32) 

RAIN(I),    PRSN(I),     SNOW(I),     SNCVd), 

MOSNd),     HHUM(I),     LHUM(I),    THRS(I), 


F)  PEVPJn,     CLCV(I),    V<IND(  I  ),  !  =  33,35) 

GOTO    30 

20    WRITF (6,123)  TPRE(32),     RaiN(32),     PRSN(32),     SN0W(32),     SNCV{32), 

A  WINO(32) 

WRITE(6,124)  (TPPF(I),     R4IN(n,     PRSN(I),     SNOW(I),     SNCV(I), 

A  HHIJM(I),    LHUM(!),     THRS(I),    CLCV(I),     W  i  ND(  I)  ,  I  =  33  ,  35  ) 


P&GE    2    OF     WEATHER    DATA 


30  CALL     HEAD     (  I  YE AP , I  I  ,  2 , MCNT H) 
CALL    CALC     (PAGE2,12) 
WRITE(6,130) 

00    31     I=1,N0AYS 

31  WPITE(6,l3l)     nAY(I),    OYR(I), 
WRITe(6,l32)     ( (PAGE2(I ,J) , J= 

PAGE    3    OF     WEATHER     DATA 


(P4Gfc2(I  ,  J),  J=l, 
1,12) , 1=32,34) 


CALL     HEAD     (I  YE  AR  ,  I  I  ,  3,  MONT  H  ) 
CALL    CALC     (PAGE3, 15) 
WRITE(6,140) 
DO    40     I=1,N0AYS 
40    WRITE(6,l4l)     DAY(I),    OYR(I),     ( P AGP3 ( I , J ) , J=  1 , 1 5 ) 
WR1TE(6,142)     ((PAGE3(I,J),J=1,15),I=32,34) 


1084 
0085 
0086 

0087 
0088 
0089 
0090 
0091 


c 
c 

STORE    DATA    FOR    YEARL 

DO    50    J=l,2 

K=II*2*J-2 

SKK.l)    =     TPRE(3l*J) 

S1(K,2J     =    RAIN(3l*J) 

SUK,3)     =    PRSN(3UJ) 

S1(K,4)     =     SN0W(3UJ) 

S1(K,5)    =     SNCV(31*J) 

Sl(K,6)     =    M0SI(3UJ) 

June  1975 


ANDERSEN,  ROPER:    METEOROLOGIC   DATA 


217 


S1(K,7) 
Sl(K,8) 
50  S2(K,5) 
S2(K,1) 
S2(K,2) 
S2(K,3) 
S2(K,4) 


MOSNOUJ) 

PEVPOUJ) 

WINO( 31+J) 

HHUM(33) 

LHUM(33) 

THRS(33) 

CLCV(33) 
DO    51    J  =  l,  12 
S3( II , J)    =    PAGE2(32,JJ 
51    S3(II,J«-12)    =    PAGE3(32,J*3) 
900    CONTINUE 


YEARLY    SUMMARY    PAGE 

00    54    J=l,8 
S1(25,J)    =    0.0 
Sl(26,J)    =    0.0 
DO    53    1=1,12 
IF(J.LT.6)     GO    TO    52 
IF  (  I  .LT.5.nP.  I  .GT.  rj)     GO    IP    53 
52    Sl(25,J)    =     SI<25,J)     ♦■    Sl(I*?-l,JI 
Sl(26, J) 


SI(26, J) 
53    CONTINUE 

Sl(26,  J)     =    Sl(26,J  ) 

IF(J.GT.5)     51(26, J) 
5^    CONTINUE 

m    56    J=l, 5 

S2(25,J»     =    n 

S2(26,  J)     =    0 

DO    55     1  =  1,  12 

S2(25,J)    =    S2(25,J) 

55  52(26, J)     =     52(26, J) 

56  52(26,  JJ     =    S2(26, J  ) 
00    58    J=l,24 

S3(  13,  J)     =    0.0 
00    57     1  =  1,  12 

57  53( 13, J)    =    S3(  13, J) 
5a    53(13,  J)    =    S3(  13, J) 

WPITE(6,160)     TYtiP, 
WRITE (6,161) 


Sl<  1*2,  J) 


12,0 
51(26, J) 


S2( 1*2-1, J) 
S2(  1*2, J) 
12.0 


53( I  ,  Jl 
12. C 


*JRITE     SUMMARY    PAGE 


DO    61     1=1, 12 

K     =     I     *     2     -     1 

L     =    K     «■     I 

IF( I.LT.5.0R. I .GT. 10)     GO    TO    60 

WRITE(6,162)     (MONTHd,  J),  J=l,3)  ,     (  5  1  (  K  ,  J  )  ,  J  =  1  ,  5  )  ,  S  1  (  K  ,  8  )  ,     52(K,5) 
A  (S1(L,J), J  =  l,7)  ,     {S2(L,J ) ,J=l  ,3) ,     51{L,8),     52(L,4), 

B  S2(L,5) 

GOTO    6  1 

60  WRITE(6,163)     ( MONT H (  I , J  )  , J  =  I  , 3 )  ,     ( 5 1 ( K  ,  J )  , J= 1 , 5 )  , 52 ( K , 5 )  , 
^  (S1(L,J), J  =  l  ,5)  ,     (S2( L,J)  ,J=1,5) 

61  CONTINUE 
WRITE  (6,16<.)     (Sl(25,J) 

A 


J=l,5) 
( S1(26,J)  , J=l  ,7) , 
( 52(26, J) ,J=4,5) 


51(25,8)  ,    S2(25,5) , 
(S2(26,J), J=l,3) ,     51(26,8) 


WRITE(6,165)     TYEAR,     lYEAR 

WRITE(6,166) 

k^RITc(6,167)     (  (MONTHd  ,J)  ,  J=l,3)  ,     (  S3  (  I  ,K  )  ,  K=  1  ,  1  2  )  ,     1=1,13) 

WRITE (6,168} 

WRITE(6,167)     ( (MONTHd  ,J), J=l,3) ,     (  S3  (  I  ,K  )  , K= 1 3 , 24) ,     1  =  1,13) 

WRTTE(6,169) 

GOTO    1 


999     STOP 

120    FORMATCO'  ,T23, 'PRECIPITATION    (  MM)  •  ,  T65  ,  •  SO  IL    MOI  STURE  •  ,  T  90  , 

A  'RFLATIVE' ,/,T7, 'DAY*  ,T90, 'HUMIDITY  MRS    OF  P0T',5X 

Pi  'CLOUD' ,/,T7, 'OF  TOTAL  R  A  IN  •  ,  3(  4X  ,  '  SNOW' )  ,  T60  , 

C  'IRRIGATED' ,T76,'N0N-IRR!G' ,T102, '98-100  EVAP',4X, 

D  "COVER  V/IND',/,'     04Y       YE  AR  •  ,  3  (  4X  ,  •  PREC  •  )  ,  •  FALL«,4X, 

E  'COVeR* ,5X, 'PERCENT    DEW     IRR*       PERCENT    DEW  MAX       MIN',4X, 

F  'REL-HUM  (MM)  (0-10)        (KM)',//) 


218  GREAT  BASIN  NATURALIST  Vol.  35,  No.  2 

0147  121    FOP.MAK '-TOTALS  •  ,  5F  8  .  1  ,  F  U  .  1 ,  2  I  4,  F  I  0.  1  ,  I  4,  T  I  .J9,  F8.  I  ,  7X  ,  I  8) 

H48  122    FORMAK 'OMEANS  '  ,  5f  8.  I ,  F  I  I  .  1  ,  T73  ,  F  8.  1  ,  T8  8,  2  I  5  .  I  8  ,  F  1 1 .  1  ,  I  7,  I  8  ,  /  , 

A  'OEXTRFMES     ',/, 

8  •  HIGH' ,5F8.l,Fll. l,T73,F8.l,T88,2I5, I8,F ll.l  ,1  7,  I  8,/, 

C  •  LOW     ' ,5F8.1,F 11. l,T73,F8. 1, T88,2IDt I8,Fll.  1 ,1  7,  I  8,/, 

0  ■-*    DATE    CF     IRRIGATION     (MAY    THROUGH    OCTOBER)') 

n49  123    FQRMAK  '-TOTALS  '  ,  5F  8  .  I ,  T  124  ,  I  8  ) 

0150  124    FQRMATCOMEANS  '  ,  5F  8  .  I  ,  T88  ,  2  I  5  ,  I  8,  1 IX,  I  7  , 1  8  ,  /  ,  •  OEX  TR  EMES  •  ,  /  , 

A  •  HIGH', 5Fd. l,T88,2I5, 18, IIX,  17,18,/ , 

R  •  LOW     ' ,5P8.1, T88, 215,  18,  IIX, 17,18,/,  •-*    DATE    OF    ', 

C  'IRRIGATION     (MAY    THROUGH    OCTOBFR)') 

0151  130    FnRMAT( 31X, 'WEATHEP     SHELTER     TEMPERATURES',  32X,'5    CM    BENE 

AATH    SOIL    SURFACE'  ,// ,9X,'nAY'  , 13X, ' THERMOMETER' , 16X ,  •  HYGROT HE RMOG" 
PAPH',16X,'IRPIG4TED',18X, 'NON-I RR 1 G AT EO' , / , 9X , • OF • , / , 5X , 
C     'DAY    YEAR ' ,4( 8X, ■ MAX.  MIN.  MEAN     '),/)    . 

D152  131    FORMAT  ( 5X , A  2 . 2X , A3 , 4 ( 5X , 3F8 .  1 )  ) 

0153  132    FOPMATJ'-  MEANS        • , 4( 5X , 3Fe . I ) , / , ' 0  EXTREMES',/, 

A  8X, 'HIGH' , 4( 5X,2F3. 1,4X,A4) ,/, 8X, 'LOW    • ,4 ( bX , 2F8 . 1 , 4X , A4 ) , 

B  /,'-  ALL     TEMPERATURES     IN    DEGREES    CELSIUS') 

0154  140    FOFMAK  16X  , 'WEATHER     SHELTE  R  '  ,  1  5X,  '  SOI  L    SURFACE    UNDER    GRASS    COVER', 

A       27X,'BARE    SOIL     SUR FACE*  , // ,5X  ,  • DA Y'  ,  I  OX ,  ' THERMOMETP R '  , 4X, 
fl       2(11X,'IRRIGATEO',14X,'NON-IPRIGATFD',3X),/,5X,'OF',/, 
C    '     DAY    YEAR' ,6X, 'MAX.  MIN.  MEAN ' , 4 ( 7X , ' MAX .  MIN.  MEAN'),/) 

3155  141    FORMAT  (  IX , A2 . 2X , A3 , 5 ( 4X , 3F7.  I )  ) 

C156  142    FGRMAT(  '-MEANS       •  , 5 ( 4X , 3F 7. I ) , / , ' OE XTREME S ' , / i '  HIGH', 

A  5(4X,2F7. 1,3X,A4} ,/,'  LOW    • , 5( 4 X , 2F 7 . 1 , 3X , A4 ) , / , 

B  '-ALL    TEMPERATURES     IN    DEGREES    CELSIUS") 

0157  IfaO    FORMAT('l       ',14,'     SUMMARY    -    PR E CI P I TA T ION ' , 95X , ' P AR T    1',///,51X, 

A        14,'     SUMMARY    OF     METEURQLOGIC    0 AT  A • , / , 55 X , • BYU    PARASITOLOGY    STATI 
BON' ,/  ,61X,'PR0V0,     UTAH',//) 

0158  161    FORMAK  32X, 'PRECIPITATICN     (  KM)  •  ,  T66  ,  '  SOI  L    MOI  STURE*  ,T84  , 

A  •RELATIVE',/,T6  8,'(PERCENT)',T84,'HUMIDITY',T9  7, 

B  'MRS    OF  POT  CLOUD' ,/,24X, 'TOTAL  RAIN', 

C  3(  •  SNOW' )  ,T97, '98-100  EVAP          COVER         UIN0',/,3X, 

0  'MONTH' ,16X,3('PREC  '),'FALL  COVER  IRRG       ', 

E  'NON-IRRG  MAX  MIN  REL    HUM       (MM)  (O-IO)        (KM)') 

0159  lf>2    FORMATCO       '  ,  3  A4  ,  '  T  HTAL  '  f  5F  8.  I  ,  5(  6X  , ' -- •  )  ,  F  8.  I  ,oX  ,  • -- '  ,  I  8  ,  /  , 

A  15X,'MEAN    '  ,7F8.  I, 3I8,F8.1,2I8) 

0160  163  FORMAT('0   ' , 3A4 , • TOTA L ' , 5F8 . I , 5( 6X , ' -- • )  , 6X, ■ ND  '  , 6X, ' -- '  ,  I  8 , /  , 

A        15X,'MEAN  • , 5F6. 1,2(6X,'ND' ) ,318,6X,«ND' ,218) 

0161  164  FORMATI'-   YEARLY  TOTAL      ' , 5F 8.  I  ,5 ( 6X , '  —  •),  F 8. 1 , ' *' , 5X , ' -- ' , 

A        l8,/,'0   GRAND  MEAN        •  , 5F 8 . 1 , I  X  ,2 ( F 7 .  1 , ' * ' ) , I  7, 21 8 , 

3        F8.  l,'*', 17, 18,/,'-   ND  =  NOT  DETERMINED',/,'    *  CALCULATE 

CD  1  MAY  THROUGH  31  OCTOBER  ONLY') 

0162  165  FORMAT('l   ',14,'  SUMMARY  -  TE MPER ATURES • , 95X , • P ART  2',///,5lX, 

A  14,  •  SUMMARY  OF  METEOPOLOGIC  0  AT  A • , / , 55X , ' BYU  PARASITOLOGY  STATIC 
BN* ,/ ,61X,' PPOVO,  UTAH',//) 

0163  166  FORMAT! 34X, 'WEATHER  SHELTER  TEMPERATURES',  32X,'5  CM  BENE 

AATH  SOIL  SURFACE',//,28X,'THERMnMETFR',16X, 'HYGR CTHERMOGR AP H' ,  16X, 
B' IRRIGATED' ,18X, 'NON-IRRIGATED' ,/, '0   MONTH ' , 8 X , 4 ( 8X, ' MAX .     MIN. 
C     MEAN  ' ) ,/) 
J164  167  FORMAT        (12('    '  ,  3A4 , 4 ( 5X , 3F 8 . I ) / ) ,  • 0   •  , 3 A4 , 4( 5X , 3FS . 1 )  , // ) 

0165  168  FORMAT(  '-'  ,32X,' SOI L  SURFACE  UNDFR  GRASS  C OVER ', 34X ,' BARE  SOIL  SUR 

AF ACE ',//,  I  IX, 2(  18X ,' IRRIGATED'  ,  18X, ' NON- I RR  IGAT ED ' ),/,'0   MONTH', 
B    7X,4(9X, 'MAX.     MIN.     MEAN'),/) 

0166  169  FORMAK'-   ALL  TEMPERATURES  IN  DEGREES  CEL  S  lUS  •  ,  /  , '  I '  ) 
U67  200  F0PMAT(20I4) 

01^8  201  F0RMAT(A2,A3,F3.2,4F3.  l,2Al,2F3.1,Al,2I3,I2,F3.l,I2,I3, 

A        /,5X,16F3.0) 
0  169  END 

JOOl  SUBROUTINE  HEAD  (  1  YE AR  ,  M,  I PG, MONTH ) 

C 

C      WRITE  A  HEADING  AND  PAGE  NUMBER  FOR  EACH  MONTH 
C 
0002                 INTEGER   NOAYS,  LAY,  DYR,  OEWl,  IRRG,  DEW2,  HHUM,  LHUM,  THRS, 

A  CLCV,  WIND,  LINE(IO),  LINl(2),  LIN2(2).  M0NTH(13,3) 

C003  RE»L      TPRE,  RAIN,  PRSN,  SNOW,  SNCV,  MC S I  ,  MOSN,  PEVP 

0004  roMMnN    NDAYS,  DAY(31),  DYR(31),  TPRE(35),  RAIN(35),  P«SN(35), 

A  SNOH(35),  SNCV(35),  MnSI(35),  0EW1(32),  IRRG(32), 

8  MOSN(35),  DEW2(32),  HHUM(35),  LHUM(35),  THRS(35), 

C  PEVP(35),  CLCV(35).  WIND(35) 


June  1975 


ANDERSEN,  ROPER:    METEOROLOGIC  DATA 


219 


WRITE(6,100)  (MONTH(M,I ) ,1=1,3) ,  lYEAR, 
\  lYEAR 

RETURN 


IPG,  {MONTH(M,  I  )  ,  1=  1,3), 


ENTRY  PRINTL  (1,11) 

PRINTL  FORMATS  DAILY  DATA  FOR  PRINTING 
INCLUDES  SCALING  AND  SUPPRESSING  ZERO  RESULTS 

CALL  ALPHA  (TPRE(I),  LINE(1»,  25.4) 

CALL  ALPHA  (RAIN(I),  L  I  NE  (  3  >  ,  25.'*) 

CALL  ALPHA  (PRSN(!),  LINE(5),  25.4) 

CALL  ALPHA  (SNOW(I),  L ! NF ( 7 ) ,  25.4) 

CALL  ALPHA  <SNCV(I),  L I NE ( 9 ) ,  25.4) 

CALL  ALPHA  (MOSKI),  LINl(l),  l.O) 

CALL  ALPHA  (HnSN<I),  LIN2(l),  l.O) 

IF( II .LT.5.0R. II.GT.IO)  GOTO  2 

I«RITE(6,101)  DAY(I),  DYR(I),  LINE,  LINI,  DEWKI),  IRRG(I), 
A  LIN2,  DEW2(I),  HHUM(  I)  ,  L  HUM  {  I)  ,  THRSd),  PEVP(I), 

3  CLCVI  I  )  ,  KIN0( I  J 

RETURN 
2  WPITE(6,I02)  DAY( I) ,  DYRCI),  LINE 
A  CLCV(  I  )  ,  WIND( I ) 

RETURN 


HHUM(I),     LHUM(I),     THRS(I), 


I  1  ,/// ,49X,  'MONTHLY     SUMMARY    0 
/,61X, 


100  FORMATI'l  • ,2A4,A2,I4,108X, 'PAGE 
AF    METEORULUGIC    DA T a • ,/ , 55X  ,  • BYU    PARASITOLOGY    STATION 
B'PPOVO,    UTAH«,//,60X,2A4,A2,I4,/) 

101  F0RMAT(lX,A2,3X,A4,5(2X,A4,A2),6X,fl4,A2,2(3X,Al),4X,A4,A2,3X,Al 
A  3X,2I5,I8,Fll.I,I7, 18) 

102  F0RMAT<1X,A2,3X,A4,5(2X,A4,A2),T88,2I5,I8,T117,I7,I8) 
END 


SUBROUTINE    ALPHA    (R,C,S) 


CONVERT  R  SCALED  BY  S 
(RETURNS  BLANK  CHARACT 


INTO  ALPHA  CHARACTERS  AND  RETURN  IN  C. 
EPS  IF  ZERO) 


INTEGER  M(30l)  / 
7'  , 
16«  , 
25', 
34"  , 
43«  , 
52*  , 
61', 
70'  , 
79*  , 


INT 


97*  , 
106«  , 
115*  , 
124'  , 
133', 
142*  , 
GER  N3 
158'  , 
167', 
176'  , 
185'  , 
194*  , 
203', 
212', 
221', 
230'  , 
239'  , 
248'  , 
257'  , 
266'  , 
EQUIVALENCE 


17« 
26* 
35' 
44* 
53* 
62* 
71' 


89«  , 
98*  , 
107*  , 
116', 
125'  , 
134"  , 
143'  , 
275)  / 
159"  , 
168'  , 
177', 
186'  , 
195'  , 
204"  , 
213*  , 
222*  , 
231', 
240'  , 
249"  f 
258'  , 
267'  , 
(NL(  I 


INTEGER  N2(  11) 


R  =  R  *  S 
C{1)  =  BLANK 


0' 

9' 

18' 

27' 

36' 

45' 

54" 

63' 

72' 

81 

90' 

99" 

108' 

117' 

126' 

135' 

144' 

151 

160' 

169' 

178' 

187" 

196' 

205' 

214" 

223' 

2  32' 

241 

250" 

259" 

268" 

2)  ,n: 

.0" 
.9'/, 


I', 

2' 

10'  , 

11' 

19'  , 

20' 

28', 

29' 

37', 

38' 

46'  , 

47' 

55', 

56' 

64'  , 

65' 

73', 

74' 

82'  , 

83" 

91', 

92. 

100'  , 

101' 

109', 

llO' 

118', 

119' 

127'  , 

128' 

136', 

137' 

145', 

146' 

152', 

153' 

161'  , 

162* 

170', 

171' 

179'  , 

180' 

188', 

189' 

197'  , 

198' 

2C6', 

207' 

215', 

216' 

224', 

225" 

233', 

234' 

242', 

243" 

251', 

252' 

260', 

261' 

31 

269', 
D) 

270' 

'. 

L','.2' 

'.3', 

, 

JLANK/' 

•/ 

'          3', 

1          4t  ,. 

'       12', 

•        13',' 

'       21', 

22'  ,' 

•       30', 

31', • 

'       39', 

•       40',' 

'       48', 

49'  ,' 

•       57', 

58',' 

'       66', 

•       67',' 

•       75', 

'        76',' 

'       84', 

85',' 

•       93', 

.       94.,. 

•    102', 

•    103', • 

•     111', 

112' ,' 

•    120', 

'     121',' 

•     129', 

'     130',' 

•    138', 

139',' 

•    147', 

148',' 

•    154', 

155', • 

•    163', 

164',' 

'     172', 

•     173',' 

•    181', 

182',' 

•     190', 

191',' 

•    199', 

2)0', • 

•    208', 

209' ,' 

•    217', 

218'  ,' 

•    226', 

227',' 

•    235', 

236',' 

'    244', 

245',' 

•    253', 

254',' 

•    262', 

263' ,' 

•    271', 

272',' 

.4', '.5 

,'.6',' 

,    C(2) 

5', 

6', 

14', 

15'  , 

23', 

24"  , 

32', 

33', 

41', 

42", 

50', 

51', 

59', 

60'  , 

68', 

69'  , 

77', 

78'  , 

86', 

87', 

95', 

96', 

104', 

105', 

113', 

114'  , 

122', 

123'  , 

131', 

132'  , 

140', 

141', 

149', 

150'/ 

156', 

157', 

165', 

166'  , 

174', 

175'  , 

183', 

184'  , 

192', 

193', 

2-)!', 

202', 

210', 

211', 

219',' 

220', 

228', 

229', 

237', • 

238', 

246", 

247'  , 

255',' 

256', 

264', 

265'  , 

273',' 

274'/ 

220 


GREAT  BASIN  NATURALIST 


Vol.  35,  No.  2 


0008 
OC09 
0010 
0011 
0012 
0013 
0014 
0015 
0016 


C(2)  =  BLANK 

P  1  =  R  ■•■  0.05 

1  1  =  Rl 

12  =  (Rl  -  ID  ♦  10 

IF( I l.EQ.O. AND.I2.EQ.0J  RETURN 

C(  U  =  Nl(  lUl) 

C(2)  =  N2( I2*l) 

RETURN 

END 


0001 


0002 
0003 
0004 
0005 
0006 
0007 
0008 
0009 
0010 
0011 
0012 
0013 
0014 
0015 
0016 
0017 
0018 
0019 
0020 
0021 
0022 
0023 
0024 
0025 


SUBROUTINE  CALC  (X,N) 

CALC  PROCESSES  REAL  DATA  IN  X  AND  RETURNS  SUMS  IN  ROW  32, 
MEANS  IN  ROW  33,  MINIMUMS  IN  ROW  34,  AND  MAXIMUMS  IN  ROW  35. 

REAL  X(34,NI,  5(35) 
REAL  BLANK/*     •/ 
LOGICAL  ZERO 
COMMON  NOAYS 
DO  10  J=l,N 
X(32,J»  =  0.0 
X(33,J)  =  -100.0 
X(34,J)  =  1000.0 
DO  10  I=1,NDAYS 
IF(MOD( J,3).NE.O)  GO  TO  9 
X(  I, J)  =  (X(I ,J-2) 
X(32,J)  =  X(32,J) 
GO  TO  10 
9  X<32, J)  =  X(32,J)  ♦  X( I, J) 

IF(X(33,J).LT.X( I,J) )  X{33,J)  =  X(I,J) 
IF(X(34,J).GT.X( I,J) )  X(34,J)  =  X(I,J) 

10  CCNTINUE 
DO  11  J=1,N 

X(32,J)  =  X(32,J)  /  NDAYS 
IF(MOD(J,3».NE.0»  GO  TO  11 
X(33,J)  =  BLANK 
X(34,J»  =  BLANK 

11  CONTINUE 
RETURN 


■  X(I,J-1) )  /  2.0 
X(  I,J) 


0026 


0027 
0028 
0C29 
0030 
0031 
0032 
0033 
0034 
0035 
0036 
0037 
0038 
0039 
0040 
0041 
0042 


ENTRY    SUMR     (S) 

PROCESS    REAL    DATA     IN    ARRAY    S    AND    RETURN    SUM    IN    S(32) 
S(33),    MINIMUM    IN    S(34),    AND    MAXIMUM    IN    S(35). 

ZERO    =    .FALSE. 

1  S(32»     =    0.0 
S(33)    =    0.0 
S(34)     =    -1000.0 
S(35)     =    1000.0 
N    =    0 

DO    2    I=1,NDAYS 

IF(ZER0.AND.S(1) .LT. 0.001)     GOTO    2 

N    =    N    ♦    I 

S(32l    =    S(32)     ♦    S( n 

IF(S«34I.LT.S( n »     S(34)     =     S(I) 

IF(S(35).GT.S( I) )     S(35)    =    S( II 

2  CCNTINUE 
IF(N.EO.O)     RETURN 
S(33)     =    Sf32)     /    N 
RETURN 


0043 
0044 
0045 
0046 


ENTRY    SUMZ     (SI 
ZERO    >=     .TRUE. 
GOTO     1 
END 


SUBKJUTINE     SUMI     (lY) 


PROCESS    INTEGER    DATA     IN    ARRAY     lY    AND    RETURN    SUM     IN    IY{32», 
ME4N     IN    IY(33),     MINIMUM    IN     IY(34I,     AND    MAXIMUM     IN    IY(35). 


INTEGER     IY(35),    PLUS/'*'/ 


ANDERSEN,  ROPER:    METEORO LOGIC  DATA 


221 


CCMMON  NDAYS 
IY(32»  =  0 
IY( 3^)  =  0 
IY(35)  =  1000 
DO  13  I=l,NOAYS 
I Y(32)  =  I Y(32»  ♦  IY(I  ) 
IF(IY(34» .LT. IY(  I )  )  IY{34) 
IF( I Y(35).GT. IY( ! ) )  IY{35) 
13  COM  iNue 

I Y( 33)  =  IY(32) • /  NDAYS  ♦ 
RFTURN 


ENTRY  SUMC  (lY) 


lYtl) 
IY(I  ) 


PROCESS  CHARACTER  DATA  IN  ARRAY  lY  AND  RETURN  THE  NUMBER  OF 
IN  IY(32). 

IY( 32)  =  0 

DO  I't  1  =  1, NDAYS 

IF(IY(  D.EC.PLUS)  IY(32)  =  IY(32)  ♦■  1 

l^  CCNTINUE 
RFTURN 
FND 


WEATHER  DATA  ANALYSIS,  PART  II 


PROGRAMMED  BY  PAUL  RCSS  ROPER 

EYRING  RESEARCH  INSTITUTE,  197^ 


Hc*t*^f****j^i^****^t^^*t^t^f7t:^^it:tti¥**  *********  *********************** 


DECK  FORMAT 


I.  CONTROL  CARD 


2.  DATA  HEADER 


COMMON  /PLOTO 


/PLOTl 
/PL0T2 
/PL0T3 
/PL0T4 
/PL0T5 
/PL0T6 
/PLOT? 
/PL0T8 
/PLQT9 
/PLOTIO/ 
/PLOTll/ 
/PL0T12/ 
/PL0T13/ 
COMMON  /LABELS/ 


CCNTROL  CAPO 
DATA  HEADER 
DATA 


(ON  TAPE 
(ON  TAPE 


IF  DESIRED) 
IF  DESIRED) 


COLUMNS  DESCRIPTION 

1-A  YEAR 

5-8  DESIRED  GRAPH  BY  NUMBER 

9-12  ...  13  GRAPHS  POSSIBLE 

1-4        YEAR  OF  DATA 

5-e  NUMBFP    OF    DAYS     IN    YEAR 

9-12  LAST    WIND    READING    OF     PREVIOUS    YEAR 

SEE    PUBLICATION    FOR    CARD    COLUMNS    AND    DATA 
DESCRIPTION. 


XPLT( 
RAIN( 
SNOW( 
YHHD( 
HUMO( 
EVAP( 
WIND( 
YMAXC 
YH5C( 
YHAK 
YHSS( 
YHA2( 
YHBG( 
YHA3( 
AXl(3 
BX(4) 
RC1(7 
RC3(l 
RC5(1 
RC7(l 


366) 
366) 
366) 
366) 
366) 
366) 
366) 
366) 
366) 
366) 
366) 
366) 
366) 
366) 
7), 

), 
!)• 
l)t 
2), 


ITAPE 

XIRR(366),     YIRR(366),    YNIR(366) 


,  YMIN(366) 

,  YL5C(366) 

,  YLA  1(366) 

,  YLSS(366) 

,  YLA2(366) 

,  YLBG(366) 

,  YLA3(366) 
AYl(37),     AX2(73) 


ZH5C(366) ,    ZL5C(366) 


ZHSS(366)  ,    ZLSS(366) 


ZHBG(366) ,    ZLBG(366) 


AY2(73) 
BY(4),  XL(4),  YL(4), 

LCl(7),  RC2( 13),  LC2( 13) 
LC3(  ID  ,  RC4(13)  ,  LC4(  13), 
LC5(18),  RC6(8),  LC6(  16) 
Lr7(24) 


CREATE    A    DEGREE    SYMBOL 


222 


GREAT  BASIN  NATURALIST 


Vol.  35,  No.  2 


0003 
0004 
0005 
0006 
0007 
0008 


0009 
0010 
0011 


0013 
0014 
0015 
0016 
0017 
0018 


0019 
0020 
0021 
0022 
0023 
0024 
0025 
0026 
0027 
0028 
0029 


0030 
0031 
0032 
0033 
0034 
0035 
0036 
0037 


1  J38 
n39 
0040 
J  HI 
D  JA2 

•)043 
0C44 
0045 
0046 
0047 
0048 


DIMENSION 


DEG(3)f    0(3),     NGRAPH(14) 


DATA  DEG  /Z05061727, 

DATA  0  /ZC1061737, 

CALL    SYMBL5    (J, 9, DEG) 
CALL    SYMBL5    (K,9,0) 
REWIND    ITAPE 

READ    CONTROL    CARD 

100    PFA0(5,200,EN0=999)     NYEAR,    N 
200    FORMAT( 1514) 
NUM    =    0 

READ    ONE    YEAR'S    DATA 

CALL    READC     ( NYE AP , NCAYS ) 


CONTROL    LCOP 

300    NUM    =    NUM    ♦    1 

M    =    NGRAPH(NUM) 

GO    TO    (1,2,3,4,5,6,7,8,9,10,11,12,13) 

GO    TO     100 
999    CALL    PLOT  (12.0,0.0,-3) 

STOP 


Z36352414,  ZC506I727/,  J  /Z9l/ 
Z46413010,  Z01061737/,  K  /ZD6/ 


C****:****!^*  *******  PLOT   I  ♦********«*****♦****»»*****»*******♦*♦**  ******* 


PRECIPITATION  AND  SCIL  MOISTURE 


( 3. 0,RC I, 4, 0.5, 7, I, LCI, 4, 0.5, 7, I) 

(RAIN,NCAYS,3.00,       3.0,       0.0,     2) 
CALL     SYMBL4    ( -0. 24 ,  I  .2 , 0. 07, 'M I LL I  MET ERS • , 90. J , II ) 
CALL     SYMBL4    ( 0. 2 , 2. 8 ,0.  1 , • PR  EC  I  PI T ATI  ON • , 0 . 0, 13 ) 
CALL    SYMBL4    (  I  .39 , 2 . 8, 0. 07 , • AND • , 0. 0, 3 ) 
CALL     SYMBL4    ( I .64 , 2 . 8, 0. 1 , • SCIL    MO  I  ST URE •  ,0.0 , 13 ) 

(NYEAR ,0.2,2.65) 

( XPLT,RAIN,NOAYS) 

(8.21,  1.3,0.0  7,' INCHES ',90. 0,6) 


CALL     AXES 
CALL    SCALE 


CALL    DATE 
CALL     BAR 


CALL     SYMBL4 

CALL     MOIST 

GO    TO    303 
C 
C 
C********»********PLOT       2*****************"****************************** 


SNOW    COVER 


(3.0,RC2,4,0.2  5,13, 1,LC2,4,0.25,13,1) 

(SNOW,N0AYS,3.00,     12.0,       0.0,     3) 
CALL     SYMBL4    ( -0 . 2 7 , I .2 , 0. 0 7 , • MI LL I  MET E RS'  , 90. 0, 1 1 ) 
CALL    SYMBL4    ( 0.2 , 2 . 8 ,0 . 1 , ' SNOW    COVER '  ,0.0 , 10) 

(NYFAR ,0.2,2.65) 

(XPLT,SNCW,NDAYS) 
CALL     SYMBL4    ( 8. 2 3, 1 .3, 0. 07 , • INCHE S' ,90.0 , 6) 


2    CALL    AXES 
CALL     SCALE 


CALL     DATE 
CALL     BAR 


GO    TO    300 


C 

C 

C**********> 

C 

c 
c 


c*****PI^OT       3*********************************************** 
RELATIVE    HUMIDITY    IN    WEATHER    SHELTER 


3  C  ML 
CALL 
C'^LL 


AXES  ( 3.0, Rr 3,^,0.25,  II,  I, LC 3,4,0.25.  I  1  ,  I  ) 

SCALE  (  YHHL  ,NCAYS,2.49,  ICO.O,        0.0,     't ) 

SCALE  ( YLHr,NCAYS,2.49,  loo.o,       0.0,     5) 

CALL     SY^BL4  { -0  . -^  5  ,  I  .  1  ,  0.  0  7  ,  '  PE  PC  ("  N  T  •  ,  90.  0  ,  7) 

CALL     SY'^PLA  (  1.2  ,  ^.8  ,T.  I,  'PhLAT!  VE     HUMIDITY     IN    WEATHER     SHELTER 

\  0.0,3b) 

CALL     DATE  ( fJYF /iP  ,  0  .2  ,  2  .  65  ( 

C^LL     LEGtrjD  (  1,7.  J, ^.6) 

CALL     LINES  ( XPL T , YHHD ,ND AYS , 0 . QI , 2 ) 

CALL     OASH  ( XPLT , YLHD  .KCAYS, J.  ri) 

CALL     SYMRL4  (  0  .  2  5  ,  I  .  1 ,  '' .  )  7  ,  '  PF  P  CF^J  T  •  ,  9  ■)  .  J  ,  7  ) 
GG    TO     300 


ANDERSEN,  ROPER:    METEOROLOGIC  DATA  223 

C  04TI  Y     nURATION    OF    RbLATIVF     HUMIDITY 

C 

r 

^    CALL     AXES  <  3.0,PC^,4,0.21,  13,  l.LC^, -^,0.21,  13,  1) 

CALL     SCALE  <  HUMO  ,  NDAY  S  ,  2  .  ^*9  ,     2^.3,        0.3,     6) 

CALL     SYM8L4  ( -0 . 2 2 ,  I  . 5 , 0. 0 7 , • HCUR S • , 9 0. 3 , 5  J 

CALL     SYM8L4  ( 0 . 2 , 2 . 8  ,  0.  I , ' D A JL Y    OHRATlnN    OF    RELATIVE    HUMIDITY     AT     9 

A8-100(',  ).1,^0) 

CALL    DATE  ( NYF AR , C.2 , 2. 6 5 ) 

CALL     BAR  ( XPLT,HUMO,NOAYS) 

CALL     SYMBL4  ( 8 . 23 ,  1 .  5, 0 . 0 7 , ' HOURS ' , 9  )  .0 , 5 ) 

GO    TO    300 
C 
C 

C«*«*«*  ««:(<*♦*  *«***PLCT  5«*****»**  «•****«*♦*«♦  **♦*♦♦«♦*:**♦«****«♦*»***«♦* 

C  POTENTIAL  EVAPCRATICN 

5  CALL  AXES  ( 3.0,RC5,4,0.28,  11,  l,LC5,  5,0.3b,l8,2) 
CALL  SCALE  (  E  VAP  ,  MCAY  S  ,  3  .  0  ) ,  21. T,   ■■^.0,  7) 

CALL  SYMBL4  ( -  J. 2 3 ,  I -2 , 0.0 7 ,  •  M I LL I MFT FRS •  ,90. 0 , 1  I ) 

CALL  SYMBLA  ( 0 . 2 , 2. 8  ,0.  I,  ' POTEN TI AL  E V APORAT I  ON'  , 0. 0, 2  I ) 

CALL  DATE  (NYEAR,0.2,2.6t)) 

C\LL  SYMBL4  ( 2 .0 , 0 . 5  ,0. C7 , • ST AR TED •  ,  D  .  T  ,7  ) 

J  =  NDAYS  -  244 

CALL  LINES  ( XPLT ( J )  ,E VAP ( J ) ,  184,  : . 01  ,  2  J 

CALL  SYMBL4  ( 7.0 , 0. 5  ,0 . C7 , • STOP PED •  ,0.0  ,  7  ) 

CALL  SYMBL4  ( 8 . 30 ,  I  . 3, 0.07 , ' I NCHF S •  ,90. 0 , 6 ) 
GO  TO  3J0 
C 

r 

f  «***«**♦***♦  j(*  +  **PLOT   6*****************  *"■**♦**«"*****♦*♦*♦**■«•  i"*  ***♦♦*♦ 

C  TOTAL  WIND  I  METFR  ABOVE  GROUND 

C 

C 

6  CALL  AXES  ( 3.0,RC6,4,0.42,e,  I  ,Lr6,5,0.42,  16,2) 
CALL  SCALE  ( W I ND  ,NDAY S ,2. 94 ,  1  7 5. 0 ,   0.0,  8) 
CALL  SYMBL4  (-0. 29,  1  .2  ,  0. 07, 'KI LOMETE RS • , 90  .0 ,  10 ) 

CALL  SYMBL4  ( 0. 2 , 2. 8  ,0. 1 ,  • TOTAL  WIND  1  METER  ABOVE  GROUND ', 0. T , 3 1 ) 

CALL  DATE  ( NYE AR  ,  0  .2 , 2. 65 ) 

CALL  LINES  ( XPLT , W  I  ND  ,  ND A YS  ,0.0  1 , 2 ) 

CALL  SYMBL4  ( 8 . 3, 1 . 4 , 0- 07 , • M I L E S ' , 90. 0 , 5) 

GO  TO  3)0 

C 

C 

C  ♦**«*♦****<.**♦**«? LOT   7*** ********************************* *«**♦*♦»*** 

C  TEMPERAT'JRE    IN    WEATHER     SHELTER 

C 

C 

7  CALL     AXES  (  4.  0  ,  R C 7 ( I ) ,4, 0. 5, 9 , I , Lf 7 ( I) , 5 , 0. 5 , 1 3  ,  2 ) 
CALL     SCALE  ( YMAX , NC AYS ,4. 00 , 122 . 0 , -22 . 0,     9) 

CALL     SCALE  ( YM IN , NC AY S  ,4. 00 , 122 . 0 , -22 . 0 ,  I  0 ) 

CALL     SYMBL4  ( -0.  25  ,  1  .2  ,  ).  07  ,  •  TE  r'PE  R  ATURE     (  JC  )  •  ,  90  .0,  16  ) 

CALL     SYMBL4  ( 0. 2 , 3. 7, 0. 1 , • TEMPE R ATURE     IN    WEATHER     SHEL TER • , 3 . 0, 301 

CALL    DATE  (  NYE  AR  ,  0  .2  ,  3.  55) 

CALL     LEGEND  (1,7.0,3.7) 

CALL    ZERO  (1.5)  • 

CALL     SYMBL4  { 8 . 32 , I  .2 , 0. 07 , • TE MPE RA TURE     ( JF  )  •  , 90 . 0  ,  16 ) 

CALL    LINES  ( XPL T , YM AX , NOA YS , 0. 01 , 2 ) 

CALL     DASH  (XPLT, YMIN,NDAYS, 0.02) 

GO    TO    3J0 
C 
C 

(-*♦♦*♦♦*♦*♦♦♦*♦♦♦♦  PLOT       8  ************************************  ****♦***♦*♦ 

C  COMPARISON    OF     MAX    AND    MIN    TEMP    5    CM    UNDER     10    CM    GRASS 
C 
C 

8  CALL    AXES  ( 3.0, RC7(3) ,4, 0.6, 6,1, LC7(5) ,5,0.6,12,2) 
CALL    SCALE  ( YH5C , ND AY S , 3. 00,    40.0,-10.0,11) 

CALL     SCALE  ( YL5r , NCAY S , 3 . 00,    40.0,-10.0,12) 

CALL     SCALE  ( ZH5C , NC AY S . 3. 00,    4  1.0,-10.0,13) 

CALL     SCALE  (ZL5C,NDAYS,3.00,    40.0,-10.3,14) 

CALL     SYMBL4  ( -0. 25 , 1 . 0 , 0. 07, • T E MPER ATURE     ( JC) ' , 90. 0, 1 6) 

CALL    SYMBL4  ( 0 . 2 , 2 . 8 ,0 . 1 , ' COMPAR I  SON    OF    MAXIMUM    AND    MINIMUM    TEMPER 

AATURES',  0.0,46) 


224 


GREAT  BASIN  NATURALIST 


Vol.  35,  No.  2 


0096 

0C97 
0C98 
0099 
0  100 
Old 
0102 
0103 
0104 
0  10  5 


0106 
0107 
0108 
0109 

oin 

Olll 

0112 

0113 
0114 
0115 
0116 
0117 
0118 
0119 


CALL  SYMRL4  ( 0 .2 , 2. 7  ,0. 07 , • 5    CM    DEEP     IN    SOIL    UNDER    10    CM    GRASS    GOV 
AER't  0.0f41) 

CALL  SYMBL4  (0 .2 , 2 . 6 , 0. 07 , ' ON    IRRIGATED    AND    NON- IRR IGATE 0    PLOTS', 
A  0-0,36) 

CALL  DATE  ( NYE AR , 0  .2 , 2. 45 » 

CALL  LEGEND  (2,6.75,2.8) 

CALL  ZERO  (0.6) 

CALL  SYMBL4  ( 8. 32 ,  1 . 0, 0. 07 , ' TE MPER ATURE     ( JF ) •  , 90 . 0, 16 ) 

CALL  LINES  ( XPL T , YH5C , NDA YS , 0. 01 , 3 » 

CALL  LINE  (XPLT,  YL5C,N0AYS,n 

CALL  DASH  (XPLT, ZH5C  ,NDAYS, 0.04) 

CALL  DASH  ( XPLT , Z L5C , NOAYS , T . 02 ) 

GO    TO    300 


C 
C 

C 

c 


COMPARISCN    OF     MEANS    5    CM    UNDER    10    CM    GRASS 


9    CALL  AXES  ( 3.0 , RC7 ( 3 ) ,4 , 0. 6 , 6 , I , LC7 ( 5 ) , 5 , 0. 6 , 1 2 ,2 ) 

CALL  SCALE       ( YHA  I , NC AYS , 3. 00,    40.0,-10.0,15) 

CALL  SCALE       ( YL A  I , NC AY S , 3 . 00,    40.^,-10.0,16) 

CALL  SYMBL4    ( -0.  2  5  ,  1 .0  ,  0.  0  7  ,  «  TEMPE  R  AT  URE     (  JO  •  ,  90  .  0,  16  ) 

CALL  SYMBL4    ( 0. 2 , 2. 8 ,0 .  I , • CCMPA R J  SON    OF    DAILY    MEAN    TEMPERATURES" 


A 
CALL 

AER', 
CALL 

A 
CALL 
CALL 
CALL 
CALL 
CALL 
CALL 


SYMBL4 

SYMBL4 

DATE 

LEGEND 

ZERO 

LINES 

DASH 

SYMBL4 


0.0,37) 
(0.2,2.7,0.07, '5  CM  DEEP  IN  SOIL  UNDER  10  CM  GRASS  COV 

0.0,41) 
(0.2,2.6,0.07,  'ON  IRRIGATED  AND  NON- I  PR IGATE 0  PLOTS', 

0.0,36) 
(NYEAR ,0.2,2.45) 
(3,6.5,2.8) 
(0.6) 

( XPLT, YHA 1 , NO AYS, 0.01, 2) 
( XPLT.YLAl ,NDAYS,0.02) 
(8-32, 1.0,0.07, 'TEMPERATURE  ( JF ) • , 90 . 0 , 16 ) 


GO  TO  300 


.«««**«******»PLGX  10***»****  ***♦♦♦*«■=•  **♦**♦*•*♦♦♦♦♦♦♦♦♦♦♦  ***♦***> 

COMPARISON    OF     MAX    AND    MIN    TEMP    AT    SOIL    SURFACE 


0120 
0121 
0122 
0123 
0124 
0125 
0126 

0127 

0128 

0129 
0130 
0131 
0132 
0133 
0134 
0135 
0136 
0137 


0  138 

0139 
0140 
0141 
0142 

")143 


10  CALL 
CALL 
CALL 
CALL 
CALL 
C«LL 
CALL 

AATUR 
CALL 

A*  . 


AXES 
SCALE 
SCALE 
SCALE 
SCALE 
SYMRL4 
SYM6L4 
ES'  , 
SYMBL4 


CALL     SYMBL4 


CALL 
CALL 
CALL 
CALL 
CALL 
CALL 
CALL 
CALL 
GO    T 


DATE 
LEGEND 
ZERO 
SYMBL4 
LINES 
LINE 
CASH 
DASH 
0    3  )0 


(3.5,RC7(3),4,C.5,8,l,LC7(5),5,0.5,16,2) 

(YHSS,N0AYS,3.50,    60.0,-10.0,17) 

(YLSS,NCAYS,3.50,    60.0,-10.0, 18) 

( ZHSS,NDAYS,3.50,    60.0,-10.0,19) 

(ZLSS,NCAYS,3.50.     60.0,-10.0,20) 

(-0.25,  I. 3, J. 0  7, 'TEMPERATURE     (JC)',9J.0,16) 

(0.2,  3.  3,0.  1, 'COMPAR!  SON    OF     MAXIMUM    AND    MINIMLIM    T^ 

0.0,46) 
(0.2,3.2,0.C7,  'AT    SOIL     SURFACE    UNDER     13    CM    GRASS    C 

0.0,39) 
( T. 2, 3.  1,0. 07,  'ON     IRRIGATED    AND    NON- I RR IGATE 0    PLOT 

0.0,36) 
(NYEAR ,0.2,2.95) 
(2,6. 75,3. 3) 
(0.5) 

(8. 32, I. 3, 0.0 7, 'TEMPERATURE     (JF)'  ,90.0,  16) 
( XPLT  ,YHSS,NDAYS,0.01,3) 
(XPLT, YLSS  .NDAYS,  1) 
(XPLT.ZHSS  ,NDAYS,0.04) 
(XPLT, ZLSS,NDAYS, 0.02) 


MPER 
OVER 
S'  , 


i^:^***  A««*> 


'♦♦***PLOT   11* ♦♦«*******♦*«**♦«****** ♦**♦*♦♦♦«***♦♦: 

COMPARISON  OF  MEANS  AT  SOIL  SURFACE 


11    CALL     AXES 


CALL 
CALL 
CALL 
CALL 


(3.'>,RC7(3),4,C.5,7,l,LC7(5),5,0.5,l4,2) 
(  YHA?.  NDAYS  .  3.  "10.     5 ')  .  n  .  -  1  '1.  U  ?  1  1 


AXES  (3.'>,RC7(3),4,C.5,7,l,LC7(5),5,0.5,l4,2) 

SCALE  (  YHA2,NDAYS,3.00,     50.  n  ,  -  I ').  0  .  2  1  ) 

SCALE  (YLA2, NCAYS,3.00,     50.0,-10.0,22) 

SYM6L4  (-0.25,  1.0,0.07,  'TEMPEPATUPt:     (  JC  )  '  ,90  .  0  ,  16  ) 

SYMRL4  ( 0.2,2.8  ,0.  I, 'CCNPARISHN    PF    DAILY     MEAN    TEMPERATURE 
0.0,37) 

CALL     SYMRL4  (  0  .  2  ,  2  .  7  ,0  .  07  ,  '  AT    SOIL     SURFACE     UNDER     10    C '^    GRASS', 


0.0,33) 


June  1975  andersen,  roper:  meteorologic  data  225 

01'*'^               CALL  SYMBL4  (0  .  2  ,  2.  6  ,  0.  07  ,  •  OM  IRRIGATED  AND  NGN- I  KR  I  G  ATFn  PLOTS', 

A  ).l,3b) 

0145  CALL  DATb  ( NYE 6P  ,  0 .2  ,  2. 4 5 ) 

0146  CALL  LEGEND  (3,6.5,2.8) 

0147  CALL  ZERO  <').5) 

0148  CALL  SYMBL4  ( 8 . 32 ,  I  .  0, 0. 0 7 , • TE MPEP A TUR E  ( J F ) •  , 90  ,  0 ,  16 ) 

0149  CALL  LINES  (  XPL T  , YH A2 , NDA YS , 0.  11 , 2  I 

0150  CALL  CASH  ( XPLT , YL 42 , NC AY S , 0 . 02 ) 

0151  GO  TO  300 
C 

C 

C  ♦**,«**«**»«*  «*«««PLfjT  I  2********'*^  ****************<'***********  ********** 

C  CUMP/KISLN  OF  MAX  AND  MIN  ON  BARE  GROUND 

C 

c 

12  CALL  AXES  (  5.  0 , P C 7 ( 2  )  ,4 , 0 . 5  ,  1  1 , 1 , LC 7( 3)  ,  5 , 0 . 5 , 22, 2) 
CALL  SCALE  (  YHBG  ,  N  F.  AY  S  ,  5  .  00  ,  8  ).  0  , -2'^  .  0,  2  3  ) 
CALL  SCALE  < YLBG , NC AY S , 5 . 00,  80.0,-20.0,24) 
CALL  SCALE  ( ZHRG, NC AYS , 5. 00,  80.0,-20.0,25) 
CALL  SCALE  ( ZLBG,NCAYS,5.00,  8  1.  )  , -2 0. 0, 26 ) 
CALL  SYMBL4  ( -0 . 25 , 2.0 , 0. 0 7 , • TE MPb P ATURE  ( JC ) •  ,  90  .  0 , I  6 ) 
CALL     SYMBL4  ( 0. 2 ,4 . 8 , 0. I , ' C CMPA R I  SON    OF    MAXIMUM    AND    MINIMUM    TEMPER 

\A TURFS',  0.0,46) 

CALL     SYMBL4  ( 0 . 2 , 4 . 6 fc, 0. 08 ,  •  AT     SOIL    SURFACE    UN    BARE    GROUND    CN', 

A  0.0,33) 

CALL     SYMBL4  ( 0 . 2 ,4 . 52 , C. 0 8  ,  •  1 P P I G AT  ED    AND    NON- I RR I  GAT  ED    PLOTS', 

A  0.0,33) 

r&LL     DATE  {NYFAR,0.2,4.37) 

CALL    LEGEND  (2,6.75,4.8) 

CALL     ZERO  (l.O) 

CALL    SYMBL4  ( 8 .32 , 2  .  0, 0. 07 , ' TE MPER ATURE     ( JF  )  •  , 90 . D , 16 ) 

CALL    LINES  ( XPLT  ,  YHBG ,NDAYS, 0. 01 , 3 ) 

CALL    LINE  ( XPLT, YLBG, NOAYS, 1) 

CALL    DASH  ( X  PL T , ZHBG , NDAYS , 0. 04) 

CALL    DASH  ( XPLT, ZLBG, NOAYS, 0.02) 

GO    TO     3  JO 

r 

c 

C *♦ 4 »*♦♦♦♦♦ »*****+PLOT     13**«*4**«******** ***♦♦♦♦*****♦♦*♦**♦* «♦******♦*♦ 

r  COMPARISON  OF  MEANS  OF  BARE  GROUND 
C 

c 

13  CALL  AXES  ( 3. 0 , R C 7( 3 ) , 4 , 0 . 5 , 7 , I , LC7 ( 5 ) , 5 , 0. 5 , I  4 , 2 ) 

CALL  SCALE  ( YH A3 , NC AY S , 3. 00,  50.0,-10.0,27) 

CALL  SCALE  ( YL A  3 , NC AY S , 3. 00,  50.0,-10.0,28) 

CALL  SYMBL4  ( -0. 25 ,  I  .0 , 0. 0 7, ' TE MPE R ATURE  ( JC ) •  ,90  .0 , I  6 ) 

CALL  SYMBL4  ( 0 . 2 , 2 . 8 ,0 . I , ' C CMPARI SON  OF  DAILY  MEAN  TEMPERATURES', 

A  0.0,37) 

CALL  SYMBL4  ( 0. 2 , 2. 7 , 0. 07 , • ON  BARE  GROUND  CN  I RR IGATED' , 0 .0 , 2 7 ) 

CALL  SYMBL4  ( 0. 2 , 2. 6  ,0. 07 , ' AND  NON- I RR I GA TED  PLOT S ' ,0. 0 , 2 3 ) 

CALL  DATE  ( NYE AR  ,  0  .2 , 2. 45 ) 

CALL  LEGEND  (3,6.5,2.8) 

CALL  ZERO  (0.5) 
CALL  SYMBL' 


CALL     LINES       ( XPL T , YHA3 , NOA YS , 0. 01 , 2 ) 

CALL     CASH          (XPLT, YLA3,NDAYS, 0.03) 

GJ    TO    300 

E^D 

BLOCK    DATA 

COMMON    /PLOTO 

/ 

XPLT(366) , 

ITAPE 

A                      /PLCTl 

/ 

RAIN(366) , 

XIPR(366) , 

YIRR(366) , 

YNIR(366) 

B                     /PL0T2 

/ 

SNCW(366) 

C                     /PL0T3 

/ 

YHHO( 366) , 

YLHD( 366) 

0                     /PL0T4 

/ 

HUM0{366) 

E                     /PLQT5 

/ 

EVAP(366) 

F                     /PL0T6 

/ 

WINO( 366) 

G                    /PL0T7 

/ 

YMAX( 366) , 

YMIN( 366) 

H                    /PL0T8 

/ 

YH5C(366) , 

YL5C(366) , 

ZH5C(366) , 

ZL5C(366) 

I                     /PL0T9 

/ 

YHAK  366)  , 

YLAl(366) 

J                     /PLOTIO/ 

YHSS(366> , 

YLSS(366) , 

ZHSS(366) , 

ZLSS(366) 

K                    /PLUTll/ 

YHA2( 366) , 

YLA2(366) 

L                    /PLGT12/ 

YHBG(366) , 

YLBG(366) , 

ZHBG(366) , 

ZLBG(366) 

M                      /oi  nri 

4/ 

VWA  -Xl   ^AA>    - 

VI  A  ^r  i<^A« 

226 

0003 


0004 
0005 


0006 


GREAT  BASIN  NATURALIST 


0012 
0013 
0014 


Vol.  35,  No.  2 


COMMON    /LABELS/       AXl(37),     AYI(37) 


DATA  RC2.LC2 


DATA  RC3,LC3 


DATA  RC'«,LC4 


DATA  RC5,LC5 


DATA  RC6,LC6 


DATA  PC7,LC7 


DATA  BX.BY 

DATA  XL.YL 

DATA  AXl.AYl 


DATA  AX2 


DATA  AY2 


BY(4)  , 
Lri(7) 


TAPE    UNIT    NUMBER 
DATA  ITAPE 

DATA  RCl.LCl 


BX(4I , 

RCl(7) 

RC3(ll),     LC3(ll) 

RC5{ll),     LC5(18) 

PC7(12),     LC7(24) 

/5/ 


AX2(73)t  AY2(73), 
XL(4),  YL(4), 

RC2(13)  ,  LC2(  13) , 
RC4(  13) ,  LC4(  13) , 
RC6(8),       LC6(16), 


/•   0  •, 

•    _t  , 

•  25-', 

'    -•  » 

•  50-', 

'    -•  t 

•  75-', 

•   0  ', 

•  -    '  , 

■-  I  '  , 

'-    '  , 

'-  2  ', 

'  -    *  » 

t   3  1/ 

/'   0  ', 

•  25-', 

•  50-', 

'  75-'  , 

'100-', 

•125-', 

•  150-', 

•175-', 

'200-', 

•225-'  , 

•25D-', 

'275-', 

•300-«, 

•   0  ', 

•-  I  '  , 

•-  2  •, 

•-  3  ', 

'-  4  ', 

•-  5  ', 

•-  6  •  , 

'-  7  '  , 

•-  8  '  , 

i_  9  •  , 

'-  10' , 

•-  11', 

•-  12'/ 

/'   0  ', 

•  10-', 

'  20-', 

'  30-' , 

•  40-', 

•  50-', 

•  60-', 

•  70-', 

'  8  0-', 

'  90-'  , 

•  100-'  , 

'   0  ', 

•-  10', 

•-  20«, 

•-  30', 

'-  40'  , 

'-  50', 

'-  60', 

•-  70', 

•-  80», 

'-  90' , 

•-100'/ 

/•  0  •, 

.   2-', 

•   ^,-1  , 

•   6-'  , 

'   8-'  , 

•  10-', 

•  12-', 

•  14-', 

'  16-', 

'  18-', 

'  20-', 

•  22-«, 

•  24-', 

•   0  ', 

•-  2  •  , 

•-  4  '  , 

'-  6  '  , 

'-  8  ', 

•-  10', 

•-  12', 

'-  14', 

•-  16'  , 

•-  18', 

'-  20', 

'-  22^, 

•-  24'/ 

/•   0  ', 

■   2-', 

•   ^_t, 

•   6-'  , 

•   3-'  , 

'  10-', 

•  12-', 

'  14-', 

•  16-', 

'  18-', 

'  20-', 

'   0.', 

•0    ', 

•-  0.', 

•1 

•-  0.' , 

•2 

•-  0.', 

•3    •, 

'-  0.', 

'4    '  , 

'-  0.'  , 

•5 

'-  0.', 

•6    •, 

•-  0.', 

•7    ', 

•-  0.', 

•8    '/ 

/•   0  •, 

•  40-', 

'  8  0-', 

•120-' 

•  160-'  , 

•200-', 

•240-', 

'280-', 

t     1  , 

'0 

•-   2'  , 

*5    •  , 

•-   5*  , 

'0    ', 

•-   7', 

•5    •  , 

'-  rv. 

•0 

•-  12*  , 

•5    ', 

'-  15' , 

•0    •  , 

•-  17', 

•  5    •/ 

/•-30-', 

'-20-', 

•-10-' , 

•   0  ' 

•  10-', 

•  20-', 

•  30-', 

•  40-', 

'  50-', 

•  60-', 

'  70-', 

•  80-', 

._  _2., 

'2 

•  -   -'  , 

'4    ' 

•-     I' , 

•4    ', 

•-   3'. 

'2 

'-   5', 

'0   • 

•-   6'  , 

'8    '  , 

•-   8', 

•6    '  , 

'-  10'  , 

'4    • 

'-  12' , 

•  2    •  , 

•-  14«, 

•0 

•-  15' , 

•8    •  , 

'-  17'  , 

'6    '/ 

/2*0.0, 

2*7.94, 

4*0.0/ 

/3.99,    2*0.0,    -0.04,     2*0.0,    2*1.34/ 

/ 2*7. 94, 3* 7. 2 7, 3*6. 6 1,3* 5. 94, 3*5. 29, 3*4. 61 , 

3*3.9  4,3*3.29,3*2.61,3*1.96,3*1.2  8,3*0.67,2*0.0, 

0.03,2*0.0,0.03,2*0.0,0.03,2*0.0,0.03,2*0.0, 

0.0  3,2*0.  1,0.03,2*0.0,0.33,2*0.0,0.03,2*0.0, 

0.0  3,2*0. 0,0.03,2*0.0,0.0  3,2*0.0,0.03,2*0.0, 

0.03/ 
/2*7.S4, 3*7.60,3*7.2  7,3*6.94 

3*5.9  5, 3*5.62,3*5.2  9,3*4.95 

3*3.94,3*3.62,3*3.29,3*2.95 

3*1.96,3*1.62,3*1.2  8,3*0.98 
/- 0.03, 2*0. 0,-0. 0  3, 2*0. 0,-0. 

-0.03,2*0.0,-0.03,2*0.0,-3. 

-0.03,2*0.0,-0.0  3,2*0.0,-0. 

-0.0  3,2*0.0,-0.0  3,2*0.0,-0. 

-0.03,2*0.0,-0.03,2*0.0,-0. 

-0.03,2*0.0,-0.0  3,2*0.0,-0. 

-C.03/ 


,3*6.61, 

3*6 

28 

1 

,3*4.61, 

3*4 

28 

,3*2.61, 

3*2 

29 

,3*0.67, 

3*0 

34 

'2*0 

3/ 

03 

2*0.0 

,-0 

03 

,2*0 

0, 

03 

2*0.0 

,-0 

03 

2*0 

0, 

03 

2*0.0 

,-0. 

03 

2*0 

0, 

03 

2*0.0 

,-0 

03 

2*0 

0, 

^1> 

2*0.0 

,-3 

33 

2*0 

0, 

0  3 

2*0.0 

,-0. 

03 

2*0.0, 

0002 
0003 


SUBROUTINE    AXES 


HT,X,NXC,DX,N1,M1,Y,NYC,0Y,N2,^'2) 


GRAPH    A    BOX    OF    HEIGHT    HT    WITH    THE    MONTHS    PLOTTED    ALONG    THE    BOTTCM. 
ALSO    LABEL    THE    BOX    WITH    APPAY    X    ON    THE    RIGHT    AND    Y    ON    THE    LEFT. 
UX     IS    THE    DISTANCE     BETWEEN    EACH    LABLE.       Nl    IS    THE    NUMBER    OF    LABELS 
ANO    Ml     IS    THE    STEP    BETWEEN    LABLES.       THE    SAME     IS    TRUE    FOR     ARRAY    Y 
WITH    CY,    N2,     AND    M2. 


DIMENSION    X(N1),  v(N2) 

COMMON    /LABELS/  axi(37), 

A  BX(4J , 

DATA  NCCUNT  /I/ 


AY1(37) , 
BY(4) 


AX2(73),    AY2(73) 


June  1975 


ANDERSEN,  ROPER:    METEOROLOGIC  DATA 


227 


0005 
0006 
3007 
0008 
0009 
0010 
0011 
0012 
0013 
0C14 
3015 


0016 
0C17 
0018 


0019 
0C20 
0021 
0022 
0023 
0024 
3025 
0026 
0027 
3028 
0029 
0J33 
0031 


0032 
0033 
0034 
0035 
0036 
0037 


!F<NCOUNT.NE.l)     GO    TO    10 
CALL     PLOTS     (12.0,0.0,-3) 
CfiLL    PLOT        (2.0,1.0,-3) 
GO    TO     11 
n    CALL    PLOT  (0.0,6.5,-3) 

11  CALL     PLTMRK    (-2.0,0^0) 
NCCJNT    =    NCOUNT    +     1 
IF(;^CCUNT.EC.5)     NCOONT    =    I 
PY(2>    =    HT 
BY(3)     =    HT 
CALL     SYMBL*,     (3.0,0.03,0.07,'        JANUARY 

APRIL  MAY  JUNE  JULY 

BT08ER         NOVEMBER  DEC  EMBER • ,0. 0, 1 3  1 ) 

CALL    LINE     ( AX1,AYI,37,  1) 

CALL     PLOT     (0.0,0-27,-3) 

CALL     SYMBL4    ( 0- 0 ,-0.  1 1 , 0. 0 7 , •  1  15  1 

A    15  I  15  1  15  1  15  1 

B    15  I  15  1  15", 3. 0,123) 

CALL    LINE     ( AX2,AY2,73, 1 ) 

CALL     LINE     (BX,BY,4,1) 

OYY    =    -0.03 

DO    12    I=1,N2,M2 

CALL     SYMBLA     ( 7.9 ,DYY ,0. 07 , Y 1  I )  ,  0.0 , NYC » 

OYY    =    OYY     ♦    DY 

12  CONTINUE 
DXX  =  -0.03 

DO  13  I=l,Nl,Ml 

CALL  SYMBL4  ( -0 . 22 , DXX , 0. 07 , X ( I)  , 0 . 0 , NXC ) 

OXX  =  DXX  *■    DX 

13  CONTINUE 
GU  TO  14 

DRAW  ZERO  LINE  AT  HEIGHT  HT. 

ENTRY  ZERO  (HT) 

BY(2)  =  HT 

BY(3)  =  HT 

CALL  LINE  (BX( 2) ,RY(2) ,2,1) 

14  RETURN 
END 


FEBRUARY 
AUGUST 


MARCH         A 
SEPTEMBER    OC 


SUBROUTINE  DASH  (X.Y.N.S) 


C 
C 
C 

C 

SUBROUTINE    0*SH    WILL    DRAW 

X    =    X    ARRAY 

c 

Y    =    Y    ARRAY 

c 

N    =    NUMBER    CF    POINTS 

c 
c 

S    =    LENGTH    OF    DASHES 

3002 

DIMENSION    X(N),    YCN),    A(2) 

0003 

SS    =    S    ♦    2.0 

0004 

DO    30    1=2, N 

0005 

DX    =    X( I)     -    X( I-l) 

0006 

DY    =    Y( n    -    Y( I-l) 

0007 

H    =    SQRT(DX*DX    «■    OY*DY  ) 

0C08 

DX    =    S    *    DX    /    H 

0009 

DY    =     S    »    DY    /     H 

0010 

A(  I)     =    X(  I-l) 

0011 

B(  I)     =    Y(  I-l) 

0012 

n 

CONTINUE 

0013 

IF(H.LE.S)     GO    TO    20 

0014 

A(2)     =    A(  1)     +    DX 

0015 

8(2)     =    B( I)    ♦    DY 

0016 

CALL    LINE     (A,B,2,1) 

0317 

IF(H.LE.SS)    GO    TO    30 

0018 

A(  I)     =    A{2)    +    DX 

0019 

B(  I)     =    B(2)    ♦    DY 

0020 

H    =    H    -    SS 

0021 

GO    TO     10 

3322 

20 

A(2)     =     X( I) 

0023 

6(2)     =     Y( I) 

0024 

CALL     LINE     (A,B,2,1) 

3025 

30 

CONTINUE 

228 

0026 


GREAT  BASIN  NATURALIST 


Vol.  35,  No.  2 


0027 

0028 
0029 
0030 
0031 
0032 
0033 

J035 

0036 
0037 
0038 
C039 
DOAO 
0041 
0042 
0  343 
0044 
0045 


SUBROUTINE  LINES  WILL  MAKE  THE  LINE  DARKER  AND  WIDER  BY  DRAWING 
N  NUMBER  OF  LINES 

X  =  X  ARRAY 

Y  =  Y  ARRAY 

N  =  NUMBER  OF  POINTS 

D  =  DISTANCE  BETWEEN  LINES 

M  =  NUMBER  OF  LINES 

ENTRY  LINES  (X,Y,N,0,M) 

DY  =0.0 

DO  40  1=1, M 

CALL  LINE  (X.Y.N.I) 

CALL  PLOT  (0.0,0,-3) 

OY  =  OY  -  D 
40  CONTINUE 

CALL     PLOT     (0.0,DY,-3) 

GO    TO    99 

ENTRY    BAR     (X,Y,N) 

SS    =    0.0 

CALL    PLOT     (X(1),SS,3) 

DO    50     1=2, N 

CALL     PLOT     (X(  I-l),Y(  I) ,1) 

CALL     PLOT     (X( I », Y( I ) ,1) 
5J    CONTINUE 

CALL  PLOT  (X(NI),SS,I) 
99  RETURN 

END 


0001 


SUBROUTINF  DATE  (NYR.X.Y) 

GRAPH  DATE  SPECIFIED  BY  NYR  AT  POINT  (X,Y) 


0002 
0003 
0004 
0005 
0006 
3007 
J008 
0C09 

oon 

0311 
0012 
0013 
0014 


0016 
>3017 
0018 
0319 
JJ2T 
0021 
0  322 
0023 
0024 
3026 
J  326 
0027 
0028 


CALL     SYMBL4  ( X , Y , 0. C 7 , ' PRC VD ,     UT AH , • , 0. 0 , 12 ) 

N    =    NYR-1969 

GOTO    (1,2,3,4,5),    N 

1  CALL    SYMBL4  ( X+0. 7 6, Y, 0. 07 , ' 1970 • , 0. 0 ,4) 
RETURN 

2  CALL     SYMeL4  (  X  ♦•O  .  7  8  ,  Y ,  C.  C  7  ,  •  1 9  7  1  •  ,  0.  0  , 4  ) 
RETURN 

3  CALL     SYMBL4  ( X+0. 78 , Y, 0. 0 7 , • 1972 • , 0. 0 ,4) 
RFTUPN 

4  CALL     SYMBL4  ( X  + J . 78  ,  Y , 3. 0 7 , • 1 973 '  , 0 . D  ,  4) 
RETURN 

5  CALL     SYMBL4  (  X*  3  .  7  8  ,  Y  ,  0.  0  7  ,  '  I  9  74  '  ,  0  .  ">  ,  4  ) 
RETURN 


ENTRY  LEGEND  (N,X,Y) 

GRAPHS    ONE    OF    FH'JR     LEGENDS     SPECIFIED    BY    N    AT    POINT     (X.YI 

G3TQ     (  10,  1  I,  12,  13)  ,N 

10  CALL     SYMPL4  (  X  ,  Y  ,  0  .  07,  '  f- A  X  I  MUM  •  ,  0  .  0  ,  7  ) 
CALL     SYMBL4  (  X  ,  Y-0.  1 ,  0.  07  ,  •  M  I  N  I  MU^' •  ,  n  .  3  ,  7  ) 
RETURN 

11  CALL     SY'^eL4  (  X  ,  Y  ,0  .  0  7  ,  •  MA  x     NCN- I  RR  •  ,  0  .  0  ,  1  1 ) 
CALL     SYMBL4  ( X , Y-0. 1 , 0. 07 , • M  IN    NON- I RP •  ,0 . 0  , 1  I  ) 
CALL    SYMBL4  ( X , Y-0 . 2 2, 0. 07 , • f A  X     I  PR  I GA TED •  ,  0. 0 , 1 3  ) 
CALL     SYMBL4  ( X , Y-0. 32, 0.0 7 , ' M I N    I  PR  I  GATED ' , 0. 0, 13  ) 
RETURN 

12  CALL     SYMBL4  (  X  ,  Y  ,0  .  3  7,  •  ME  AN    NOr'- I  RR  IG  A  TED  •  ,  0.  0  ,  1  8  ) 
CALL     SYMBL4  ( X , Y-0. 1 ,0 . 37 , • Mf AN    I RP IG ATED* , J. 3  ,  L 4  ) 

13  RETURN 
END 


0001 


SUBROUTINE     SCALE     (  X , N, S , YM AX  ,  YM I N , I D) 

S'"ALE     DATE     IN    A^KAY    X     DIMENSICNFO    BY    N    AND    RETURN    SCALEO    OMA     IN 
X.        S     IS    THE    MAXIMUM    HEIGHT     OF     SCALED     DATA,     YMAX     HAS     MAXIVUM 
ALLOWED    DATA    AND    YMN     THE     KIMMUM.        SHOULD     DATA     EXC>=FD    THESE 
BOUNDS,    THE     ID    NUMBER    WILL     BE    PRINTED    ALONG    WITH    THE    DATA. 


June  1975  andersen,  roper:  meteorologic  data  229 

O0T2  DIMENSION  X  <  f' ) 

0003  SS  =  S  /  (  YMAX-YMNJ 

TJ34  on  10  1=1, N 

00^5  IF(X(  I  )  .LT.VMINJ  WRITE(6,l-jn  ID,  X(I),  YMIN 

0006  IF(X(  n.GT.YMAX)  VIRITE(&,100)  ID,  X(T),  YMAX 

0007  X(  I )  =  SS  »  ( X(  I)  -  YMIN) 
0CC8            10  CCNTINUE 

3:39  RFTURN 

0010  no    FORMAT!'  CALL     •  ,  1  2  .  5  X  ,  F  I  0.  2  ,  •     EXCFEOED    '.Fg.D 

0011  lOI    FORMAT(«  CALL     • ,  I  2 , 5 X, F  10 . 2 , •     WAS    BELCW     '.FQ.l) 
3312                                        END 

OCjI  SJ'ShLuTI.^L     .M-AuC     (  I  Y£A^  ,i\CAYS) 

C 

r,  KcAu    Ifi'z    Y£Ak'b    uiTA    SPECIFIEU    BY     lYEAR.       THE    FIRST    CARD     IS     i     CATA 

C  HEAL.Lr,    LUi\,TA  INI  M:,     THE     Y£AK,     NUMBER     OF     DAYS     IN     YEAR    AND    LAST 

C  kECukDEG    WlNb    VALUE    CE     PRtVlCUS    Yi^AR.       RETURim    NJMOlR    UF    DAYS    IN 

C  NO  AYS. 

C 
CrC2  CLM^.-JN    /PLCK     /        XPLTOto),     ITAPt 

A  /P^GTl    /        TPRE(3oo),     XlRROfab),     YlRRCjOb),     YNIP(36d) 

P  /PLCT2    /        Si\CV(366) 

C  /PLLT3     /        Hho^(3cfc),     LHUN(3o6) 

D  /PLCT4    /       THPS<'3o6) 

E  /PLLT5     /        PEVP(j.6fcJ 

F  /PLCTo    /        wINC(j)o6) 

G  /PLCT7    /       YMAX(366),     YMlN(3b6) 

H         /PLJTo  /   YH5C(JC6),  YL5C(j6t),  LhbCiibti,     ^LbC(36b) 
I         /PLUT9  /   YHAl(366),  YLA1(366) 

J        /PLCTIO/   YpSS(366),  YLSS(366),  ZHSS136o),  ZLSS(36o) 
K         /PLuTU/   YHi2(:>ob),  YLA^(366) 

L         /PLJTi2/   YhfcG(366),  YLBG(366),  ZhBG(36o),  ZLB3(366) 
M         /PLOUi/   YHA3<366),  YLA3(366) 
0C03  CGMHCN  /LABELS/   DUNY(22b),  XL(A),    YL(4J 

CCO^  REAL     CbUl)    /•   0  ',  •    -',  •  10-',  •    -',  •  2  3-',  '    -', 

A  '  30-' ,  •    -',  •  ^0-',  •    -•  ,  '  50  •/ 

Jjyj  REAL  FHUM,  LhLf'',  M  k  Vv ,  M  C  V. ,  IRnW,  IROW  ,OLDw 

OCCo  AVE(X,Y)  =  (X+YJ/^.C 

C 
->j7  1  PEAu(  ITaPl  ,2J  J,LND=?9>7  )  I  YF  ,  NDA  YS  ,  CLUV^ 

Ci-Oe  200  F3RKATi21A,F^.O) 

::39  ifciyeak.ew.iyr)  gu  to  3 

OCIO  00    ^     I=l,NCAYS 

DCll  2    KcAOd  TAPE  ,200, END  =  SS9) 

CC12  Gu    TO     1 

C 
C 

0C13  3    DO    ^     I=i,NoAYS 

3314  REAC(  ITAPc  ,201  ,c,mD=';99)    RAIN,     SNOkJ,     SNCVCI),     IK^«,     IRDW,    NlViU, 

A  NIDU,     HHUK(I),     LHUM(I),     THRS(I),    PtVP(I), 

B  LLCV,     UINC(I),     YMAX(I),     YMINCH, 

C  YH5C(!),     YL5C(I),     YHSS(I),    YLSS(I),     YHBGd), 

D  YLBG(I),     ZH5C(I),     ZL5C(I),    ZHSS(I),     ZLSS(I), 

E  ZhBG(I),     ZLBGd) 

)  n-J  2  31    FQRNAT(bX,F3.2,4F3.  1,2X,2F3.1,1X,2F3.C,F2.0,F3.1,F2.0,F3.0,/, 

A  5X,^F3.0,bX, 12Fi.O) 

CClo  XPLT(I)     =     I 

.:317  TPRE(n     =     RAIN    •»•     SNLVs*C.l 

0C13  YIRR(I)=-i.J 

0319  YMR<  I  )    =    -1.0 

0C20  IFINI wW.NE . J.O J     YNIR(I)     =     < N IWW-N I DW ) * IJO . j/NIJW 

C021  IFdRww.NE.O.O)     YIRR(I)     =     d  RWW- I  RDW  )  *  I  JO.  J/IRCW 

::22  lF(uLJW.GT.ksINOd  )  )     CLClv    =    OLDW    -     ICOO.O 

0C23  HOLD     =    WIND(I) 

0C24  wIND(I)     =     (wINi.(  n-CLDW)     *     l.6093«-0.05 

3025  DLOW    =    HOLD 

0026  YHAKIJ    =    AVE(  YHijC<  I  ),YLbCd)) 

3327  YLAKIJ    =    A  VE  (  ZH5C  (  I  )  ,  ZL5C  (  I  )  > 

^C2d  YHi2d)     =     iVt  (  YHSS(  I),  YLSSI  n  ) 

^C^9  YLi^«I)     =     AVE(ZHoS(  n.ZLSSd)  J 

3)jJ  YhAj)(I)     =     AVt<  Yhoo(  I  ),  YL'3Gd  )  I 

CC31  YlAjJI)     =    AVE<  ZHB.:,(  I  ),ZL&G(  n  ) 


230 


GREAT  BASIN  NATURALIST 


Vol.  35,  No.  2 


0C3^ 

4 

COMIi\Ub 

JCiJ 

Y3AVS     =    MJ>S 

0CJ4 

CALL     SCfiLL     tXPLl  ,\[./^YS  ,7.94,YCiYS,        I.'.,     U 

r-Tss 

-itTLKN 

J-J6 

9  99 

^»o  !  Tfc  (6,5  )j) 

0C^7 

500 

FCkMAK  •-• »**'»»-*i''(  LRHCR     IN     INPUT    DATA    fhClA     F  AP  c:     *  v**  *•  ,  /  /  ) 

::3o 

C 

srjp 

CC39 

c 

c.n;TKY    must 

d:;^j 

CALL     PLOT          (u. 0,1. 2, -3) 

:j4i 

N    =    0 

0C42 

DJ    ZL     1=12  1,^06 

J  J-ifi 

iFiYikh (I)  .LT. J..)   g:  tc  z: 

cc^^ 

N     =    K    +     1 

CC45 

YIFR(NJ     =     Y1K5( I ) 

CC46 

XIRk (N)     =     XPLT ( I ) 

0C47 

20 

CONTINUE 

0C4b 

CALL     SCALE     (YIRF  ,,>j,  l.^d,50.0,C.0,29) 

CC49 

CALL     LINt     (XlKR,YlKr,,\,l) 

CC5C 

K     =    C 

J  Obi 

OC    JJ     I=U1,jOo 

0C52 

1  F(  YMR<1  >  .LT.C.  0)     GC     TO    30 

j:i)3 

N    =    N    +     1 

jC54 

<IRtN(N)     =     XPLTd) 

0Cb5 

Y  N I K  <  N )    =   Y  ^a  R  (  I  ) 

3':  36 

3  J 

CCNTINUE 

J..b7 

CALL     SCALE     I  YN  I  k  ,  ,m  ,  1  .^  t  ,  5  J.  0  ,  0.  J  ,  3.T  ) 

CC5b 

CALL     CASH     (  XlRK,  YM  h  ,N,C.04) 

OJbS 

CALL     PLCT           (^.6  1  ,./.:, -j) 

J06  J 

CALL     LINE           (XL.YL,',  ,1) 

CC61 

OY    =    -0.0  J 

0-)6^ 

DO  n   1  =  1,  ii 

ccej 

CALL     SYMbL4     <  -  J.  <i2  ,  C  Y  ,  J  .  J  7  ,  CR  (  I  )  ,  0  .  0,  4  J 

OCot 

DY    =    DY    +    0.13 

CC63 

10 

CCNTINUt 

C066 

CALL     SY:^bL4     (-0.22  ,  C.45,G.C7,  'PERCENT  •  ,',J.^  ,7  J 

C:o7 

CALL     5YMBL4     ( 3. 2 , 1 . 5 i  ,  C, C 7  ,  '  Sul L    MblSTURE     (  IRR I wATE J )  •  , 0. 0 

0C63 

CALL     SY^1oL4     (  j.2,1.6j,C.:7, 'SOIL    MOISTURE     (  NJN- 1  RR  I  GA  TE  0  )  ' 

CC6  9 

CALL     PLOT     (-^.ol,-1.2,-3J 

•wC7J 

RETURN 

CC71 

ENC 

Literature  Cited 

Andersen,  F.  L.,  P.  D.  Wright,  and  J.  C.  Fox. 
1974.  A  comparison  of  meteorologic  mea- 
surements from  irrigated  and  non-irrigated 
plots.  Provo,  Utah.  1970-1972.  Brigham 
Young   Univ.   Sci.   Bull.,   Biol,   Ser.    19:    1-37. 

Fo.\'.  J.  C,  F.  L.  Andersen,  and  K.  H.  Hoopes. 
1970.  A  survey  of  the  helminth  parasites 
of  cattle  and  sheep  in  Utah  Vallcv.  Great 
Basin  Nat.  30:131-145. 

FuRMAN,  D.  p.  1944.  Effects  of  environment 
upon  the  free-living  stages  of  Ostertagia  cir- 
cumcincta  (Stadelmann)  Trichostrongylidae: 
II.  Field  Experiments.  Am.  J.  Vet.  Res.  5: 
147-153. 

HoNEss,  R.  F.,  AND  R.  C.  Bergstrom.  1966. 
Trichostrongylosis    of    cattle    in    Wyoming. 


Science  Monograph  2.  Agr.  E.x.  Sta.,  Univ. 
Wyoming,  Laramie. 

Mitchell,  J.  K..  and  F.  L.  Andersen.  1969. 
A  computer  program  for  meteorologic  data 
reduction.  Transactions  of  the  111.  State  Acad. 
Sri.    62:    15-28. 

Rainey,  M.  B.,  and  a.  D.  Hess.  1967.  Public 
health  pi-oblems  related  to  irrigation.  Pages 
1070-1081  in  R.  M.  Hagan,  H.  R.  Haise, 
and  T.  W.  Edminster.  eds.  Iirigation  of  ag- 
ricultural lands.  American  Society  of  Agron- 
omy. Madison.  Wise. 

Would  He.m.tii  Organization.  1950.  Joint 
study  group  on  bilharziasis  in  Africa.  Rep. 
1st.  "Sess.  Tech.  Rep.  17:16. 

Wright,  P.  D.,  and  F.  L.  Andersen.  1972. 
Parasitic  helminths  of  sheep  and  cattle  in 
Central   Utah.   J.   Parasitol.   58:959. 


CLARENCE  COTTAM,  1899-1974 

A  Distinguished  Alumnus  of 
Brigham  Young  University 

Vasco  M.   Tanner^ 


Dr.  Clarence  Cottani  was  one  of  the  na- 
tion's most  outstanding  and  })roductive 
biologists  and  conservationists.  While  this 
is  an  achievement  in  itself,  he  further  dis- 
tinguished himself  in  the  field  of  adminis- 
tration during  his  service  as  assistant  di- 
rector of  the  U.S.  Fish  and  Wildlife  Ser- 
vice between  1945  and  1954,  and  as  direc- 
tor, for  almost  twenty  years,  of  the  Rob 
and  Bessie  Welder  Wildlife  Refuge  at 
Sinton,  Texas. 

Clarence  Cottam  was  born  in  Utah's 
Dixieland  at  Saint  George  on  January  1, 
1899.  His  father,  Thomas  P.  Cottam,  and 
his  mother,  Emmaline  Jarvis  Cottam,  were 
prominent  St.  George  citizens,  his  father 
having  served  as  mayor  of  St.  George  and 
counselor  with  the  LDS  stake  president, 
Edward  H.  Snow.  He  was  also  a  success- 
ful farmer  and  mason.  Clarence  spent  his 
boyhood  days  working  on  the  farm  and 
enjoying  the  plants  and  animals  of  the 
Virgin  River  and  Santa  Clara  Creek.  The 
St.  George  area  is  the  sole  region  of  Utah 
in  which  the  plant  and  animal  species  of 
the  Lower  Sonoran  Zone  are  found.  Clar- 
ence became  acquainted  early  with  the 
vermilion  flycatcher,  gambel  quail,  road- 
runner,  phainopepla,  and  western  mock- 
ingbird and  as  a  high  school  student  evi- 
denced an  interest  in  and  general  acquain- 
tance with  the  birds  of  his  homeland.  The 
following  is  extracted  from  one  of  his 
papers  written  in  1970: 

My  work  in  the  field  of  conservation, 
teaching,  and  management  of  re- 
sources certainly  had  its  start  in  the 
biological  training  I  gleaned  under  the 
able  leadership  of  Dr.  Tanner  at  Dixi'e 
and  BYU. 

In  my  first  course  in  high  school  bi- 
ology, Dr.  Tanner  asked  me  what  species 
of  hummingbirds  I  had  seen.  To  me  they 
were  all  hummers,  so  I  answered,  "Little 
hummers;  and  bigger  ones  and  some  had 
white  throats  and  some  with  iridescent 
black  and  brilliant  red  throats."  He 
answered  kindly  but  challengingly  asked, 

'Department    of    Zoology,    Brigham    Young    University,    Pr^ 
'Great  Basin  Nat.    30:201.      1970. 


"Wliat  species  are  here?"  and  he  added 
"It  will  be  a  lot  of  fun  to  find  out." 
Then  he  inquired,  "what  kind  of  flowers 
do  they  go  to?"  I  could  only  reply  that 
they  came  to  flowers  of  different  colors 
and  I  had  seen  them  in  flowers  of  dif- 
ferent shades  of  red,  blue  and  white.  He 
challenged  me  further  by  asking  "what 
do  these  tiny  birds  feed  on?"  I  could 
only  suggest  "something  inside  of  the 
flowers."  He  then  assigned  me  the  task 
of  finding  out  what  I  could  about  hum- 
mingbirds. There  were  few  books  on 
birds  at  that  time  in  St.  George  and  still 
fewer  of  those  dealing  with  these  diminu- 
tive but  attractive  creatures,  so  he  remind- 
ed me  that  the  best  place  to  learn  of 
them  was  out  of  nature,  he  appropriately 
added  that  nature's  books  were  always 
open   at   the   appropriate   time. 

I  doubt  that  my  esteemed  friend  and 
teacher  has  the  faintest  recollection  of  this 
little  incident,  and  I  am  still  more  sure 
that  any  knowledge  I  imparted  on  this 
assignment  was  elementary  indeed.  Still, 
this  challenging  excursion  into  nature 
.  left  an  indelible  impression  on  my  mind. 
It  has  over  the  years  made  me  ask  many 
questions  about  nature's  ways  and  her 
varied  progeny.  How-  do  they  survive  and 
compete?  What  good  or  harm  do  they 
do?  What  relationship  do  they  have  to 
their  environment  and  to  other  species? 
Why  were  they  where  I  found  them? 
How  could  they  be  increased,  decreased 
or  controlled?  What  were  the  basic  fac- 
tors of  their  population  dynamics? 

In  looking  back  on  these  rich  early 
experiences  I  feel  that  the  most  valuable 
training  I  received  was  on  our  summer 
field  trip  when  we  were  collecting  insects, 
birds,  rodents,  and  plants  for  the  Uni- 
versity. On  these  trips  the  graduate  stu- 
dents were  collecting  and  studying  spe- 
cific groups  of  organisms  or  problems  for 
their  respective  theses.  I  was  studying 
and   collecting  birds. - 

After  (ompleting  his  school  work  at 
Dixie  College,  Clarence  was  called  to  .spend 
two  vears  in  the  central  states  as  a  mis- 
sionary for  the  LDS  Church.  In  1922  he 
accepted  the  principalship  of  the  schools 
at  Alamo,  Nevada,  and  in  1925  he  ma- 
triculated at  Brigham  Young  University 
and   was   appointed    an    instructor   in   bi- 


231 


232 


GREAT  BASIN  NATURALIST 


Vol.  35,  No.  2 


Fig.   1.  Clarence  Cottam.   1899-1974. 

ology.  For  the  next  four  years  ho  was  ac- 
tive in  research  and  field  work.  He  grad- 
uated with  a  B.S.  degree  in  zoology  and 
entomology  in  1926  and  a  year  later  was 
awarded  a  Master  of  Science  degree.  His 
thesis  dealt  with  the  birds  of  Utah. 

In  1929  Clarence  received  an  appoint- 
ment as  junior  biologist  in  the  U.S.  I3ureau 
of  Biological  Survey  in  Washington,  D.C. 
During  his  first  few  years  with  the 
Bureau  he  devoted  himself  to  an  investi- 
gation of  food  habits  research  but  also 
found  time  to  continue  graduate  studies 
at  George  Washington  University,  from 
which  he  received  a  Ph.D.  degree  in  193fi. 
This  early  study  resulted  in  his  most 
prized  contribution,  "Food  habits  of  North 
American  diving  ducks"  (1936).  At  this 
time  Clarence  also  became  clo.sely  asso- 
ciated with  leaders  and  authorities  on 
wildlife  management  and  conservation 
such  as  Ira  N.  Gabrielson,  Aldo  Leopold, 
and  .T.  N.  (Ding)  Darling.  Embracing 
their  point  of  view,  he  became  one  of 
this  country's  most  dynamic  and  sought 
after  champions  of  wildlife  conservation. 


Clarence  was  an  ambitious,  friendly 
individual.  He  made  friends  with  the  of- 
ficials of  organizations  and  societies 
throughout  the  United  States,  Canada,  and 
some  foreign  countries  that  were  con- 
cerned with  the  flora  and  fauna  of  the 
nation.  He  was  a  fluent  speaker,  well  in- 
formed on  the  management  of  wildlife. 
Few,  if  any,  conservationists  of  the  Clar- 
ence Cottam  era  were  his  peers  in  promul- 
gating conservation  measures  in  behalf  of 
the  wildlife  of  our  country.  He  vigorously 
opposed  the  indiscriminate  use  of  certain 
poisons  for  the  control  of  insects,  birds, 
fish,  and  mammals  and  spoke  out  against 
destroying  food  and  nesting  habitats  of 
native  fish  and  birds  through  the  drainage 
of  wet  lands  and  dredging  and  contami- 
nation of  the  nation's  waterways. 

During  Dr.  Cottam's  25-year  affiliation 
with  the  U.S.  Biological  Survey,  later  the 
U.S.  Fish  and  Wildlife  Service,  many  or- 
ganizations sought  after  his  services  as  a 
consultant  or  officer.  He  was  president  of 
the  Wildlife  Society  (1949-1950),  Texas 
Ornithological  Society  (1957-1959),  and 
National  Parks  Association  (1962-1963); 
trustee  of  the  J.  N.  (Ding)  Darling  Foun- 
dation and  the  Rachel  Carson  Trust.  He 
was  an  active  member  of  the  following 
organizations:  American  Ornithological 
Union,  Cooper  Ornithological  Society,  Wil- 
son Ornithological  Society,  Forestry  Asso- 
ciation, Outdoor  Writers  Association, 
Wildlife  Management  Institute,  Izaak 
Walton  League,  Cosmos  Club,  Soil  Con- 
servation Society,  Friends  of  the  Land, 
Wildlife  Society,  Utah  Academy  of 
Sciences,  Arts,  and  Letters,  American  As- 
sociation for  the  Advancement  of  Science, 
Ecological  Society  of  America,  Limnolog- 
ical  Society,  American  Society  of  Mam- 
malogists,  American  Society  of  Range 
Management,  National  Wildlife  Feder- 
ation, National  Audubon  Society,  Sigma 
Xi,  Sierra  Club,  American  Institute  of 
Biological  Sciences,  and  the  Southwestern 
Association  of  Naturalists. 

Ill  (-('cognition  of  his  leadership  and 
(ouiisel  as  an  imtiring  government  of- 
fi(  iai  and  stimulating  member  of  many 
societies,  ho  was  the  recipient  of  the  fol- 
lowing awards:  the  Leopold  Medal  from 
the  Wildlife  Society  (1955);  Utah  State 
Uni\ersity  Distinguished  Service  Award 
(1957);  IMational  Audubon  Society  Dis- 
tinguished Service  Medal  (1961);  Con- 
servation Service  Citation  of  the  National 


June  1975 


TANNER:    CLARENCE   COTTAM 


233 


Wildlife  Federation  (1964);  Distinguished 
Service  Award,  BYU  (1964);  Conserva 
tion  Senice  Award  of  the  Department  of 
the  Interior  (1965);  Distinguished  Ser- 
vice Award  of  the  Texas  Chapter  of  the 
Wildlife  Society  (1971);  "Eminent  Dis- 
tinction" status  in  the  National  Register  of 
Prominent  Americans  and  International 
Notables  (1971);  James  A.  Talmage 
Scientific  Achievement  Award,  BYU 
(1971);  Distinguished  Service  Award  of 
the  Texas  Ornithological  Societ}'    (1972). 

In  1954  Clarence  resigned  his  govern- 
ment post  to  accept  the  deanship  of  the 
College  of  Biological  and  Agricultural 
Science  at  Brigham  Young  University.  As 
dean,  he  had  begun  instituting  a  number 
of  significant  changes  in  the  college  when 
he  received  an  offer  to  become  the  direc- 
tor of  the  Welder  Wildlife  Foundation  at 
Sinton,  Texas — a  new  and  promising  pro- 
ject. Because  this  program  provided  many 
possibilities  for  research  and  the  means  of 
demonstrating  what  must  be  done  to  de- 
velop conservation  measures  for  the  per- 
petuity of  many  of  the  country's  plant 
and  animal  species.  Dr.  Cottam  accepted 
the  directorship,  assuming  his  duties  as 
director  of  the  Rob  and  Bessie  Welder 
Wildlife  Foundation  in   1955. 

For  almost  20  years  Clarence  directed 
the  development  of  the  Welder  Wildlife 
Refuge  on  the  Aransas  River  in  San  Pa- 
tricio County,  near  Sinton,  Texas.  The 
refuge,  located  in  a  transition  zone  be- 
tween coastal  prairie  and  the  Rio  Grande 
plain,  is  a  strip  of  rich  alluvial  land, 
seven  miles  long  and  about  two  miles 
wide.  For  many  years  it  was  part. of  a 
cattle  ranch,  and  today  as  a  part  of  the 
biota  of  the  refuge  there  are  more  than 
500  head  of  beef  cattle. 

The  refuge  is  favorably  located  to  serve 
as  a  conservation  research  area.  Dr.  Cot- 
tam observed  that  "this  is  one  of  the  rich- 
est areas  for  flora  and  fauna  in  Amer- 
ica."^ In  this  wilderness  he  has  identified 
more  than  1,300  species  of  plants,  in- 
cluding 200  species  of  grass  and  more 
than  400  species  of  birds.  Through  the 
efforts  of  Dr.  Cottam  and  his  staff  the  re- 
fuge has  been  developed  into  one  of  the 
most  renowned  wildlife  research  and  ed- 
ucational laboratories  in  the  United  States. 
Nearly  150  students  from  many  univer- 
sities have  obtained  graduate  degrees 
under  leadership  at  the  foundation. 

^Deseret  News,  16  June  1973,  Church  Section,  p.  7. 


Dr.  C>)ttam  was  a  bibliophile;  he  built 
up  a  complete  set  of  ornithological  jour- 
nals and  textbooks  on  American  birds  for 
the  use  of  students  at  the  refuge.  He  was 
also  concerned  with  the  migration,  breed- 
ing, and  feeding  of  game  birds  and  cam- 
paigned for  the  j)rotection  of  rare  and 
vanishing  species,  such  as  hawks,  owls, 
eagles,  whooping  cranes,  and  brown  pel- 
icans. He  constantly  emphasized  the  ne- 
cessity of  ha\ing  a  broad,  well-grounded 
program,  based  upon  verifiable  facts,  when 
dealing  with  conservation  problems. 

Clarence  also  found  time  to  coauthor, 
with  Dr.  Angus  M.  Woodbury  and  ,Tohn 
Sudgen,  a  manuscrijit  dealing  with  the 
birds  of  Utah.  The  untimoh',  accidental 
death  of  Dr.  Woodbury,  the  senior  author, 
delayed  the  publication  of  this  important 
treatise.  Through  the  efforts  of  Dr.  Cot- 
tam this  manuscript  was  turned  over  to 
Dr.  C.  Lynn  Hayward  of  the  Department 
of  Zoology  at  BYU  to  review,  edit,  and 
publish. 

Clarence  had  the  help  and  companion- 
ship of  a  gracious,  artistic  helpmate  and 
wife,  Margery  Brown  Cottam,  whom  he 
married  on  May  20,  1920,  while  they  were 
both  teaching  at  Alamo,  Nevada.  The}^ 
were  the  parents  of  four  girls:  Glenna, 
Mrs.  Ivan  L.  Sanderson  of  San  Francisco, 
California;  Margery,  Mrs.  Grant  Osborne, 
Amherst,  Massachusetts;  Josephine,  Mrs. 
Douglas  S.  Day,  Salt  Lake  City,  Utah; 
and  Caroline,  Mrs.  Dwayne  Stevenson, 
McClaine,  Virginia.  They  had  23  grand- 
children and  4  great  grandchildren.  Mar- 
gery was  an  active  member  of  the  LDS 
Church,  serving  as  stake  YWMIA  pres- 
ident and  stake  Relief  Society  president. 
She  died  February  28,  1975,  and  was 
buried  beside  Clarence  at  Orem,  Utah. 

Always  religious  and  devoted  to  the  LDS 
Church,  Dr.  Cottam  was  a  pillar  of 
strength  in  the  LDS  Corpus  Christi 
(Texas)  Stake.  He  served  as  first  coun- 
selor in  the  San  Antonio  Stake,  after  which 
he  served  for  nine  years  as  president  of 
the  Corpus  Christi  Stake.  At  the  time  of 
his  death,  March  30,  1974,  he  was  pa- 
triarch of  the  Corpus  Christi  Stake. 

He  was  the  essence  of  tolerance  in  his 
dealings  with  his  fello\MTien.  As  an  edu- 
cator, administrator,  researcher,  and  inter- 
mediary between  scientific  groups  and  the 
the  sports  public.  Dr.  Cottam  was  most 
successful. 


234 


GREAT  BASIN  NATURALIST 


Vol.  35,  No.  2 


Dr.  Clarence  Cottam's  name  is  indelibly 
inscribed  on  the  roster  of  distinguished 
alumni  of  Brigham  Young  University. 
For  the  past  50  years  I  have  enjoyed 
\vatching  him  become  one  of  the  most 
honored  biologists  of  the  nation. 

The  long  list  of  his  publications  which 
follows  reveals  his  wide  experience  and 
insight  in  providing  solutions  to  man's 
mistakes  in  dealing  with  his  animate  en- 
vironment. 


Published  Biographical  Reports 
of  Clarence  Cottam 

Anonymous.  1974.  Clarence  Cottam:  Nation- 
ally acclaimed  conservationist.  National  Parks 
Conservation   I^lagazine   28:26. 

Bolen,  Eric  G.  1975.  In  memoriam:  Clarence 
Cottam.   Auk  92:118-125. 


Bibliography  of  Clarence  Cottam 

1928-1975 

For  the  preparation  of  the  following 
list  of  publications  by  Dr.  Clarence  Cot- 
tam, I  am  obligated  to  Mr.  Douglas  S. 
Day,    Dr.    Cottam's    son-in-law. 

1928. 

1928. 
1928. 


1929. 
1929. 
1929. 
1929. 

1929. 
1930. 

1930. 
1931. 


1931. 
1931. 
1932. 


Christmas  bird  census,  Provo,  Utah.  1927. 
Bird  Lore  30(1): 65. 

Killdeer  swimming  on  Green  River.  Utah. 
Auk     45  (2):  207-208. 

White  pelicans  and  great  blue  herons  win- 
tering in  northern  Utah.  Condor  30:160. 
April. 

Christmas  bird  census,  Provo.  Utah.      1928. 
Bird  Lore  31(1):  61-62. 
A   shower  of  grebes.    Condor  31  (2):  80-81. 
March. 

The  status  of  the  ring-necked  pheasant  in 
Utah.    Condor  31  (3):117-123.   May. 
The    fecunditv   of   the   English   sparrow    in 
northern   Utah.     Wilson    Bull.    41:193-194. 
September. 

A  studv  of  the  water  birds  of  Utah  Countv. 
Utah.  Proc.  LTtah  Acad.  Sci.  6:8-11.  July. 
Food  habits  of  the  shoal-water  and  diving 
ducks  of  Florida.  Florida  Woods  and  Wa- 
ters 1(2):  37-38.  Summer. 
A  friendly  humming  bird.  Bird  Lore  32 
(5):352.  September-October. 
Montlil}"  and  yearly  percentages  of  food, 
page  124;  Animal  food,  pages  158-159;  In- 
dex, pages  547-559  in  H.  L.  Stoddard,  The 
bobwhite  quail:  its  habits,  preservation, 
and  increase.  Charles  Scribner's  Sons,  New 
York    City. 

Some  new  and  uncommon  bird  records  for 
South  Dakota.  Wilson  Bull.  43  (4): 311. 
Birds  and  motors  cars  in  South  Dakota. 
Wilson  Bull.  43(4):313-334. 
(with  F.  M.  Uhler,  Leon  Kelso,  and  E.  R. 
Kalmbach).  Christmas  bird  census.  Port 
Tobacco,  Maryland.  1931.  Bird  Lore  34 
(1):49. 


1932.  (with  Leon  Kelso,  and  W.  H.  Ball).  The 
Louisiana  heron  in  Washington.  D.C., 
region.   Proc.  Biol.  Soc.  Washington  45:207. 

1932.  Nocturnal  habits  of  the  chimnev  swift. 
Auk   49(4):  479-481.    October. 

1933.  (with  F.  M.  Uhler.  A.  L.  Nelson,  and  Leon 
Kelso).  Christmas  bird  census.  Port  To- 
bacco, Maryland.  1932.  Bird  Lore  35 
(1):30. 

1933.  (with  Phoebe  Knappen).  Oil  gland  usually 
tufted  in  Hj'dranassa  tricolor  ruficollis. 
Auk  50(l):94-95. 

1933.  (with  Leon  Kelso).  An  incubating  wood- 
cock.   Auk    50(2):  170-173;    4    pi. 

1933.  Night  migration  of  eastern  chipping  spar- 
rows.   Bird   Banding  4(l):54-55. 

1933.  Feeding  habits  of  the  lesser  scaup  duck. 
Condor     35 (3): 1184 19. 

1933.  Winter  records  for  the  coastal  region  of 
North   Carolina.   Auk   50(2) :  231 -232. 

1933.     A  pot  cock  pheasant.  Bird  Lore  35(3):  148. 

1933.  Disappearance  of  eelgrass  along  the  At- 
lantic coast.  Plant  Disease  Reporter  17(6): 
46-53.  (Mimeographed,  Bureau  of  Plant 
Industry.) 

1933.  The  blue  goose  in  North  Carolina.  Auk 
50(3): 353. 

1933.  Recent  observations  on  eelgrass  conditions. 
Plant  Disease  Reporter  17(10)  :119-120. 

1933.  Further  reports  on  eelgrass.  Plant  Disease 
Reporter   17(11). 

1933.  Nelson's  sparrow  and  tree  swallow  winter- 
ing in  New  Jersej-.  Bird  Banding  4(2):  115. 

1933.  Spring  migration  of  the  great  blue  heron. 
Auk  50(4):427-428. 

1934.  Summer  bird  records  for  North  Cai-olina. 
Auk  51(0:94-96. 

1934.      Incursion  of  American  scoters   in  Norfolk. 

Connecticut.   Auk   51  (2) :  228-229. 
1934.     Possible  e.xtension  of  regular  winter  range 

of  the  great  black-backecl  gull.  Auk  51(3): 

376. 
1934.      Past   periods   of   eelgrass   scarcitv.    Rhodora 

36(427) :261-264. 
1934.     The  eelgrass  shortage  in  relation  to  water- 
fowl. Trans.  American  Game  Conf.  20:272- 

279. 
1934.     Eelgrass   disappearance   has   serious   effects 

on    waterfowl    and    industry.    Yearbook    of 

Agriculture     (1430) :  191-193. 

1934.  Summary  of  reports  of  baiting  and  other 
gunning  practices.  Bureau  of  Biological 
Survev  17  pp.   (Unsigned,  mimeographed.) 

1935.  (withT.  H.  Scheffer).  The  crested  myna, 
or  Chinese  starling,  in  the  Pacific  North- 
west. USDA  Tech.  Bull.  467.  27  pp;  3  pi., 
2  fig. 

1935.  The  present  situation  regarding  eelgrass 
(Zns/eia  marina).  Wildlife  Research  and 
Management   Leaflet   BS-3.    7   pp. 

1935.  (with  F.  M.  Uhler).  Mosquito  control  and 
its  effects  on  aquatic  wildlife.  Proc.  Amer. 
Game   Conf.   21:291-294. 

1935.  The  effects  of  arsenic,  as  used  in  poison- 
ing grasshoppers,  upon  birds.  .Xuk  52(1): 
118-119. 

1935.  Wasting  disease  of  Zoslrra  marina.  Nature 
135(3408):, 306. 

1935.  Late  migration  of  tree  swallows  and 
mourning   doves.    Auk   52(2):  189. 

1935.  Umjsual  food  habits  of  California  gulls. 
Condor    37(3):  170-171. 


Juno  1975 


TANNER:    CLARENCE   COTTAM 


235 


1935.     The  eelgrass  situation  in  1934.    Proc.  Anier. 

Game    Conf.     21:295-301. 
1935.     Winter  notes  from  coastal  North  Carolina.        1937. 

Auk  52(3):318-319. 
1935.     Waterfowl   problems   rlarified   by   stud\'   of        1937. 

gunning  practices.  Yearbook  of  Agriculture 

(1519):328-330. 
1935.     The    present    eelgrass    situation    along    the 

American    Atlantic    Coast.     Plant    Disease        1937. 

Reporter     19(  14)  :230-231.     (Presented     by 

Neil  Stevens  at  the  International  Botanical 

Conference  at  Amsterdam.  August.  1937. 

1935.     Further   notes   on   past   periods   of   eelgrass 

scarcity.    Rhodora    37(440) :  269-271. 
1935.     Blue    and    snow    geese    in    eastern    United        1937. 

States  in  the  winter  of  1934-35,  with  notes 

on  their  food  habits.  Auk  52(4) :  432-441. 
1935.      (with  F.  M.  Uhler).    Bird  records  new  or       1938. 

uncommon   to   Maryland.   Auk   52(4)  .-460- 

461.  ^  1938. 

1935.  Review  of  Food  preferences  and  require- 
ments   of    the    white-tailed    deer    in    New 

York    State,    bulletin    by    L.    A.    Maynard.        1938. 
Gardiner  Bump,  Robert  Darrow,  and  J.  C. 
Woodward.  J.  of  Mammal.   16(4)  :330-331. 

1936.  (with  I.  N.  Gabrielson.  A.  L.  Nelson,  and        1938. 
E.    R.    Kalmbach).    Christmas   bird   census. 

Port  Tobacco.  Maryland.  Bird  Lore  38(1). 

1936.      (with    F.    M.    Uhler).     The    role    of    fish- 
eating    birds.    Progressive    Fish    Culturist.        1938. 
M.    1-131     (14.3202):  1-14.    (Multigraphed. 
Bureau   of   Fisheries.) 

1936.     Earlj-  migration  of  the  great  black-backed 

gull.   Auk   53(1):81.  1938. 

1936.     Economic   oinithology   and   the   correlation 
of  laboratory  and  field  methods.    Wildlife 
Research  and   Management  Leaflet  BS-30.        1938. 
13    pp.;    1    fig. 

1936.     Food  of  the  limpkin.   Wilson  Bull.  48(1):        1938. 
11-13. 

1936.  Food  of  arctic  birds  and  mammals  col- 
lected by  the  Bartlett  Expeditions  of  1931- 
32-33.  J.  Washington  Acad.  Sci.  26(4) :  165- 
177.  1938. 

1936.  Notes  on  tlie  birds  of  Nevada.  Condor  38 
(3):  122-123. 

1936.     The  place  of  food  habits  research  in  wild-        1938. 
life   management.    Utah   Juniper   7:16-19. 

1936.     Status    of    the    black-backed    gull.    Auk    53 

(3):332-333.  .,  1938. 

1936.     Broken  wing   ruse   in   the   yellow   warbler. 

Auk  53  (4):  481.  "  1939. 

1936.      (with   H.    R.    Lewis).    Eelgrass   and   other 

waterfowl  foods:   present  status  and  future       1939. 
prospects.   Proc.   North  American  Wildlife 
Conf..    Wildlife    Restoration    and    Conser- 
vation,  pp.   498-500. 

1936.  Food    habits    of    North    American    diving        1939. 
ducks.     George     Washington     Univ.     Bull. 

pp.   66-70.    (Summarv  of  doctoral  disserta- 
tion.) 1939. 

1937.  (with  L  N.  Gabrielson,  A.  L.  Nelson,  and 

C.  F.  Smith).    Christmas  bird  census.  Port       1939. 

Tobacco.    Maryland.    Bird    Lore    39(1  ):45. 
1937.     Review  of  Home  life  and  economic  status 

of  the  double-crested  cormorant,  bv  H.   L.        1939. 

Mendall.  Auk  54(2)  :21 3-214. 
1937.     An    unusual    concentration    of    blue    jays.        1939. 

Bird  Banding  8  (2):  79-80. 
1937.     Speed  of  the  gray  fo.x.  J.  Mammal  18(2):        1939. 

240-241 . 
1937.      (with   F.   M.   Uhler).     Birds  in   relation  to 


fishes.      Wildlife     Research     and     Manage- 
ment Leaflet  BS-83.  16  pp. 
American  egret  and  black-bellied  plover  in 
Delaware  in  winter.  Auk  54(3)  :382. 
(with    J.    J.    Lynch).     Status    of    eelgrass 
{Zostera    marina)    on    the    north    Atlantic 
Coast,    January.      Wildlife    Research     and 
Management    Leaflet    BS-94.    15    pp. 
(with  A.   L.   Nelson  and  C.   S.   Williams). 
Uncommon   winter   birds  in   coastal   North 
Carolina.    Auk    54(4): 548. 
(with  A.  L.  Nelson).    Winter  nesting  and 
winter    food    of    the    barn    owl    in    South 
Carolina.   Wilson  Bull.  49(4) : 283-285. 
(with  J.  E.  Shillinger).    The  importance  of 
lead  poisoning  in  waterfowl.    Proc.   North 
Amer.    Wildlife   Conf.    2:398-403. 
Coot    swallowed   b}'   fish.    Wilson   Bull.    50 
(1):60. 

(with  W^  S.  Bourn).  Some  effects  of  mos- 
cjuito  control  on  wildlife.  Conserv^ation  4 
(2): 20-22. 

Review  of  Utah  birds  in  control  of  certain 
insect  pests,  by  G.  F.  Knowlton.  Utah  Acad. 
Sci.  14:159-166.  1937.  Auk  55(2) :300. 
Review  of  Biological  control  of  the  beet 
leafhopper  in  Utah,  by  G.  F.  Knowlton, 
Utah  Acad.  Sci.  14:111-139.  1937.  Auk 
55 (2): 303. 

Status  of  eelgrass  [Zostera  marina)  on  the 
north  Atlantic  Coast.  February.  Wildlife 
Research  and  Management  Leaflet  BS-110. 
7   pp. 

(with  H.  C.  Hanson).  Food  habits  of  some 
arctic  birds  and  mammals.  Zool.  Ser.  Field 
Mus.  Nat.  Hist.  .20 (31): 405-426. 
A  fatal  combat  between  a  heron  and  a 
snake.  Wilson  Bull.  50(2):  140. 
The  coordination  of  mosquito  control  with 
wildlife  conservation.  Proc.  New  Jersey 
Mosquito  Exterm.  Assn.  25:217-223.  Wild- 
life Research  and  Management  Leaflet  BS- 
119.   6  pp. 

(with  W.  S.  Bourn).  What's  wrong  with 
mosquito  control?  Trans.  North  Amer. 
Wildlife  Conf.  3:81-87,  98-99. 
(with  A.  L.  Nelson).  Why  study  the  food 
of  fur  animals?  Trans.  North  Amer.  Wild- 
life Conf.   3:527-530. 

Nesting   of    an    eastern   kingbird   in    a   de- 
serted   oriole   nest.     Condor   40(6)  :259. 
Great  blue  heron  swimming.  Condor  41  (1): 
37. 

(with  I.  N.  Gabrielson,  A.  L.  Nelson,  and 
A.  C.  Martin).  Christmas  bird  census.  Port 
Tobacco,  Marvland.  1938.  Bird  Lore  41 
(l):22-23. 

(with  Phoebe  Knappen).  Food  of  some  un- 
common North  American  birds.  Auk  56(2): 
138-169. 

Late  occurrence  of  nighthawk  in  Connect- 
icut.   Auk   56(2):188. 

(with  William  Vogt,  Victor  Cahalane,  and 
Aldo    Leopold).     Report   of   Committee   on 
Bird    Protection.    Auk    56(2)  :212-219. 
Food    habits    of    North    American    diving 
ducks.  USDA  Tech.  Bull.   643.   140  pp. 
The    eelgrass    situation    on    the    American 
Pacific   Coast.     Rhodora   41  (487)  :257-260. 
(with    A.    L.    Nelson    and   T.    E.    Clarke). 
Nptes   on   early   winter  food   habits  of  the 
black    bear    in     the    George     Washington 


236 


GREAT  BASIN  NATURALIST 


Vol.  35,  No.  2 


National    Forest.     .T.    Mammal.    20(3)  :310-        1942. 
314. 

1939.      (with  W.  S.  Bourn).    Need  mosquito  con-        1942. 
trol  be  incompatible  with  wildlife?  Trans. 
North    Amer.     Wildlife     Conf.     4:121-130. 
130-140,    143.  1942. 

1939.  (with  W.  S.  Bourn).  The  effects  of  lower- 
ing water  levels  on  marsh  wildlife.  Trans. 
North  Amer.  Wildlife  Conf.  4:343-350. 

1939.      (with    F.    M.    Uhler    and    T.    E.    Clarke).        1942. 
Food  of  snakes  of  the  George  Washington 
National  Forest.   Trans.  North  Amer.  Wild-        1942. 
life  Conf.  4:605-622. 

1939.      (witli  C.  S.  Williams).  Food  and  habits  of       1942. 
some    birds    nesting    on    islands    in    Great 
Salt  Lake.  Wilson  Bull.  51  (3):  150-155. 

1939.  (with  W.  S.  Bourn).    Marshes  of  the  past        1942. 
and    future.    Appendix    F,    pages    471-488 

in  Van  Campen  Heilner.  A  book  on  duck 
shooting.  Penn  Publishing  Company.  Phila-        1942. 
delphia. 

1940.  (with  I.  N.  Gabrielson,  A.  C.  Martin,  and 

A.  L.  Nelson).    Christmas  bird  census.  Port        1942. 

Tobacco,      Maryland.     1939.     Bird      Lore 

Suppl.  42(l):89-90. 
1940.      (with  V.  H.  Cahalane,  Aldo  Leopold,  and        1942. 

W.   L.  Finley).    Report  of  Bird  Protection 

Committee  in    1939.    Auk  57 (2): 279-291.  1942. 

1940.     Obituary    of    John    William    Sugden.    Sr. 

Auk    57(3):  448.  1942. 

1940.      (with  A.   L.   Nelson,   and   L.   W.    Saylor). 

The    chukar    and   hungarian    partridges   in 

America.     Trans.     North    Amer.     Wildlife        1942. 

Conf.  5:432.    Modern  Game  Breeding  and 

Hunting  Club  News   10(7)  :6-7,   12.    Wild-        1942. 

life  Research  and  Management  Leaflet  BS- 

159.    6   pp. 

1940.  (with  F.  M.   Uhler).   Birds  as  a  factor  in 
controlling    insect    depredations.     Wildlife        1942 
Leaflet     BS-162.     6    pp. 

1941.  Incubation   feeding   of   calliope    humming-        1943 
bird.    Auk    58(1):  59-60. 

1941.      (with  L  N.  Gabrielson.  A.  C.  Martin,  and        1943. 
A.  L.  Nelson).  Christmas  bird  census.  Port 
Tobacco,      Maryland.        1940.        Audubon 
Magazine  Supplement  43(1): 99.  1943. 

1941.  The  eelgrass  situation,  fall  1940.  Plant  Dis- 
ease Reporter  25  (2): 46-52.  1943. 

1941.     LeConte's  sparrow  in  Utah.  Condor  43(2): 

116-117.  1943. 

1941.  Indigo  bunting  and  band-tailed  pigeon  in 
Utah.  Condor  43 (2): 122. 

1941.      (with    Phoebe    Knappen).    Eskimo    curlew        1943. 
food  note  corrected.  Auk  58(2)  :256. 

1941.     Color     attractive     to     hummingbirds.     Auk 

58(2) :261.  1943. 

1941.      (with  V.  H.  Cahalane,  W.  H.  Finley.  and 
Aldo   Leopold).     Report  of   the   Committee 
on   Bird   Protection.    1940.   Auk   58 (2): 292-        1943 
298. 

1941.     California   cuckoo  in  southeastern  Nevada.        1943. 
Condor  43(3): 160. 

1941.     How  fast  can  a  fox  squirrel  run?  J.  Mam-        1944. 
mal.  22(3):  323. 

1941.      (with     C.     S.     Williams).      Wilson     snipe 

perches  on   telephone  pole.   Condor  43(6):        1911. 
293. 

1941.  European  starling  in  Nevada.  Condor 
43(6):293-294. 

1941.      (with   James   Moffitt).     Eelgrass   depletion        1944. 
on   the   Pacific    coast    and    its   effect   upon 
black  brant.    Wildlife  Leaflet  204.  26  pp. 


European  widgeon  at  the  upper  Souris 
National  Wildlife  Refuge.  Auk  59(1):  104. 
(with  V.  H.  Cahalane,  Aldo  Leopold,  and 
W.  L.  Finlev).  Report  of  the  Committee 
on  Bird  Protection.  Auk  59(2)  :286-300. 
(with  I.  N.  Gabrielson,  F.  M.  Uhler,  and 
A.  L.  Nelson).  Christmas  bird  census.  Port 
Tobacco.  Marvland.  1941.  Audubon  Maga- 
zine   44(1  ):27. 

Supplementary  notes  on  the  food  of  the 
limpkin.  Nautilus  55(4) :  125-128. 
Records  from  extreme  northeastern  Neva- 
da. Condor  44(3):  127-128. 
(with  C.  S.  Williams,  and  C.  A.  Sooter). 
Flight  and  running  speeds  of  birds.  Wilson 
Bull.   54(2):121-131. 

(with  Seth  Low,  and  R.  E.  Griffith). 
Glaucous  gull  in  Oklahoma.  Wilson  Bull. 
54(2):  139-140. 

(with  C.  S.  Williams,  and  C.  A.  Sooter). 
Cooperative  feeding  of  white  pelicans. 
Auk  59(3):444-445. 

(with  C.  A.  Sooter,  and  R.  E.  Griffith). 
The  European  starling  in  New  Mexico. 
Condor   44(4):  182. 

Slate-colored  junco  in  Nevada.  Condor  44 
(4): 185. 

Food  of  the  ruddy  turnstone.  Auk  59(4): 
581. 

(with   C.   A.    Sooter.    and   R.    E.   Griffith). 
The  yellow  rail   and   the   Caspian  tern   in 
New   Mexico.   Condor  44(5)  :230. 
Coyote  without  external  ears.  J.  Mammal. 
23(4): 450. 

(with  C.  S.  Williams,  and  C.  A.  Sooter). 
Some  unusual  winter  visitors  or  late  mi- 
grants to  the  Bear  River  Marshes.  Utah. 
Great  Basin  Naturalist  3(2):  51 -53. 
New  or  uncommon  Utah  bird  records. 
Wilson  Bull.  54(4):254-255. 
Bullock's  oriole  as  a  fighter.  Auk  60(1): 
94-95. 

(with  V.  H.  Cahalane,  W.  L.  Finley,  and 
Aldo  Leopold).  Report  of  Committee  on 
Bird  Protection.  Auk  60(1):  152-162. 
(with  C.  S.  Williams).  Speed  of  some 
wild  mammals.  J.  Mammal.  24(2) :  262-263. 
Greater  vellow-legs  as  a  fish-eater.  Wilson 
Bull.  55(2):  128. 

Is  the  starling  population  decreasing  in 
northeastern  United  States?  Auk  60(3): 
439-440. 

(with  C.  S.  Williams,  and  G.  H.  Jensen). 
Some    birds    not    commonlv    observed    in 
Utah.    Condor   45(4):  159-160. 
(with    L.    E.    Givens,    and    D.    V.    GrajO- 
Vermilion  flvcatcher  at  St.  Marks,  Florida. 
Wilson    Bull.    55 (3):  192. 
Ihiusual    feeding    habits    of    grnckles    and 
crows.   Auk   60(4) :  594-595. 
Tlie    least    bittern    at    Long    Lake.    North 
Dakota.  Wilson  Bull.  55(1): 54. 
(wath    A.    M.    Woodbury).     Type   locality 
of  Pcrisoreus  canadensis  capitalis  Ridgway. 
Auk   61(1):  131 -132. 

(with  J.  J.  Lynch,  and  A.  L.  Nelson). 
Food  habits  and  management  of  American 
sea  brant.  J.  Wildlife  Mgmt  8(1): 36-56; 
3    pi.  - 

fwith  C.  C.  Sperry).  The  greater  and 
lesser  yellow-legs  as  fish  caters.  Wilson 
Bull.    56(1):45. 


Iiiiie  1975 


TANNER:    CLARENCE   CX)TTAM 


237 


1944.     Birds  and  the  west  Tennessee   River  Lake. 

Migrant    15(  1):L5;    2    maps. 
1944.     Gulls  as  vegetarians.  Condor  46(3) :  127-128.        I  9  17. 
1944.     Eastern    lark    sparrow    and    upland    plover 

in  western  Tennessee.  Migrant  15(2)  :29. 
1944',     Starlings    feeding    on    the    liacks    of    cattle. 

Migrant   15(2) : 24-25.  1947. 

1944.  The    role    of    impoundments    in  'post-uar 
plaiming     for     wateifowl.     Trans.     Noitli        1947. 
Amer.  Wildlife  Conf.  9:288-295. 

1945.  Diving  habits  of  the  shoveller  duck.  Condor        1947. 
47(1  ):39. 

1945.      (with   A.   L.    Nelson,    and    W.    S.    Bourn).        1947. 
Red  fox  breeding  in  salt   marsh.   J.   Mam 
mal.   26(1): 91-92.  1948. 

1945.     Speed    and    endurance    of    the    covote.    J. 
Mammal.   26(1): 94. 

1945.      (with  J.  H.  Steenis).    A  progress  report  on        1948. 
the  marsh  and  aquatic  plant  jnohlcm.  Reel- 
foot  Lake.   .1.  Tennessee  Acad.   Sci.   20(1): 
6-19.  1948. 

1945.     The     ruddv     turnstone     in     Ltah.     Condor 

47(2): 79.  1948. 

1945.     Feeding   habits    of    the    Clark's    nutcracker. 

Condor   47(4): 168.  1948. 

1945.     Eelgrass  conditions  along  the  Atlantic  sea- 
board   of    North    America.     Plant    Disease        1949. 
Reporter    29(12) :  302-310. 

1945.     Some  records  of  birds  in  LTtah.    Condor  47 

(4):  172-173.  1949. 

1945.     California  gulls  feeding  on  midges.  Condor 
47(5):216.' 

1945.     Research    problems    on    the    LI.S.    National        l')4<». 
Wildlife     Refuges.     Trans.     North     Amer. 
Wildlife  Conf.    10:347-355.  1949. 

1945.     American  eider  in   Delaware.   Auk   62(4): 
634. 

1945.     The  whistling  swan  in  Maine.  Auk  62(4):        1949. 
634-635. 

1945.  (with   H.   S.   Zim).     Bad  news   for  brother 

rat.     Saturdav    Evening    Post    2I8(19):17.        1949. 
113-116. 

1946.  Abundance  of  wildlife  in  Illinois  a  century 
ago.  Illinois  Conservation.  Winter   1945-46. 

p.  41.  1949. 

1946.      (with  Elmer  Higginsj.  DDT  and  its  effect 

on  fish  and  wildlife.  .1.  Econ.  Ent.   39(1):        19-1-9. 

44-52. 
1946.     Late    nesting    of    Caspian    tern    in    Utah.        1949. 

Condor  48(2): 94-95. 

1946.  (with    Clifford    Presnall).     Partnership    in 
conservation   of  renewable   resources.   State        1949. 
Government    19(6) :  153-157.    162.   ,Iune. 

1947.  Missouri     River     Basin     development:     Its        1949. 
effects  on  fish  and  wildlife.   Outdoor  Amer- 
ica 12(3).  Februaiy.    Pi'oc.  36th  Conv.  Int. 

Asso.  Game,  Fish,  Cons.  Comm.  September        1950. 

9-11,  1946.  pp.  200-206.  North  Dakota  Out- 
doors.   July.    pp.    12-14.    Wyoming    Wild-        1950. 

life  11  (6): 20-27.  June. 
1947.     Unusual  flight  of  American  egret.  Migrant 

(Published     by    Tennessee     Ornithological        1950. 

Society).   June.   p.   27. 
1947.     Zone-tailed   hawk   feeds   on    rock    squirrel. 

Condor    59(5)  :210.    September-October. 
1947.     Late   nests   in  Yellowstone   National   Park.        '^'^O- 

Wilson  Bull.   59 (3):  172-1 73.   September. 
1947.     Some  improvements  needed  in  wildlife  re-        1950. 

search.    J.   Wildlife   Mgmt.    1 1  (4) :  339-347. 

October.  1950. 

1947.     Waterfowl    at   the   crossroads.    Trans.    12th 


North  Amer.  Wildlife  Conf.  Ivhruary. 
pp.   67-85. 

Piesent  eelgrass  condition  and  problems  on 
the  Atlantic  Coast  of  North  America. 
Trans.  12th  North  Amer.  Wildlife  Conf. 
February,   pp.   387-398. 

Some    bird    records    for    southern    Nevada. 
Condor  49(6): 2+4.   November-December. 
Utah    swans    and    geese.    Utah    Magazine 
9(9):  10-13,   25-29.   September. 
Famous    geese    of    Utah.    LTtah    Magazine 
9(10):  12-15.  24-27.  October. 
The  puddle  ducks  of  LUah.  Utah  Magazine 
9(10):20-23.   October. 

Unusual  consumption  of  fish  bv  three 
species  of  birds.  Wilson  Bull.  fi0?2):117- 
118.    June. 

Our  waterfowl  problena  and  program. 
Proc.  19-1-8  Northeastern  Game  Conf.  pj). 
125-131. 

Aquatic     habits    of    t.lie     Norwav     rat.      J. 
xMammal.   29(3):299.   August. 
The    mourning    dove    in    Alaska.    Wilson 
Bull.   60(3):  188-189.   September. 
Yellow-headed    blackbird    on    Long    Island. 
Auk  65  (4):  605. 

The  trumpeter  swan,  greatest  of  American 
waterfowl,  stages  a  comeback.  The  Field, 
London.  England.  26  March: 350-351. 
(with  Angus  M.  Woodbury,  and  John  W. 
Sugden).  Annotated  checklist  of  the  birds 
of  Utah.  Bull.  Univ.  Utah  39(16).  March. 
Swinmiing  pigeons.  Condor  51  (  3) :  150-1  51. 
May- June. 

Does  stocking  pay?  Texas  Game  and  Fish 
7(6)  :6.  33.  May.  Wildlife  in  North  Caro- 
lina 8(5):  118-120. 

Limiting  factors  of  present  waterfowl 
knowledge.  Trans.  14th  North  Amer.  Wild- 
life Conf.   j)p.  42-57. 

The  effects  of  uncontrolled  introductions 
of  plants  and  animals.  Proc.  Papers  Inter- 
nal. Tech.  Conf.  Protect.  Nat..  Lake  Suc- 
cess. N.Y.  August,  pp.  408-413. 
The  eelgrass  story.  Virginia  Wildlife  10 
(10):  16-18. 

Wildlife   research   pays  off.    Sports   Afield. 
November,  pp.  34-35,  58-60. 
Conservation     in     Mississippi.     Mississippi 
Game    and    Fish     13(4)  :4-5.     12.    October. 
13(5):  10-12.  November. 

Further     needs     in     wildlife     research.     J. 
Wildlife    Mgmt.    13(4):333-341.    October. 
Conservation    in    New    Zealand    and    LTtah. 
The   Pioneer,    Salt    Lake    Citj-.    Utah.    Sep- 
tember,  pp.  5-6. 

Rehabilitation  in  the  wild.  Auk  67(2): 
238.   April. 

(with    James    Silver).     The    last    stand    of 
the    key    deer.     National    Parks    Magazine 
24(102) :  100-103.   July-September. 
Planned    enterprises    and    their    effects    on 
the  balance  of  nature.  Proc.  and  Papers  of 
Internal.   Tech.   Conf.   Protect.    Nat..   Lake 
Success.   N.Y.   August    1949.    pp.   323-325. 
Our  federal  government  and  consei-vation. 
Proc.  Utah  Acad.  Sci.  25:59-67.  1948. 
The  waterfowl  regulations.  Maryland  Con- 
servationist 27(2)  :4-5,  28.   Summer. 
The  why   of  migrator)^  waterfowJ   regula- 
tions. Proc.  International  Assoc,  of  Fish  & 


238 


GREAT  BASIN  NATURALIST 


Vol.  35,  No.  2 


Game  Commissioners.  Texas  Game  and  Fish 
9(l):4-7,  31. 

1951.  (with  Warren  S.  Bourn).  Some  biological 
effects  of  ditching  tidewater  marshes.  Re- 
search Report  19.  January. 

1951.  (with  Herbert  Zim).  Insects.  A  guide  to 
familiar  American  insects.  Simon  &  Schue- 
ster.   New   York. 

1949.  A  New  Zealand  appraisal.  The  Wood 
Thrush  5(2).  November-December.  Au- 
dubon Society,  District  of  Columbia. 

1951.  Waterfowl's  future  depends  upon  manage- 
ment. Trans.  16th  North  Amer.  Wildlife 
Conf.  March  5-7.  pp.    109-121. 

1951.  Snow  goose  record.  The  Oriole  16(2)  :23. 
June. 

1951.  Fifty  years  of  progress  and  handicaps  in 
wildlife  management  in  the  U.S.  Trans. 
41  Con.  Int.  Asso.  Game,  Fish  and  Cons. 
Commissioners,  Rochester.  N.Y.  Wyoming 
Wildlife  15(10)  :4-ll,  31-34.  October.  16 
(l):22-29.  January  1952. 

1952.  Present  status  of  migratorj'  game  birds. 
Atlantic  Nat.  9(3) :  118-123. 

1952.  Management  of  our  waterfowl.  Happy 
Hunting  Ground  (magazine  of  Kentucky 
fish  and  game  division).  Januar3\ 

1951.  Conservation  of  our  wildlife  and  other 
renewable  resources.  The  County  Officer. 
December,  pp.  301-307. 

1952.  Chemical  controls  vs.  wildlife.  Ducks  Un- 
limited Quarterly  15  (2):  7.  Spring.  South 
Dakota  Conservation  Digest.  May.  pp.  14- 
15.  North  Dakota  Outdoors.  June.  pp.  10- 
11.    Tennessee  Conservationist    17    (7):6-7. 

18.  July  Iowa  Conseivationist   11(7).  July. 

1952.  (with  Warren  S.  Bourn).  Coastal  marshes 
adversely  affected  by  drainage.  Presented 
at  North  Amer.  Wildlife  Conf.    March  17- 

19.  Published  in  Proceedings  and  in 
Florida  Nat.  25(2).  April. 

1953.  Jamaica  Bay  sanctuary.  Bulletin  to  the 
Schools  (Arbor  and  Wild  Life  Dav  issue) 
39(7):213-217.  March.  University  of  the 
State   of   New   York. 

1953.  (with  Philip  DuMont).  What  has  hap- 
pened to  the  key  deer?  National  Parks 
Magazine,  p.   82.  April-June. 

1953.  Wildlife  and  chemicals.  Virginia  Wildlife 
14(5):10-12. 

1953.  Conservation  of  America's  faunal  re- 
sources. Proc.  Symposium  on  Cons,  of  Re 
newable  Nat.  Res.  of  the  French-Canadian 
Assoc,  for  the  Adv.  of  Sci.,  Laval  Univ.. 
Quebec,  Canada.   October   1952. 

1953.  Report  of  committee  on  wildlife  conser- 
vation planning.  Proc.  Int.  Assoc,  of  Game. 
Fish   &  Cons.   Comm.   pp.   60-65. 

1953.  (with  Geo.  A.  Rounsefell,  and  W.  Harry 
Everhart).  Fishery  science:  its  methods 
and  applications.  .lohn  Wiley  &  Sons.  Inc.. 
New    York. 

1953.  Does  upland  game  beai-  its  fair  share  of 
hunting  pressure?  Proc.  Midwest  Assoc,  of 
Fish,  Game.  Cons.  Conmi.   pp.    184-206. 

1954.  Letter  to  the  editor  of  journal  wildlife 
management.  Re  Carp  removal  at  Lake 
Mattemuskeet.  A  controversv  1«(1):134- 
135. 

1954.     Twin    opportunities:    conservation    and    en- 


gineering. Proc.  Amer.  Soc.  Civil  Engineers 
80(419).  March. 
1954.      (with   David   A.    Munro).     Eelgrass   status 
and    enviromnental    relations.     J.    Wildlife 
Mgmt.    18(4):  449-460. 

1954.  Bird  records  for  Nevada.  Condor  56(4): 
223-4. 

1955.  The  Welder  Wildlife  Foundation.  Inter. 
Assoc.  Game.  Fish.  Cons.  Comm.  Proc.  45: 
133-134. 

1955.  Progress  in  wildlife  restoration  and  train- 
ing. Natl.  Wildl.  Conf.  Pub.  20th  Trans. 
North  Amer.  Nat.  Wildl.  Conf. 

1955.  On  the  food  habits  of  the  diving  ducks  in 
Denmark.  J.  Wildlife  Mgmt.  19(4).  Oc- 
tober. 

1955.  Conservation  and  engineering  partnership. 
Virginia   Wildlife. 

1955.  The  Welder  Wildlife  Foundation.  Inter- 
national   Proceedings. 

1956.  Little  thing  may  become  big.  Instructor 
91(1):9. 

1956.  History  of  waterfowl  management  in 
America.  Univ.  Minnesota.  Processed  Proc. 
by  Univ.  in   1957. 

1956.  Water  and  waterfowl.  Minnesota  Conser- 
vation Federation.  January  21. 

1956.  Who's  doing  what  in  wildlife  research. 
Minnesota,  North  Carolina  State  Wildlife 
Societv.  January  20. 

1956.  The  Welder  Wildlife  Foundation.  Atlantic 
Naturalist.  Washington,  D.C.  1U4):163- 
168.   March   and  April. 

1956.  Prairie  ducks:  a  study  of  behavior,  ecology, 
and  management.  Review.  Wilson  Bull. 
68(l):86-87. 

1956.  Seeing  America's  wildlife  in  oia-  national 
refuges,  a  review.  Ecology. 

1956.  Chemical  controls  in  relation  to  wildlife. 
Virginia  Wildlife.  March  and  April. 

1956.  Marking  birds  for  scientific  purposes.  Ecol- 
ogy  37 (4): 675-681. 

1956.  The  problem  of  wildlife  introductions:  its 
success  and  failures.  Proc.  Int.  Assoc,  of 
Game,  Fish,  Cons.  Comm. 

1956.  Some  wildlife  research  problems  of  the 
southeast.  Proc.  1956  Virginia  Wildlife, 
and    other   states. 

1957.  Chemical  controls  in  relation  to  wildlife. 
Texas  Game  and  Fish  Magazine.  Con- 
gressional Record.  Appendix  pp.  A4974- 
A4976. 

1957.  The  role  of  piivate  agencies  in  conser- 
vation. Proc.  Int.  Assoc,  of  Game,  Fish.  & 
Cons.   Comm. 

1957.  Future  i-ole  of  federal  agencies  in  forest 
wildlife  management.  Proc.  Soc.  Amer. 
Foresters.  Syracuse.  N.Y.  pp.  97-100. 

1057.  Late  nesting  in  south  Texas.  Texas  Acad. 
S(i.   (processed).  Deceinber   H. 

1958.  (with  W.  C.  Glazencr).  Exotics.  Texas 
Game  and  Fish    16(l):-1'-7.  Jaiuiar\ . 

1958.  Wildlife:  its  lole  in  our  dynaniii  e(oiiom>'. 
Conference  of  Southwest  Foundations 
(processed).   April. 

1958  Waterfowl  research:  whv  research  is 
needed.  Naturalist  9(1):28-31. 

1958.  Wildlife  and  water  conservation.  J.  Soil 
and    Water    Cons.    1-3(2)  :65-69. 

1958.  Science  and  resources  in  America.  Pro- 
cessed 23rd  No.  Amer.  Wildl.  Conf.  Vir- 
ginia Wildlife.  May. 


Line 


1975 


TANNER:    CL.\RENCE   COTTAM 


239 


1958.     Coturnix     quail     in    America.     Proc.     Int.        1960. 
Assoc.  Game,  Fish,  Cons.  Comm.  pp.   111- 
119. 

1958.  A   commentary   on    the   fire   ant   problem.        1960. 
Proc.   Symposium  on   Fire  Ant,  Southeast- 
ern   Assoc,    of    Game    and    Fish    Commis- 
sioners. Columbia,  South  Carolina.  October        1960. 
19-22. 

1959.  The     Welder     Wildlife     Foundation     ap- 
proach to  research  and  education.  Gazette        1961. 
8(1):  17-19.  53-54.  January.  Salt  Lake  City. 

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western  Naturalist.    In   press. 


EVOLUTIONARY  DIVERGENCE  IN  CLOSELY  RELATED 

POPULATIONS  OF  MIMULUS  GUTTATUS 

(SCROPHULARIACEAE) ' 

Karen  W.  Hughes-  and  Robert  K.  Vickery.  Jr.^ 


Abstract. —  The  evohitionary  divergence  of  five  closely  related  populations  of  the  3'ellow  monkey 
flower.  Mimulus  guttatus.  was  compared  with  the  length  of  time  the  populations  could  have  occu- 
pied the  site  and  with  the  ability  of  the  populations  to  intercross.  Populations  from  the  younger 
sites  were  found  to  be  morphologically  intermediate  to  i)opulations  from  the  older  sites.  Ability  to 
intercross  was  not  correlated  with  morphological  divergence   at  this   stage   of  evolution. 


Glaciation  patterns  of  the  Wasatch 
Mountains  of  Utah  provide  an  estimate 
of  the  geological  age  of  existing  plant  pop- 
ulations. If  evolutionary  relationships  be- 
tween isolated  populations  can  be  deter- 
mined, an  estimate  of  rates  of  evolution  for 
these  populations  may  be  obtained.  The 
purpose  of  this  investigation  is  to  deter- 
mine the  amomit  of  divergence  between 
populations  from  previously  glaciated  and 
unglaciated  sites  and  to  relate  this  di- 
vergence to  the  age  and  possible  origin  of 
the  populations.  Divergence  is  estimated 
by  calculating  a  Prim  network  (Prim, 
1957),  based  on  morphological  charac- 
teristics of  the  populations,  and  by  analy- 
zing the  ability  of  the  populations  to 
intercross.  The  Prim  network  was  calcu- 
lated for  24  morphological  traits  and  for 
6  selected  traits  to  determine  w^iether 
larger  numbers  of  characters  make  a  sig- 
nificant difference  in  the  network. 

Methods  and  Materials 

Five  populations  of  Mimulus  guttatus 
Fischer  ex  D.C.,  the  yellow-  monkey 
flower,  from  two  adjacent  canyon  drain- 
ages were  selected  for  investigation  (Table 
1 ) .  M.  guttatus  grows  in  small  isolated 
populations  near  streams  or  springs  and 
is  thus  ideal  for  studies  of  evolutionary 
divergence.  Plants  were  grown  from  seeds 
in  the  greenhouse.  Quantitative  data  were 
gathered  for  24  morphological  traits  on  an 
average  of  16  plants  per  population*.  We 
were  unable  to  select  a  larger  number  of 
traits  because  of  the  close  morphological 
relationships  of  the  populataions.  Traits 
selected  include  height,  dr}'  weight,  degree 
of  pigmentation,  timing  and  extent  of 
flowering  and  seed  set,  etc.    (Crook,  1964) . 

^This   invesligation   was   supported   in   part    by   a   Public   Heal  I 
Division    of   General    Medical    Sciences,    Public   Health    Service. 
-Department    of    Botany.    University    of    Tennessee,    Knoxvill 
^Department    of    Biolopy.    University   of   Utali.    Salt   I-akc    Cil 


Plants  from  each  population  were  inter- 
crossed in  all  comfjinations  but  one,  i.e., 
diallel  design. 

The  estimated  age  of  the  populations 
was  determined  from  the  period  of  time 
each  site  was  available,  geologically,  for  oc- 
cupation by  the  populations.  The  Thou- 
sand Springs  and  Storm  Mountain  sites 
have  never  been  glaciated  and  apparently 
have  been  available  for  occupation  for 
50,000  years  or  more,  while  the  remaining 
three  sites  were  glaciated  and  have  been 
available  only  since  the  retreat  of  the  Wis- 
consin period  glaciers  (Table  2).  Pop- 
ulations may  not  be  as  old  as  the  site  and 
may  have  migrated  up  and  down  the 
moinitainsides  with  changing  environmen- 
tal conditions  at  the  end  of  the  glacial 
epoch. 

Table    1.    Populations   of   Mimulus  guitalus 


Culture    number 


Location 


6648  Big  Water  Gulch,  Millcreek 
Canyon.  Along  stream  south  of 
parking  lot  at  end  of  road. 
Altitude   7,680  feet. 

5840  Thousand      Springs,      Millcreek 

Canyon.  Hillside  adjacent  to 
roacl.  Site  is  marked  with  a 
road  sign.    Altitude  7,200  feet. 

6649  Brighton  Loop,  Big  Cotton- 
wood Canyon  on  Clayton  Peak 
drainage.  East  edge  of  Brigh- 
ton Loop  at  top  of  canvon.  Al- 
titude 8.760  feet. 

5839  Spruces,    Big    Cottonwood    Can- 

yon. Stream  and  marsh  areas 
of  Spruces  Campground.  Al- 
titude  7.360  feet. 

6127  Storm     Mountain,     Big    Cotton- 

wood Canyon.  Stream  in  small 
canyon  south  of  Storm  Moun- 
tain Campground.  Altitude  6,240 
feet. 


Service    Followsliip     1  No.     D-Fl-GM- 


3/920. 
841 12 


from     the 


240 


June  1975 


HUGHES,  VICKERY:    MIMULUS  GUTTATUS 


241 


Table  2.  Maximum  estimates  of  time  available 
for  occupation  of  sites  by  Mimulus  guttatus  pop- 
ulations. 


Site 


Most  recent  glaciation 


Brighton 

Spi-uces 

Storm  Mountain 

Big  Water  Gulch 

Thousand   Springs 


9,000  j'ears  before  present* 
11.000  j'ears  before  present* 

Not  glaciated 
11,000  years  before  present* 

Not   glaciated 


•     Deglaciation    occuitchI    later    lluin    ihesc    (lales. 

The  24  morphological  traits  were  ana- 
lyzed by  the  principal  components  me- 
thod of  factor  analysis  as  outlined  by 
Harman  (1964)  to  obtain  factors  composed 
of  mutually  correlated  groups  of  traits. 
Six  major  factors  were  obtained  with 
eigenvalues  greater  than  one  (Crook, 
1964). 

The  morphological  similarities  of  the 
populations  were  determined  by  calcula- 
tions of  a  shortest  distance  network  de- 
veloped by  Prim  (1957)  and  adapted  to 
evolutionary  studies  by  Edwards  and  Ca- 
valli-Sforza  (1964).  In  this  method  the 
sum  of  the  normalized  character  differ- 
ences between  the  populations  is  the  esti- 
mate of  the  difference.  Populations  with 
large  numbers  of  differing  traits  are 
widely  separated  on  the  network  and  vice 
versa.  The  assimiption  implicit  in  the  use 
of  the  Prim  network  for  evolutionary 
studies  is  that  the  sum  of  the  character 
differences  is  jDroportional  to  the  evolu- 
tionar}'  differences  between  the  popula- 
tions. It  should  be  noted  that  Edwards 
and  Cavalli-Sforza  made  assiunptions  of 
independence  and  selective  neutrality  not 
made  here.  The  assumption  of  neutrality 
is  not  necessary,  as  we  are  interested  in 
divergence,    whether   random    or   the   re- 


sult of  selective  pressures.  The  24  mor- 
phological traits  are  clearly  not  indepen- 
dent, as  they  may  be  combined  into  6 
groups  of  mutually  correlated  traits  by 
factor  analysis.  Some  of  the  24  variables 
are  represented  in  more  than  one  factor 
(Crook,  1964).  The  6  traits  are  as  inde- 
pendent as  can  be  obtained  in  that  they 
represent  high  factor  loading  on  one  fac- 
tor and  very  low  loadings  on  the  other 
factors.  Prim  networks  were  calculated 
for  all  24  characters  and  for  the  6  char- 
acters  obtained   from   factor   analysis. 

Results  and  Discussion 

The  6  major  factors  derived  from  the 
factor  analysis  of  24  characters  can  be 
identified  as  follows.  Factor  1  is  a  compos- 
ite factor  representing  general  bushiness 
of  the  plant,  with  high  factor  loadings  on 
the  number  of  leaves  and  branches  and  on 
measurements  related  to  the  age  of  the 
plant.  Factor  2  represents  dry-weight 
measurements,  wdth  emphasis  on  roots 
and  runners.  Factor  3  represents  almost 
entirely  pigment  measurements,  with 
negative  loadings  on  age.  Factor  4  is  a 
general  measure  of  height.  Factor  5  is  a 
general  measure  of  flower  size.  Factor 
6  has  the  highest  loadings  on  duration  of 
flowering  and  seed  set.  The  characteris- 
tic with  the  highest  factor  loading  in  each 
factor  was  selected  for  analysis  by  the 
Prim  network  (Table  3). 

The  Prim  networks  based  on  24  mor- 
phological characters  and  on  the  6  se- 
lected morphological  characters  are  given 
in  Figures  1  and  2.  Both  networks  are 
identical  in  pattern,  but  the  distances  be- 
tween  populations  vary   somewhat. 


•H 
U 

PQ 


o 

iH 

P 

o 

u 

<u 

w 

+J 

0) 

<0 

o 

^ 

:3 

u 

en 

0* 

•H 

CO 

CQ 

37.33 


17.48 


21.70        I     23.77 


Total   distance  =    100.28 


Fig.      1 .     Prim    network    for    24    morphological    traits. 


242 


GREAT  BASIN  NATURALIST 


Vol.  35,  No.  2 


Table  3.  Morphological  traits  selected  by  fac- 
tor analysis. 

1.  Total    number    of    branches    at    end    of    seed 
production 

2.  Total    dry    weight    recorded    at    end    of    seed 
production 

3.  Number    of    internodes    on    main    stem    that 
show   anthocyanin   pigment 

4.  Length   in  mm  of  longest  internode 

5.  Length  of  last  flower  produced  by  the  plant 

6.  Time   in   days    of    seed    production 

All  populations  tested  intercrossed.  In 
only  two  cases  was  the  seed  production 
reduced  (Figure  3).  Thousand  Springs 
2  X  Spruces  c^  produced  an  average  of 
89  seeds  per  capsule.  Spruces  9  X  Brigh- 
ton cT  produced  an  average  of  90.61 
seeds  per  capsule.  The  average  parental 
seed  production  for  all  populations  in  the 
study  was  173.67.  Big  Water  Gulch  and 
Thousand  Springs  were  not  intercrossed. 

The  correlation  of  Prim  network  dis- 
tances between  populations  and  the  num- 
ber of  seeds  produced  was  not  significant 
when  Prim  network  distance  was  calcu- 
lated from  all  24  traits  (r=  -.026)  and 
barely  significant  when  Prim  network 
distance  was  calculated  from  6  traits 
(r=   -.443,  p  =  .05). 

Conclusion 

The  similarity  of  the  two  Prim  net- 
works indicates  that  fewer  characteris- 
tics may  be  used  without  distorting  the 
observed  pattern  of  relationships  between 
the  populations;  however,  the  distances  be- 
tween populations  do  change  somewhat. 
In  this  study  the  characteristics  for  the 
second  network  were  selected  by  factor 
analysis;  however,  the  factors  represented 


broad  generalized  categories,  and  an  in- 
vestigator with  an  extensive  knowledge 
of  his/her  organism  might  intuitively 
select  similarly  representative  factors.  Fac- 
tor analysis  might  also  be  used  in  a  pre- 
liminary study  by  investigators  who  lack 
resources  for  collecting  large  amounts  of 
data. 

In  both  Prim  networks  the  oldest  pop- 
ulations, i.e.,  populations  occupying  sites 
that  have  been  available  for  the  longest 
period  of  time,  appear  at  either  end  of 
the  network,  while  populations  from  the 
newer  sites  occupy  intermediate  jiositions. 
This  finding  is  in  agreement  with  the  pat- 
tern of  evolution  expected  if  the  younger 
populations  received  portions  of  their 
gene  ])ool  from  the  older  established  pop- 
ulations. Mimulus  is  an  edible  plant,  and 
Lindsay  (1960)  has  shown  that  seeds  can 
pass  undigested  through  the  digestive 
tract  of  birds.  It  is  thus  possible  that  birds 
and  mammals  could  carry  the  seeds  from 
site  to  site.  Seeds  deposited  in  recently 
deglaciated  sites  would  find  little  com- 
petition from  other  plants  and  woitld  be 
likely  to  survive.  Seeds  carried  to  well-es- 
tablished older  sites  from  the  younger 
sites  would  have  to  compete  with  well- 
adapted  genotypes.  Thus  the  younger 
populations  are  likely  to  be  a  mixture  of 
genotypes  from  the  older  populations, 
while  the  older  populations  would  tend  to 
maintain  their  well-adapted  genotypes.  If 
this  pattern  of  evolution  is  correct,  we 
would  expect  that  the  younger  popula- 
tions ^^•ould  be  a  mixture  of  genotypes 
from  the  older,  well-established  popula- 
tions and  would  be  intermediate  to  the 
older  populations  on  the  Prim  network. 
The  network  suggests  that  older  popula- 


c 
c 

en 

•H 
U 

m 


8.52 


4.45 


4.05 


o 

o 
u 

0) 
4-1 

CT" 

•H 


3.98 


Total   distance 


21.00 


Fig.  2.     Prim  network  for  six  morphological   traits. 


June  1975 


HUGHES,  VICKERY:    MIMULUS  GUTTATUS 


243 


BRIGHTON 


SPRUCES 


THOUSAND 
SPRINGS 


STORM  MT. 


AVERAGE  SEED  SET  PER  CROSS 


0  to  100 
101  to  200 
201  to  300 


301  to  i+00 
UOl  to  500 


Fig.  3.  Seeds  produced  by  crosses  between  populations  of  Mimulus  guttatus.  The  direction  of 
tlie  arrow  is  from  male  pollen  donor  to  female.  Crosses  within  populations  are  indicated  by  a  bar 
within   the  circle.     Self-pollinations   are  indicated  by  the  thickness  of  the  circle  circumference. 


tions  are  more  closely  related  to  the  young- 
er populations  in  adjacent  canyons  than 
to  younger  populations  in  the  same  can- 
yon. This  may  reflect  the  results  of  se- 
lection on  a  mixture  of  genotypes  rather 
than  patterns  of  population  establishment. 
The  failure  of  crossing  data  to  corre- 
late with  the  Prim  network  results  is  not 
surprising.  None  of  the  populations  have 
shown  significant  morphological  differ- 
ences, and  barriers  to  gene  exchange  have 
not  evolved.  At  such  a  point  a  random  ac- 
cumulation of  gene  differences  might  or 


might  not  cause  a  reduction  in  ability  to 
intercross,  depending  upon  which  genes 
are  involved. 

Summary 

Prim  networks  based  on  24  and  6  mor- 
phological traits  were  identical  with  re- 
spect to  order  of  the  populations.  How- 
ever, the  distances  between  the  popula- 
tions did  vary  from  population  to  popula- 
tion. The  two  oldest  populations  appeared 
at  either  end  of  the  network,  while  the 
more  recently  established  populations  ap- 


244 


GREAT  BASIN  NATURALIST 


Vol.  35,  No.  2 


peared  in  the  middle.  Probably  the  older, 
well-adapted  populations  donated  a  por- 
tion of  their  gene  pools  to  the  younger 
populations. 

References  Cited 

Crook.  K.  W.  (Hughes).  1964.  A  statistical 
study  of  variation  and  evolution  in  Mimulus 
gultatus.  Unpublished  M.S.  Thesis.  tTniver- 
sity    of   Utah,    Salt    Lake   City,    Utah. 

Edwards.   A.   W.   F..   .and   L.    L.    C.avalli-Sforza. 


1964.     Reconstruction    of    evolutionary    trees. 

Syst.  Assoc.  Publ.  6:67-76. 
Harman.  H.  H.      1964.       Modern  factor  analysis. 

University    of    Chicago    Press,    Chicago,    111. 

471   pp. 
Lindsay,    D.    W.     1960.     The    crossing    behavior 

and  cytogenetics  of  Mimulus  guttatus  Fisher 

in   the   Bonneville   Basin.     Unpublished   Ph.D. 

Dissertation.    University    of    Utah,    Salt    Lake 

City.   Utah. 
Prim,  R.  C.      1957.     Shortest  connection  networks 

and  some  generalizations.    Bell  Syst.  Tech.  J. 

36:1389-1401. 


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TABLE  OF  CONTENTS 

A  revision  of  the  Phacelia  Crenulatac  group  (Hydrophyllaceae)   for  North 

America.     N.    Duane   Atwood    127 

Rodent  populations,  biomass.  and  connnuiiitv  relationships  in  Arleniisin  tri- 
dentata.  Rush  Valley.  Utah.  D.  W.  Nichols.  H.  D.  Smith,  and  M.  F. 
Baker   l^^l 

Computerized  reduction  of  meteorologic  measurements  from  irrigated  and 
nonirrigated  plots  in  central  Utah.  Fenon  L.  Andersen  and  Paul  R. 
Roper 203 

Clarence   Cottam.    1899-1974:    a   distinguished   alumnus  of   Brigham   Young 

University.     Vasco   M.   Tanner   231 

Evolutionary  divergence  in  closely  related  populations  of  Mimulus  gut- 
talus  (Scrophulariaceae).  Karen  W.  Hughes  and  Robert  W.  Vickery. 
Jr 240 


IE  GREAT  BASIN  NATURAUS 


me  35  No.3    September  30, 1975 


Brigham  Young  Universit 


7    /^^  V 


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,  Zoology;  Stanley  L. 

Welsh,  Botany;  Clayton  M.  White,  Zoology. 
Ex  Officio  Editorial  Board  Members.  A.  Lester  Allen,  dean.  College  of  Biological  and 

Agricultural  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  continuously  published  from  one  to  four  times  a  year  since  then  by  Brigham 
Young  University,  Provo,  Utah.  In  general,  only  original,  previously  unpublished 
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Manuscripts.  All  manuscripts  and  other  copy  for  the  Great  Basin  Naturalist 
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The  Great  Basin  Naturalist 

Pubmshj:d  at  Pr()\  o,  Utah,  by 
Brigham  Young  University 


Volume  35 


September  50,  1975 


No.  3 


UROSAURUS  AND  ITS  PHYLOGENETIC  RELATIONSHIP  TO 

UTA  AS  DETERMINED  BY  OSTEOLOGY  AND  MYOLOGY 

(REPTILIA:   IGUANIDAE) 


Charles  Fangli 


D,i 


V.  Av, 


i.ul  Wihupi-  W.  T,i 


Abstract.-  -  Tlu>  sUilus  ol'  the  f;;eiius  rrosctunis 
Baird  and  Giiaid  as  piest'iitcd  in  i)ro\  ions  studit 
and  tln-oat  osteology  and  myology  are  included  ii 
ratios  detei'mined.  and  the  position  noted.  Five  dis 
Uta  and  Urosaurus.  and  it  is  concluded  that  both  of 
generic  status.   Vta  is  considered   to   be  phylogenetic 

The  climbing  utas  {Urosaurus)  and  the 
the  ground  utas  [Uta)  are  two  genera 
which  have  had  an  uncertain  sy.stematic 
relationship.  Studies  by  Mittleman  (1942), 
Savage  (1958),  Etheridge  (1962),  and 
Larsen  and  Tanner  (1975)  have  used 
various  sets  of  morphological  data  to  ex- 
plain their  affinities,  but  myological  char- 
acteristics have  been  mostly  neglected. 
Other  studies  involving  taxonomy  l)y  Bal- 
anger  and  Tinkle  (1973)  and  I'anner  and 
Jorgensen  (1963),  ecology  and  external 
anatomy  by  Smith  ( 1 946 )  \  Tinkle  ( 1 967 ) , 
Turner  et  al.  (1970),  and  Tanner  (1972), 
of  Uta  and  Urosaurus  and  other  related 
genera  have  been  done.  However,  the  va- 
lidity of  the  genus  Urosaurus  and  its  po- 
sition in  the  phylogeny  of  the  Igiianidae 
has  not  been  completely  established.  The 
present  study  is  designed  to  in^-estigate 
the  anterior  osteological  and  myological 
anatomy  of  these  genera  in  order  to  de- 
termine their  relationship  to  each  other 
and  to  show  their  j)hylogenetic  ])osition 
in  the  family  Iguanidae. 

The  literature  dealing  with  the  anatomy 
of  iguanid  lizards  has  been  reviewed  by 
Avery  aiul  Tanner  (1971);  therefore,  we 
will  confine  our  study  primarily  to  prob- 
lems relating  to  phylogeny.  Data  from 
previous  studies,  as  indicated  abo\e,  will 
be  added  to  our  myological   findings. 


|i  In  the  genus  Ula 
l.il.i  lidni  the  head 
muscle  is  measured, 
are  noted  between 
different  to  warrant 


Ilallowell  and  its  i-elalinnsh 
s    <ue    reviewed.    Additimi.il     , 

the  analysis.  Each  bone  and 
tinct    anatomical     differences 

these  genera  are  sufficiently 
allv   more   jirimitive. 

Baird  and  Girard  (1852 1  erected  the 
gonus  I 'la  for  the  species  stansburiana, 
which  was  characterized  by  gular  folds, 
auricular  openings,  and  a  fine,  homogen- 
eous dorsal  scalation.  Later  that  year  they 
described  another  form,  Uta  ornata.  which 
differed  from  stansburiana  in  having  the 
dorsal  scalation  com}X)sed  of  fine,  strongly 
keeled,  j)rominently  ind)ricated  scales 
which  were  (h^  ided  into  two  parallel  series 
on  either  side  of  the  median  dorsal  line 
by  a  series  of  somewhat  smaller,  vertebral 
scales.  However,  Hallowell  (1854)  en- 
countered a  different  lizard  similar  to 
Uta  ornata  Baird  and  Girard.  It  had 
enlarged  dorsals  extending  the  length 
of  the  dorsum  in  a  broad  and  unin- 
terrupted band  and  lacked  the  smaller, 
(H\i(ling  series  of  scales.  For  this  species 
Hallowell  established  the  genus  Urosaurus. 
Dumeril  (1856)  described  the  genus  P/z/- 
tnatob'psis  for  a  species  similar  to  Uta 
ornata  in  that  it  has  similar  dorsals  on 
either  side  of  the  smaller  \  ertebral  scales. 
These  actions  were  challenged  by  Baird 
(1858)  who  described  Uta  synunrtrica.  a 
(lose  relative  of  Uta  ornata;  and  the  next 
year  he  (Baird.  1859)  placed  Hallowell's 
Urosaurus  in  synonomy  with  Uta  because 
of  similar  dorsal  scalation  and  promi- 
nently characterized  gular  folds.  Urosau- 
rus was  used  as  a  subgenus  by  Van  Den- 


JDcpartiiient  of  Biology,  Branford  lligli  Scliool,  Bianfoid,  (  (iim 
-Department  of  Biolog\%  Southern  OmnccticiU  Stale  College.  Ni 
•'Life  Science  Museum,  Brigham  Young  Universil}-,  Provo,  Clah 


245 


246 


ASIN  NATI'RALLST 


Vol.  35,  No.  3 


burgh  (1922).  P/nniatolcpsis  was  placed 
in  synoiioniy  under  Uta  by  Cope  (18()4). 
Although  Fischer  (1882)  and  Boulenger 
(1883)  used  the  name,  it  was  again  jilaced 
in   synonomv   by  Boulenger    (1885). 

Cope  (18fr3)  described  the  distincti\e 
f'fa  tljcddssina  from  a  general  habitat  suf- 
liciently  different  from  other  known  forms 
that  Boulenger  (1885)  considered  it  gen- 
erica  lly  distinct  and  ])roposed  the  name 
Petrosaurus.  Boulenger's  usage  was  ig- 
nored by  Cope  (1887),  who  retained  Tha 
thalassina.  Excej^t  for  Van  Denbnrgh 
(  1922),  who  used  Petrosaurus  subgeneri- 
cally.  the  name  has  been  considered  a 
synonym  of  Uta. 

Mittleman  (1942)  reviewed  the  phylo- 
genetic  relationships  of  North  American 
iguanid  genera  and  considered  Ctenosaura 
( Wiegmann,  1828)  to  have  evolved  from 
a  jirimitive  iguanid  genus  and  to  have 
given  rise  to  two  phyletic  lines.  The  first 
contains  Crotaphytus  (Holbrook.  1842). 
and  the  other  scelojiorine  line  contains 
the  following  genera:  Uta  ( Baird  and 
Girard,  1852),  Urosaurus  (Hallowell, 
1854,  and  Sator  (Dickerson,  1919).  Uta 
and  Urosaurus  are  considered  to  ha^e 
evolved  from  an  early  progenitor  related 
to  Sceloporus.  which  soon  diversified  suf- 
ficiently to  produce  Uta  and  Urosaurus. 
The  latter  geiuis  is  ])robably  the  older  of 
the  two.  Uta  j^robably  did  not  chsperse  un- 
til the  beginning  of  the  Miocene,  for  it  is 
restricted  to  the  continental  United  States 
and  Mexico  and  many  of  the  adjoining 
islands.  According  to  Mittleman  (1942) 
Uta  and  Urosaurus  "...  may  be  con- 
sidered as  very  nearly  biological  ecjuiv- 
alents,  for  they  are  widely  distributed, 
highly  prolific,  of  about  the  same  age,  suc- 
cessful, and  derived  from  cIoscIn'  related 
progenitors." 

Sator  (Dickerson,  1919).  because  of  its 
relationship  with  Sceloporus,  is  also  of 
interest.  Although  Dickerson  ( 1919j  men- 
tioned certain  osteological  characteristics 
unique  to  Sator.  the  constant  osteological 
variations  wit  bin  the  genera  Sator,  Scelop- 
orus, Uta.  and  U rosaurus  have  not  been 
properly  ascertained.  Mittleman  (  I  9  1-2  i 
designates  Sator  as  a  direct  deri\ali\e  ol 
the  primitive  pyrocephalus  group  of  .SVv 
loporus  and  considers  it  to  hv  not  (  loscly 
related  to  any  other  lizard. 

Stejneger  and  Barbour  (1943i  and 
Smith  and  Taylor  (1950),  in  their  (  bee  k- 
list   of    lizards    of    the    U.S.    and    Mexico. 


adopted  Mittleman's  (1942)  arrangement 
of  the  "utas,"  while  Smith  (1946)  pre- 
sented a  somewhat  modified  phylogeny  of 
North  American  iguanid  s  that  is,  never- 
theless, in  basic  agreement  with  Mittle- 
man's work.  Since  Mittleman's  study, 
heri)etologists  have  been  divided  in  their 
acceptance  of  his  work  as  opposed  to  that 
of  Oliver  (1913).  who  did  not  recognize 
Mittleman's  genera  because  he  saw  few 
characters  separating  them.  Oliver  re- 
tained all  the  s])ecies  now  assigned  to 
either  TUa  or  Urosaurus  within  the  genus 
Uta.  Schmidt  (  1953)  and  Stebbins  (1954), 
among  others,  adhered  to  his  view.  Mittle- 
man's classification  was  based  upon  his  in- 
terjiretation  of  external  characteristics 
rather  than  upon  inarked  structural  dif- 
ferences between  the  se^•eral  species 
groups. 

Savage  (1958)  is  in  general  agreement 
with  Alittleman's  two  lines  of  jihylogeny: 
the  iguanine  line  and  the  sceloporine  line. 
Flowever,  genera  ])laced  in  the  sceloj:)orine 
line  by  the  two  authorities  do  not  agree. 
Savage  states  that  based  upon  the  type  of 
sternal  arrangement  there  are  two  major 
subdivisions  within  the  scelojiorine  grou]). 
Within  the  line  ha\  ing  a  utiform  sterum, 
two  distinct  stocks  are  indicated.  One  of 
lh(>NC>  is  represented  by  the  genus  Phryjjo- 
sonia.  which  lacks  xiphisternal  ribs.  The 
()tbc>r  group,  with  the  utiform  sterum 
bearing  xijihisternal  ribs,  contains  the 
genera  CaUisaurus.  Holbrookia.  Unia.  and 
Uta.  Within  Uta..  the  subgenus  Petro- 
saurus is  considered  by  Savage  to  be  the 
most  primitive,  although  highly  adapted 
for  a  rock  habitat.  The  genera  Urosaur- 
us. Sator.  and  Sceloporus  are  closely  allied 
,uid  differ  from  the  other  sceloporines  in 
ha\  ing  a  urosaurine  type  of  sternum 
which  possesses  xiphisternal  ribs.  Thus  in 
Savage's  studies.  I)ased  upon  the  type  of 
sternal  arrangements.  Uta  and  Urosaurus 
are  distinctly  different  genera. 

Presch  (19()9)  reports  that  the  osteo- 
logical characteristics  indicate  that  the 
horned  lizards  (Phrynosoma)  form  a 
highly  s])ecialized  genus  within  the  Sce- 
loporus  grou{)  of  genera  (Sceloporus.  Sa- 
lor.  Uta,  Urosaurus.  Uma,  CaUisaurus, 
llolhrookia.  and  Petrosaurus).  Phryno- 
soma is  distinguished  from  all  other  mem- 
bers of  \\\o  familv  in  having  both  a  large 
sternal  f()iit<niell(>  and  femoral  pores.  He 
lists  Pe/r(jsaurus  as  primitive  because  of  its 
incideratelv    sized    sternal     f()ntan(dle    and 


Sept.  1975 


FANOIIELLA.  ET  AL:    LI/.ARn  ANATOMY 


247 


the  presence  of  four  sternal  ribs.  Of  the 
two  groups  (leriA'ecl  from  Pctrosaui  us.  the 
least  altered  are  Urosaurus,  Uta^  Scclop- 
orus,  and  Sator,  which  have  hooks  on  the 
clavicle  and  a  covering  oxer  the  antero- 
lateral processes  of  the  frontal  in  some 
species,  while  Holbrookia.  Unia.  and  (\iUi- 
saurus  are  the  most  higlilA  ('\nlved.  They 
have  lost  the  lacrimal  and  postfrontal 
bones  and  the  first  ])air  of  cervical  ribs. 
Also,  the  interclavicle  is  shortened,  and 
the  anterolateral  processes  of  the  frontal 
are  covered.  Thus  Uta  and  Urosaurus  are 
placed  in  the  same  group  but  in  distinct- 
ly different  genera. 

Etheridge  (1964)  claims  tthat  osteo- 
logical  comparisons  do  not  {irovide  a 
strong  enough  argument  for  or  against 
the  recognition  of  Uta  and  Urosaurus  as 
separate  genera.  However,  the  few  osteo- 
logical  differences  between  sceloporines 
that  do  exist  suggest  that  three  sul)groups 
might  be  recognized:  (1)  Holbrookia..  Cal- 
lisaurus,  and  Uma  with  two  cervical  ribs 
(three  in  all  others);  (2)  Uta.  Urosaurus. 
Sator,  and  Sccloporus  with  cla\icular 
hooks  present  (absent  in  all  others);  and 
(3)  Petrosaurus  with  four  sternal  ribs 
(two  or  three  in   others). 

Hotton  (1955)  in  his  studies  of  den- 
tition and  food  habits  has  implied  that 
although  Uta  and  Urosaurus  are  inter- 
preted as  direct  but  independent  descen- 
dants of  sceloporines,  the  dentition  and 
diet  of  the  utas  are  similar  to  Callisaurus. 
Low^e  (1955)  studied  the  problem  of  gen- 
eric status  of  Uta  and  Urosaurus  using 
ecological  relationships.  He  w^as  able  to 
recognize  genera  on  the  basis  of  ecologic 
divergence  alone,  without  the  support  of 
any  other   character. 

On  this  ecological  concept  Lowe  and 
Norris  (1955)  based  their  classification 
of  the  assemblage  of  lizards  formerly 
placed  in  the  genus  Uta.  They  confirmed 
Mittleman's  arrangement  of  these  species 
because  of  supporting  ecological  differ- 
ences between  and  similarities  within  the 
groups  involved.  As  a  result  of  their 
studies,  they  recognized  the  following 
taxonomic  arrangement:  genus  Petro- 
saurus with  subgenus  Streptosaurus-  genus 
Uta  and  genus  Urosaurus. 

Petrosaurus  and  Streptosaurus  \^•ere 
placed  together  because  of  their  cliff- 
dwelling  habits.  Urosaurus  was  retained 
as  a  distinct  genus  because  the  species 
within  the  group  are  plant  dwellers  and 


clind)ers.  lUa  was  distinguished  from  the 
other  two  genera  \)\  \\<.  gi-()und dwelling 
I  de-style. 

MittlemaiTs  c  lassification  of  these  igua- 
nids  has  ]iot  been  generally  accej)ted  by 
Savage  and  others  Ixnause  he  failed  to 
present  lonxincing  e\  idence  that  the  sev- 
eral groups  were  mor})hologically  dif- 
ferent from  one  another.  The  most  strik- 
ing morphological  feature  listed  by  Mit- 
tleman  as  se]Kirating  Uta  from  Urosaurus 
was  the  homogeneous  scutellation  of  the 
former  and  the  differentiation  of  the  para- 
\ertebral  scales  in  the  latter. 

We  extend  our  gratitude  to  those  who 
have  helped  us  in  ihe  preparation  of  this 
pa])er.  We  are  grateful  to  Dr.  Ernest 
Williams,  at  the  Dei)artment  of  Herpe- 
tology,  Museum  of  ComJlarati^■e  Zoology 
at  Harvard,  for  providing  us  with  various 
])re])ared  skeletons  of  Uta  anrl  Urosaurus. 
We  also  thank  Mr.  (diester  .1.  Bosworth 
aiid  Dr.  Dwight  G.  Smith,  who  have  been 
so  kind  as  to  read  and  criticize  this  study, 
distribute  necessary  literature,  and  make 
suggestions.  We  are  grateful  to  Kenneth 
R.  Larsen  and  Wilmer  W.  Tanner  for 
making  available  a  copy  of  the  manuscript 
of  Larsen  and  Tanner  (1975).  Lastly  we 
thank  Southern  Connecticut  State  College 
for  financial  aiti  and  the  loan  of  materials 
and  s]:)ace  for  ])art  of  this  study,  and  Brig- 
ham  Young  LTniAersity  for  editorial  and 
l)ublication    courtesies. 

Ma'ikrials  and  Methods 

Skeletons  used  in  the  study  were  bor- 
rowed from  the  Museum  of  Comj)arative 
Zoology  at  Harvard  (MCZ),  and  ])reserved 
speciinens  were  borrowed  from  Southern 
Connecticut   State   College    (SCSC). 

One  skeleton  of  Urosaurus  ornata 
wrighti  and  three  of  Uta  s.  stanshuriana 
were  prepared  by  carefully  stripping  away 
the  skin,  connective  fascia,  and  large  mus- 
(le  the  first  (\i\\  of  skeletonizing.  After 
dr^'ing,  the  remaining  tissues  were  re- 
moved by  stri])])ing  and  picking  until  the 
skeletons   were   clean. 

All  measurements  were  taken  iji  milli- 
meters with  an  ocidar  micrometer 
mounted  in  a  chssecting  microscope.  All 
measurements  were  taken  from  the  ex- 
treme points  of  the  width  <nid  length  of 
each  structure. 

Specimens  are  accessioned  in  the  nat- 
ural   history    collection    of    MCZ    and/or 


248 


GREAT  BASIN  NATUHALIST 


Vol.  35,  No.  3 


SCSC.    The    materials 
study  are  as  follows: 


utilized     for 


Osteologv 
Uta   stansburiana   stansburiana    15airil    niid 
MCZ  62-141  I'tah 
SCSC   381.  East  of  Ell)erta.   t'tah 
SCSC  382,  East  of  Elberta.  l^tah 
SCSC  383,  East  of  Elberta.  Utah 
Urosaurus  ornata  syninielrica   (Baird) 
MCZ  26695,  Fort  Yuma.  Arizona 
Urosaurus  ornata  linearis  (Baird) 

MCZ  04947.  Tucson  Mt.  Park.  Arizona 
Urosaurus  ornala  lateralis   ( Boulenger) 

MCZ  14345.  Guaynias.  Mexico 
Urosaurus  ornata  scholli  (Baird) 

MCZ  64122.   Sonora.   Southern   Guavr 
Mexico 
Urosaurus  ornata  u-ri^hti  Schnhdt 
SCSC    384.   Moab.   ftah 


Uta 


Myology- 
stansburiana  stansburiana  Ba 


<1  C, 


SCSC  381.  L'tah  Counts 
SCSC  382,  Utah  Count' 
SCSC  383.  Utah  Count' 
Urosaurus  ornata  (Baird) 
SCSC  921.  Moab,  Utah 
SCSC  922.  Moab.  I^tali 
SCSC  923.  Moab.  T'tah 


Utah 
Utah 
I'tah 


Ostkolo(;y 

A  sIikIa  of  the  osseous  elements  of  Uta 
and  U rosdiiriis  re\  eals  a  basic  pattern  that 
was  described  by  Savage  (1958),  Ethridge 
(1964),  and  Avery  and  Tanner  (1971) 
for  thes(^  and  other  iguanids.  As  a  result 
\\e  confine  otir  descriptions  to  deviations 
from    that    pattern. 


Skull 


d  T 


An  auidysis  of  the  skull  and  jaw  was 
made  from  data  obtained  by  examining 
their  size  and  sh.ape.  After  skidls  were 
measured,  a  pei((>ntaoe  \\as  computed  be- 
tween length  and  width  and  compared 
with  similar  data  for  both  genera.  Mea- 
surements and  ratios  were  taken  for  iden- 
tical bones  in  both  genera.  Those  bones 
haAing  an  average  mean  greater  than  40 
points  are  jiresented  in  Tables  1  and  2. 
venience  of  reference  the  skull  has  been 
subdivided  into  a  posterior  occipital  unit 
and  <\\\  anterior  maxillary  unit. 

The     skulls     are     strej)tostylic     with    a 
freeh'  movable  (luadrate   bone  which  ar- 


T.\Bi.E    1. —  Mininuiui.   mean,  and  nuixinnmi  measurcnu'nts 
Uta. 

Length 
Name  of  structure  Min.  Mean  Max. 

Basisphenoid 1.22-1.35-  1,46 

Basioccipital  1.46-  1.75-2.14 

Pterygoid    3.95  -  4.58  -  5.12 

Ectopterygoid  2.20  -  2.26  -  2.'U1' 

Vomer  1 .76  -  1 .92  -  2.04 

Palatine  1.7!  -  1.92  -2.10 

Premaxilla   1.12-  1.86- 2. 4-!' 

Maxilla  \^.7^'  -  5.08  -  V.SO 

Nasal  - 1 .80  -  1 .97  -  2.01- 

Prefrontal  -...      2.39  -  2.62  -  2.88 

Lacrimal     487  -  .574  -  .681' 

Frontal    3.90  - -k07  -  k35 

Postorbital    1.07-  1.92-2.62 

Jugal   6.10-6.58-  7.22 

Parietal     3.24  -  3.86  -  4.10 

Postfrontal    487  -  .682  -  .926 

Squamosal     2.30-2.60     2.93 

Quadrate     735  ■  .81  i     .')75 

Supratemp.   Fossa  2.78  -  2.91     3.0  I 

Orbit    4.35-4.41  -  4.50 

Nasal  Opening  975  -  1.38  -  1.6! 

Dentarv    6.60-7.08  -  7.80 

Articular 3.16-  3.46  -  3.86 

Angular   Pr 5.70  -  6.89  -  9.50 

Surangular  2.20  -  3.17  -  4.1  :• 

Splenial     2.20-2.42  -2.58 

Angular    5.70  -  6.89  -  9.50 

Coronoid   2.20  -  3.23  -  t.40 

Pvriform  Becess    2.68  -  3. 1  7     3.42 

Parasplienoid  Pr 925-  1.2!      1.46 

Entu-e  Skull  11.2-11.6-11.9 


,kuil 


uctures    of 


Mi 


Width  Width-length  ratio 

.  Mean  Max.       Min.  Mean  Max. 


1.7i 

2.20 

1.27 

1.61 

.780 

.975 

1.1-6 

1.9", 

.486 

.490 

.390 

3.66 

1.66-  1.7') 

.440-  .610 

5.15-5.66 

.098  -  .  1 95 

.68')     .8  11 


1.83- 
2.46 
1.62- 
1.77- 
.830 
1.18 
1.67  - 
1.00 
.658  - 
1.41  • 
.57  3- 
1.02 


.58() 

.140 
.10-, 
1.13 
1 .46 
2.04 
.487 
6.35 


1.1  1 
.720 
6.07 
.',98 
.I'^l 
1 .88 
1.67 
2  17 

~m 

7.26- 


1.9', 
2.78 
2.30 
1.9', 
.880 
1.22 
2.21- 
2.04 
.975 
1 .90 
.732 
4.6  1- 

.7  32 
5.00 
.20  3 
1.07 
2. 1 1 

3. 17 
1.42 
1.22 
.880 
7.\-^ 
,780 
,585 
2.30 
2.01- 
2.1 1 
.487 
8.05" 


.715 
.610 
.274 
.700 
.100 
.500 
.600 
.352 
.239 
.202 
.072 
.8  12 
,',',0 
,070 
,()',() 


.10  1 
.720 


,388 
,700 
331 
566 


,/6/  -  77 1 
,701  -  .770 
.  354  -  .445 
.704 -.885 
.431  -  .472 
.579  -  .687 
.733-1.00 
.396 -.41 6 
.3  32 -.476 
.433-  .710 
.101  -.150 
,003-  ,940 
,711  -  .925 


.092 
,680 
288 


H) 


.101- 

,  H)0 
.  V)0 
,410 
.505 
,762 -.820 
.770  -  .995 
.163 -.185 
.207  -  .243 
.824  -  .855 
.185  -.214 
.  1 74  -  .226 
.280  -  .334 
.560  -  .926 
.722 -.765 
.424  -  .525 
.623  -  .675 


Sept.  1975 


Table  2. —  Minimum,  nu\ 
saurus. 


FANGHELLA.  ET  AL:   LI/ARD  ANATOMY  249 

u  and  mnxinunn  measiurements  and  ratios  for  tlie  skull  structures  of  Uro- 


Length  Widtli 

Name  of  structure  Min.  Mean  Max.        Min.  Mean  Max. 

Basisphenoid     1 .56 

Basioccipital   1 .22 

Pterygoid 4.00 

Ectoptervgoid  1 .41 

Vomer  1 .22 

Palatine  \  .46 

Premaxilla    1,22 

Maxilla  5.26 

Nasal     2.04 

Prefrontal  2.74 

Lacrimal  487 

Frontal    3.80 

Postorbital    2.44 

Jugal   5.37 

Parietal  3.90 

Postfrontal    585 

Squamosal    2.44 

Quadrate     930 

Supratemp.  Fossa   2.78 

Orbit    3.90 

Nasal  Opening    1.49 

Dentary    6.84 

Articular     3.26 

Angular   Pr 5.70 

Surangular     2.68 

Splenial     2.20 

Angular    5.70 

Coronoid     2.68 

Pvrifomi  Recess   2.44-2.76-3.18  1.76 

Parasphenoid  Pr 925-1.25-1.42         .440 

Entire  Skull  11.2-11.6-12.4         6.75 


Width-lengtli  ratio 
Min.  Mean  Max. 


1.71  -  1.95 
1.57-  1.80 
4.74-5.37 
1.84-2.24 
1.76-2.-M. 
1.84-2.20 
1.44-1.56 
5.84-  6.10 
2.32  -  2.78 


2.91- 

.615- 

4.15- 

2.75- 

6.00- 

4.08- 

.791  - 

2.55- 

1.07- 

3.14 

4.59- 

1.52- 

7.15- 

3.44 

6.30-  7.15 

2.86  -  3.26 

2.53  -  2.92 

6.30-7.15 

2.96-3.18 

2.76-3.18 

1.25-  1.42 

11.6-  12.4 


3.12 
.737 
4.40 
3.18 
6.35 
4.30 
.975 
2.93 
1.32 
3.36 
4.90 
1.61 
7.60 
3.76 


1.90 
2.68 
■1.17 
1.17 
.733 
.975 
1.71 
2.20 
.830 
1.07 
.487 
4.40 
1.71 
.540 
5.35 
.024 
.780 
2.20 
1.32 
3.42 
.585 
.810 
.974 
5.70 
.684 
.2-H 
1.45 


-2.01 

-  2.89 
-1.32- 
-1.50 
-.811  - 

-  1 .24 

-  2.23  - 

-  2.21  . 

-  .946  - 

-  1  22 

-  .615  - 
4.60 
1.86- 
.635 
5.64 
.190 
.940- 
2.34 
1.54 
3.68 
.975 
.889 
1.39 
6.77 
.794- 
.478 
1.03 
1.73 
2.08- 
.597 
7.38 


2.14 
3.17 
1.46 
1.80 
.930 
1.71 
2.58 
2.34 
1.07 
1.36 
.732 
4.87 
1.95 
.732 
5.96 
.-440 
1.07 
2.64 
1.76 
3.90 
1.22 
.928 
1.85 
8.10 
.880 
.585 
1.90 
1.95 
2.44 
.925 
7.80 


.800 
.455 
.280 
.655 
.340 
.445 
.565 
.360 
.360 
.344 
.073 
.780 
.538 
.088 
.661 
.024 
.327 
.405 
.430 
.720 
.388 
.118 
.300 
.705 
.225 
.084 
.252 
.400 
.635 
.360 
.582 


.872-  1.00 
.555  -  .675 
.304 -.366 
.753  -  .880 
.488  -  .680 
.848  -  1 .00 
.656  -  .730 
.405  -  .437 
.4 10 -.464 
.419  -  .462 
.286-  1.00 
.902  -  .990 
.688  -  .800 
.106-. 127 
.723  -  .765 
.255  -  .500 
.370  -  .440 
.460  -  .590 
.448  -  .525 
.806  -  .875 
.656  -  .897 
.124-  .131 
.399  -  .507 
.875-1.00 
.271  -  .328 
.156 -.244 
.284 -.333 
.591  -.726 
.760 -.910 
.474  -  .650 
.633 -.670 


ticulates  dorsall}'  with  the  paroccipital 
process  and  ventrally  with  the  quadrate 
process  of  the  pterygoid.  Thus,  they  form 
a  compact,  light,  and  strong  cage  for  the 
brain  and  sense  organs. 

The  actual  shape  of  the  skull  is  either 
elongated  and  flattened  dorsoventrally,  as 
in  Uta,  or  shortened  and  lateraHy  com- 
I)ressed,  as  in  Urosaurus.  Measurements  of 
the  length  of  the  skull  were  from  the  top 
of  the  premaxillary  bone  to  the  most  pos- 
terior extension  of  the  occipital  condyle. 
Measurements  of  the  width  were  from 
the  widest  extension  between  the  sub- 
orbital bars  in  the  area  of  the  orbit. 

The  means  in  Tables  1  and  2  indicate 
that  Uta  has  a  slightly  lower  skull  ratio 
(.623)    than   Urosaurus    (.633).   For  con- 

The  occipital  jiortion  forms  a  median 
wall  for  the  attachment  of  the  neck  and 
articulation  for  the  remainder  of  the  skull. 
It  consists  of  two  parts:  (a)  braincase 
(basisphenoid,  basioccipital,  prootic,  exoc- 
cipital,  supraocci])ital)  (b)  foramen 
magnum  (enclosed  by  the  basioccipital, 
exoccipitals,  and  supraoccipital).  A  tripar- 
tate  occipital  condyle  is  located  on  the  pos- 


terior end  of  the  basioccipital  and  the 
lateral  exocci])ital  in  all  iguanine  genera. 

Basisphenoid:  Length  is  from  the  suture 
between  basisj)henoid  and  basioccijntal, 
to  the  beginning  of  the  paras])henoid  ])ro- 
cess  (Fig.  1).  Wichh  is  the  distance  be- 
tween the  widest  expansion  of  the  basio- 
{)tygoid  ])rocesses.  The  lowest  ratio  mean 
is  in  Uta  {.767),  the  highest  in  Urosaurus 
(.872).  A  low  ratio  indicates  that  the 
bone  is  much  longer  than  it  is  wide, 
whereas  the  higher  ratios  indicate  bones 
with  lengths  and  widths  more  nearly 
equal. 

Basioccipital:  Length  is  from  the  suture 
between  the  basisphenoid  and  basioccipital 
to  the  posterior  tip  of  the  occipital  condyle 
(Fig.  1);  and  width  is  between  the  tips  of 
the  lateral  extensions  of  the  sphenoccipi- 
tal  tubercles.  The  ratio  in  Uta  is  .701  and 
Urosaurus  .555. 

Pterygoid:  Length  (Figs.  1,  2,  3)  is  be- 
tween the  anterior  [portion  of  the  i)terygoid 
where  it  sutures  with  the  palatine  and  the 
most  j)osteri()r  tij)  of  the  quadrate  process. 
Width  is  between  the  articulation  with 
the  basipterygoid  process  of  the  basisphen- 


250 


GREAT  BASIN  NATURALIST 


Vol.  35,  No.  3 


BO — Basioccipital 

BS — Basisphenoid 

EC — Ectopterygoid 

FE — Fenestra  e.xonarina 

FEO — Fenestra  e.xochoanal 

FR— Frontal 

FVE — Fenestra  vorner- 

onassalis  e.xterna 
JU—Jugal 


MX— Maxilla 

NA— Nasal 

OB   -Orbit 

PAL — Palatine 

PAR— Parietal 

PF-  -Pineal   foramen 

PM-  -Premaxilla 

POT— Postorbital 

PP   -Parasplienoid   process 


PR     Pyriforni  recess 

PRF— Prefrontal 

PT— Pterygoid 

PTF— Postfrontal 

QU— Quadrate 

STF — Supratemporal   fossa 

SQ — Squamosal 

VO-  Vomer 


Fig. 


Dorsal    and    ventral    views   of   skulls.   A  ,ind    C:    l' la.   B    and    1)    / 


Sept.  1975 


FANGHELLA,  ET  AL:    LI/.ARD  ANATOMY 


251 


AR — Articular  JU — Jugal 

CO — Coronoid  LA- -Lacrimal 

DE— Dentary  MX— Maxilla 

EC- -Ectoptorygoid  NA — Nasal 

EP— Epipterygoid  OB— Orbit 

FE--  PVnestra  exonarina  FM — Premaxilh 

FR— Frontal  PO   -Postorbital 

Fig.  2.     Lateral  view    of    skull    aiul 
saurus. 


ial    view    of    niandib 


PRE— Prefrontal 
PR— Parietal 
PT— Pterygoid 
PTF— Postfrontal 
QU— Quadrate 
SP— Splenial 
SR     Surangular 

A    and    C    f'!a. 


and     I)    Uro- 


252 


GREAT  BASIN  NATURALIST 


Vol.  35,  No.  3 


CC — Constrictor  colli 
EP — Episternocleidoniastoideus 
GE — Genioglossus 
lAP — Intormandibularis 
anterior  profundus 


IAS — Intermandibularis  an- 
terior superficialis 

NP — Intermandibularis  pos- 
terior 

MHI— Mandiliulohyoideiis  I 


OM — Omohyoideus 
PE— Pectoralis 
SH — Sternohyoideus 
ST — Sternothyroidcu: 


Fig.  3.     Ventral   view   of   thioat   musculature;    superfi 
Uta;    B.    Urosaurus. 


'pth 


left    aud    fi 


;pth 


•igh 


oid  and  the  suture  with  the  ectopterygoid. 
The  ratio  is  Urosaurus  .304  and  Uta  .354. 

Ectopterygoid:  Length  (Figs.  1,  2)  is 
between  the  suture  with  the  pterygoid  and 
the  suture  with  the  jugal  and  maxilla. 
The  greatest  (Hameter  is  at  its  point  of 
union  with  the  jugal  and  maxilla.  The 
lower  ratio  is  in  Uta  (.704)  and  the  higher 
in  Urosaurus  (.753). 

Vomers:  Length  (Fig.  1 )  is  from  the 
anterior  suture  with  the  premaxilla  to  the 
most  posterior  point  of  the  suture  with  the 
palatine.  Width  is  between  the  median 
border  of  the  vomer  at  the  ventral  mid- 
line and  the  most  lateral  border  where  it 
attaches  to  the  maxilla.  The  ratio  in 
Urosaurus  is  .488  and  in  Uta  .431.  The 
vomers   possess   a    small    blunt    projection 


which  jirotrudes  from  its  lateral  border 
into  the  ojiening  of  the  fenestra  exocho- 
analis  and  fenestra  vomeronasalis  externa 
and  divides  the  opening.  This  anterolateral 
projection  is  seen  in  both  genera. 

Palatine:  Length  is  from  the  anterior 
suture  with  the  \()mer  at  the  midline  to 
the  most  posterior  extension  of  the  suture 
with  the  pterygoid  (Fig.  1).  Width  is 
from  the  skull's  midline  to  the  lateral  su- 
ture between  the  palatine  and  the  maxilla. 
The  ratio  in  T^rosaurus  is  .848  and  in  Uta 
.57<». 

Pn/nan//ae:  Length  (Figs.  1,  2)  is 
from  its  anteroventral  tip  to  its  dorsal 
union  with  the  nasal  at  the  dorsal  midline. 
Wi(hh  is  between  th{>-  lateral  sutures 
shared  b\    the  |)reniaxillae  with  the  maxil- 


Sept.  1975 


FANGHELLA.  ET  AL:    LIZARD  ANATOMY 


253 


la  on  the  ventral  surface  of  the  jiremaxilla. 
The  ratio  in  Uta  is  .7  3^  and  in  T^rosaurus 
.656. 

Maxillac-.l.ewgXh.  (Figs.  1.  J)  is  from 
the  most  anterior  extension  of  the  j)re- 
maxillarv  process  to  the  })osterior-most 
extension  of  the  maxillae  where  it  sutured 
with  the  jugal  and  ectopterygoid.  Width 
is  the  vertical  distance  from  the  ventral 
border  of  the  maxillae  to  the  dorsal-most 
extension  at  the  ]ioint  of  suture  with  the 
nasals  and  prefrontals.  The  ratio  in  Via  is 
3.96  and  in  Urosaurus  .405. 

Nasal:  Length  (Figs.  1,  2)  is  from 
the  tip  of  the  ventral  border  as  it  forms 
the  fenestra  exonarina  to  the  posterodor- 
sal  extension  that  sutures  with  the  jire- 
frontal.  Width  is  from  its  medial  suture 
with  its  opposite  member  to  its  most  lat- 
eral extension  where  it  sutured  with  the 
maxilla  and  prefrontals.  The  ratio  in  T'ro- 
saurus  is  .410  and  in  Uta  .332. 

Prefrontal:  Length  (Fig.  1)  is  from 
the  suture  between  the  ])refrontal  and 
lacrimal  bones  at  the  anterior  lip  of  the 
orbit,  to  the  suture  between  the  prefroiital 
and  frontal.  Width  is  from  the  suture  be- 
tween the  prefrontal  and  lacrimals  to  the 
median  point  wdiere  the  frontal,  nasal, 
and  prefrontal  bones  suture  together.  The 
ratio  in  Uta  is  .433  and  in  Urosaurus  .419. 

Lacrimal:  Length  is  from  the  antero- 
dorsal  border  as  it  sutures  with  the  pre- 
frontal and  maxilla  to  the  posterior  border 
on  the  rim  of  the  orbit  as  it  sutures  with 
the  jugal  (Fig.  2).  Width  is  the  distance 
between  the  dorsal  border  of  the  lacrimal 
at  the  rim  of  the  orbit  to  its  ^'entral  border 
at  its  suture  wdth  the  maxilla.  The  ratio  in 
Uta  is  .101  and  in  Urosaurus  .286. 

Frontal:  Length  (Figs.l,  2)  is  from  the 
most  anteromedian  suture  shared  with  the 
parietal.  Width  is  between  the  most  lateral 
j)osterior  projections  which  suture  with 
the  parietal  and  })ostfrontal.  The  ratio 
in  Uta  is   .903  and  in  Urosaurus  .902. 

Postfrontal:  Length  (Figs.  1.  2)  is  the 
extremities  of  its  longest  axis.  Width  is 
the  distance  between  the  parallel  borders 
on  the  axis  at  right  angles  to  the  length. 
The  ratio  in  Urosaurus  is  .255  and  in  Uta 
.288. 

Jugal:  Length  (Figs.  1,  2)  is  between 
its  most  anterior  projections  as  it  sutures 
with  the  lacrimal  and  maxillae,  to  the 
posterior  j)rojection  which  sutures  to  the 
anteroventral  border  of  the  postorbital. 
Width   is   the   distance   between   the   two 


parallel  borders  at  right  angles  to  the 
length.  The  ratio  in  I'ta  is  .092  and  in 
Urosaurus   .106. 

Parietal:  Two  measurements  (Figs.  1, 
2)  were  taken.  The  anterior  two-thirds  of 
the  bone  was  subjected  to  length-width 
measurements,  with  the  length  being  the 
distance  along  the  midline,  from  the  an- 
terior suture  with  the  frontal  to  the  suture 
between  the  ]iarietal  and  the  supraoccipi- 
tal.  Width  is  the  distance  between  the  two 
anterolateral  ])rojections  that  suture  with 
the  ]iostorbital  and  jiostfrontal.  The  ratio 
in  Urosaurus  is  .723  and  in  Ufa  .680. 

Postorbital:  Length  (Figs.  1,  2)  is 
between  the  anteroventral  and  postventral 
projections.  Width  is  from  the  ^•entral 
border  to  the  tij)  of  the  dorsal  ]irojection 
where  it  sutures  with  the  ])arietal  and 
postfrontal  bones.  The  ratio  in  Uta  is  .711 


md 


rosaurus 


.688. 


Squamosal:  Length  ( Fig.  1  )  is  between 
the  most  anterior  and  ])osterior  extrem- 
ities. Width  is  between  the  parallel  bor- 
ders on  an  axis  at  right  angles  to  the 
length.  Ratio  in  Urosaurus  is  .370  and  in 
Uta  .^25. 

Quadrate:  Length  (Fig.  2)  is  from 
its  dorsal  border  where  it  attaches  to  the 
squamosal  and  the  \('ntral  extremity  of 
the  condyle  which  articulates  with  the 
articular.  Width  is  between  its  medial 
and  lateral  borders.  The  ratio  in  Urosaurus 
is  .460  and  in  Uta  .^46. 

Supratemporal  fossa:  Its  length  (Fig.  1) 
is  the  inside  distance  on  the  longest  axis 
and  width  the  inside  distance  on  the  long- 
est axis  at  right  angles  to  the  length.  The 
ratio  in  ['ta  is  .  1()7  and  in  Urosaurus  .488. 

Orbit:  Length  (Figs.  1,  2)  is  between 
the  lacrimal  and  jiostorbital.  Width  is  be- 
tween jugal  and  frontal  bones  .The  ratio 
(most  circular  o])ening)  in  Urosaurus  is 
.806  and  (most  (41i]itical  opening)  in  Uta 
.762. 

Feru'stra  e ronarina:  Length  (Fig.  1) 
is  the  nitenial  distance  between  the  lateral 
jirojection  of  the  |)remaxilla  and  maxilla 
and  the  sutiu'e  between  the  nasal  and 
maxilla.  Width  is  the  inside  distance  be- 
tween the  lateral  border  of  the  premaxilla 
and  the  anterior  border  of  the  maxilla.  The 
ratio  (most  circular  opening)  in  Uta  is 
.770  and  (most  elliptical  opening)  in 
Urosaurus  .65(). 

The  lower  jaw  consists  of  two  paired 
rami  united  anteriorly  in  a  mental  sym- 
])h\  sis.   Each   articidates   posteriorly  with 


254 


GREAT  BASIN  NATURALIST 


Vol.  35,  No.  3 


the  quadrate.  The  dentarv  of  each  ramus 
bears  a  single  row  of  pleurodont  teeth, 
whereas  the  remaining  bones  (articular, 
surangular,  angular,  splenial,  and  coro- 
noid)   are  edentate. 

Dentary.  Length  is  from  the  anterior  tip 
to  the  posterior-most  projection  on  the 
lateral  surface  of  the  mandible  (Fig.  2). 
\\'i(hh  is  the  Acrtical  distance  bewteen 
the  to])  and  the  bottom  of  the  mandible, 
immediately  in  front  of  the  coronoid.  The 
ratio  in  Uta  is  .163  and  in  Urosaurus  .124. 

Articular:  Length  (Fig.  2)  is  from 
it  most  anterior  projection  on  the  median 
surface  where  it  sutured  to  the  coronoid 
and  splenial  to  the  most  posterior  tip  of 
the  retroarticular  process.  Width  is  from 
the  most  ventromedial  projection  of  the 
angular  ]:)rocess  to  the  opposite  border  of 
the  articular  where  it  sutured  with  the 
surangular  on  the  lateral  surface.  The 
ratio  in  Urosaurus  is  .399  and  in  Uta  .207. 

The  angular  process  of  the  articular 
bone  differs  in  shape  and  size  in  each 
genus.  This  projection  was  also  subjected 
to  length-width  measurements.  The  length 
is  the  greatest  length  of  the  mandible  and 
was  contrasted  with  the  width  of  the  artic- 
ular, which  in  part  is  a  result  of  the  size 
of  the  angular  process.  Urosaurus  .875  has 
the  greatest  ratio  (shortest,  widest) 
while  the  smallest  (longest,  narrowest)  is 
in  Uta  .824. 

Surangular:  Length  is  the  longest  an- 
terior-posterior axis  on  the  lateral  surface 
of  the  mandible  (Fig.  2).  Width  is  the 
longest  dorsal-ventral  axis  in  the  area  of 
the  anterior  sutures  wdth  the  dentary  and 
coronoid  on  the  lateral  surface.  The  ratio 
f shortest,  widest)  is  in  Urosaurus  .271  and 
in  Uta  .185. 

Spenial:  Length  is  the  longest  anterior- 
posterior  axis  and  the  greater  dorsal-ven- 
tral axis  is  the  width.  The  ratio  (shortest, 
widest)  is  in  Uta  .174'  and  in  Urosaurus 
.156. 

Angular:  The  angular  is  roughl}'  fusi- 
form; its  length  is  between  the  most  an- 
terior and  most  posterior  projections. 
Width  is  between  the  opi)osit(>  borders  on 
an  axis  at  right  angles  to  \\\v  length.  The 
ratio  (shortest,  widest)  in  Urosaurus  is 
.284  and  in  Uta  .280. 

Coronoid:  Length  is  ftoin  the  dorsal 
tip  of  the  bone  to  tbe  tip  of  the  ventral- 
most  projection  on  the  later-al  surface 
(Fig.     2).     Width     is     betw(MMi     anterioi- 


and  ])osterior  borders  where  they  contact 
the  dorsolateral  surface  of  the  mandible. 
The  ratio  in  Urosaurus  is  .591  and  in  Uta 
.560. 

Ah()LO(A- 

To  a\()id  (onfusion,  the  terminology 
used  lor  the  following  descriptions  of  the 
muscles  is  that  of  Robison  and  Tanner 
(1962),  .Jenkins  and  Tanner  (1968).  and 
Avery  and  Tanner  (1954,  1971).  The 
musculature  also  follow^s  the  basic  iguanid 
pattern  described  h\  the  aboAe.  Only 
deviations  will  be  noted  in  the  test. 

Throat  Muscidature 

M .  /u/crniandibularis  anterior  superfic- 
ialis  is  tonstant  in  both  genera  examined 
with  the  following  exception:  slightly 
broader  in  Urosaurus  than  Uta  (Fig.  3). 
However,  in  both  genera  the  muscle  is 
sheetlik(>  with  the  width  at  least  half  the 
length. 

M.  Iut(-rniandil>ularis  anterior  profun- 
dus is  relatively  consistent  in  its  location; 
however,  in  Uta  it  is  a  wide  band  of  mus- 
cle attached  to  the  intermandibularis  an- 
terior superficialis.  In  Urosaurus  it  is  a 
thin  sheet  separated  anteriorly  from  the 
intermandibularis  anterior  superficialis 
and  posteriorly  from  the  intermandibularis 
posterior  by  a  thin  membrane  (Fig.  3). 

M.  Intermandibularis  posterior  is  con- 
tinuous posteriorly  with  the  constrictor 
colli  from  wlii(  h  it  can  be  delineated  by  a 
natural  sej)aration  of  the  muscle  fiber 
bundles  (Figs.  3,  9  and  10).  The  posses- 
sion of  this  separation  is  varied  in  the 
genera  examined.  In  Urosaurus  the  con- 
strictor colli  and  intermandibularis  j)os- 
lerior  are  (■l()sol^■  associated  along  their 
entire  (onnnon  border.  In  Uta  the  two 
muscles  are  separated  totally  laterally  but 
are  continuous  for  a  short  distance  near 
the  midlin(>  raphe. 

M.  Mandihulohyoideus  I  in  Uros(nirus 
has  ap[)r()\iinatcly  one-half  of  its  body 
covered  l)\  the  omohyoideus.  whereas  in 
the  Uta  ()nl\  a  small  posterior  ])ortion  is 
covered    i  b'ig.    >  ) . 

M .  Ma/idil>ul(jlnoideus  II  ^vas  described 
l)V  AM'r\-  and  Tanner  (1971)  for  other 
ignanids.  I  l(t\\('\ cf.  we  were  iniable  to  lo- 
<alc  this  nniN(l«>  in  (Mther  Ufa  or  Urosaur- 
us^. 

M.  Mfnidihuinhyoideus  III  in  both  Uta 
and     Urosaurus    arises    from    the    ventre- 


Sept.  1975 


FANGHELLA.  ET  AE:    EIZARD  ANATO!\IY 


255 


medial  surfaces  of  the  dentary  and  angu- 
lar between  the  anterior  and  posterior 
mvohoid  foramina  (Fig.  4).  Tiio  narrow 
insertion  is  on  the  lateral  surfaic  of  the 
ceratohyal  distal  to  its  midpoint. 

M.  Genioglossus  is  a  thick  bandlike 
muscle  in  both  generji  which  occupies  a 
large  area  between  the  mandibular  rami 
(Figs.  3  and  4).  Its  position  is  ventral  to 
the  tongue  and  anterior  to  the  basihyal. 
The  first,  second,  and  third  mandibulo- 
hyoideus  and  the  intermandibnlari^ 
muscles  are  all  dorsal  to  it. 

M.  Hyoglossus  is  as  described  in  ollu>r 
iguanids. 

M.  Branchiohyoideus  in  Uta  (as  in  Sau- 
romalus)  has  a  narrow  insertion  on  the 
first  ceratobranchial,  whereas  in  Urosaur- 
us  the  insertion  covers  over  half  the  (hstal 


portion  of  the  first  ceratobranchial    (Fig. 
4 ) . 

M.  stcrnohyoidcus.  as  reported  in  the 
literature,  is  subject  to  considerable  con- 
fusion concerning  its  limits  (Figs.  3,  4, 
6).  Davis  (1934:19)  considers  the  super- 
ficial layer  to  be  divisible  into  three  parts 
in  (^rotaphytus.  One  of  these  muscles  he 
calls  the  omohyoideus.  Robison  and  Tan- 
ner (1962:6)  consider  this  muscle  con- 
tinuous in  the  same  genus.  Oelrich 
(1956:51-52)  treats  this  muscle  in  Cteno- 
saura  as  being  continuous,  but  owing  to 
the  different  origin  and  direction  of  the 
fibers  he  separates  the  layers  into  omo- 
hyoideus and  sternohyoideus.  Kesteven 
(1944:245-246)  studied  the  agamid,  Phy- 
signathus,  suggesting  a  separation  in  young 
sj)ecimens  and  treats  these  layers  as  con- 


BH — Branchiohyoideus 
GE — Genioglossus 


Fig.  4.     Ventral   view   of   throat   miisculaturr 
A.    Uta;    B.    Urosaurus. 


256 


(,KI.\r  BAMN  NAirUAl. 


Vol.  35,  No.  3 


sisting  of  three  patis  which  he  considers 
to  represent  the  similar,  though  distinct, 
divisions  present  in  \'ar(inus.  In  the  igua- 
nines  Avery  and  Tanner  (U)71i  treated 
the  sternohyoid(His  complex  as  three  sej)- 
arate  muscles,  sternohvoideus,  sterno- 
throideus,  and  omohyoideus.  This  arrange- 
ment is  followed  here.  In  both  genera  ex- 
amined, the  sternohyoid  ens  forms  a  broad 
elongated  sheet  of  muscle  covering  the 
posterior  portion  of  the  mandibulohyoideus 
I  muscle. 

\1.  omohyoideus  is  sheetlike  and  forms 
the  lateral  extension  of  the  sternohvoideus 
complex  (Figs.  3,  5,  10).  In  both  genera 
it  originates  medially  from  the  lateral  tip 
of  the  transverse  ]irocess  of  the  interclavi- 
cle  with  some  fibers  of  the  episternocleido- 
niastodeus.     Laterallv,    the    omohyoideus 


originates  from  the  anterolateral  surface 
of  the  cla\  i(  Ic  and  anterior  border  of  the 
su])rascapnla.  Its  fibers  pass  obliquely  an- 
terior to  insert  on  the  posterior  margin  of 
the  first  ceratobraiK  hial  and  the  ])roximal 
end  of  the  second  c cratohranchial  cartil- 
ages. 

In  both  geiiera  the  niecHan  border  is 
separated  from  the  lateral  border  of  the 
sternohvoideus.  7'he  delineation  of  both 
muscles  must  b(^  made  l)y  comparing  the 
origins  and  insertions.  In  U rosaurus  it  is 
easil^•  se])arated,  as  the  fibers  of  this  mus- 
c  ie  pass  oblicjue  to  those  of  the  sterno- 
hvoideus covering  most  of  the  mandibulo- 
hyoideus I.  In  Uta  it  is  a  thin  band  just 
lateral  to  the  sternohvoideus. 

M.  Stcrnotliyroidcus  is  the  most  medial 
extension  of  the  sternoh}-oideus   complex 


Ix- 7*— , 


C;L     Claviclp 
IC-  -Interclavicle 
LX — Larynx 

Fig.   5.     Ventral   view 
A.   I'to;    B.    IJrosaurus. 


oM    {)nniii\.ii(i(Mis  rr. 

I'M      I'h.nvn-..,,!    uu-mUam- 
PT      Pl(MVK()i(lin,in.lihul,ins 

•i\\    iiius<  iihitin-o;    foiirlh      <iiM)tli     at      Ic 


fif'th     (Irptli     at     right 


Sept.  197-5 


I  A  \(,1II,L1.\. 


\L:    LI/.AHI)    \.\  AIO.MV 


257 


Neck    Musdihilurc 
U.  Con^lru  tar  colli  is  xni-iiihlc  in  width 


iiiholh     j^cncfii      i 
widest.    ('()V<"riiio 
face  ..I    the  iKH  k. 
(■()nsti-i(  lor  (olli   ( 
M. 


and  can  be  separated  Irdiii  llie  otiiei-  mem- 
l^ers  of  the  group  l)\  its  different  origin 
and  insertion  (Fig.  ^i.  Ihe  name  st(n-no- 
thyroideus  is  used  as  in  (-<nnp  i  l')i);  1 ')  1  i. 
who  figured  it  as  the  dee|)  mend)er  of  the 
complex   in   BrachyUjplius. 

The  origin  is  considered  to  he  those  fi- 
bers arising  from  the  iutercla\  i(  le  and 
sternum.  These  fibers  pass  anterior! \  <nid 
parallel  to  the  trachea  to  insert  on  the 
liyoid  at  the  point  of  union  bet^^c>en  the 
basihvoid    and    hypohyal. 

In  Urosaurus  the  lateral  bordc>r  of  the 
sternothyroideus  and  the  mechau  l)ord(M- 
of  the  sternohyoideus  are  difficidt  to  de- 
termine. In  Uta  their  sei)aratiou  is  chs- 
tinguishal)le  since  the  three  muscles  have 

fibers  obliciue  to  one  another  as  described       scured    b\    the   more    superfic 
for    Saurornnlus    by    A^■er^'    and    Taimer       tor  colli. 
(1971).  M.  Levator  scapulae  superficiaHs  is  not 


is.  \.  t.  9).  it  IS 
s|  of  the  later-al  siir- 
/  /<)^(/urus.  A  nari"o\N 
us  HI  /  '/a. 
h'.pistcriiix  lctil(tin(i.\t<ii<l<us  was 
found  .IS  ,1  thin  band  of  muscle  extenchng 
o\  er  the  shouldc-r  m  lid.  whereas  in  I  '  ro- 
sdurus  it  is  ill  a  gt'e.itef  depth  (Figs.  ^,  7, 
M.     10.    and     111. 

M.  /J(/)ns\(jr  iiKindihuldris  (  b'igs.  (i.  9, 
10)  is  (h\i(lecl  into  three  l)undles  as  clo- 
se ribc^d  by  A\ery  and  Tanner  (1971). 
rhc>  third  bundle  ( cervicomandibularis) 
in    I  Id  and    Urosaurus   is   com])leteh'   ob- 

onstric- 


AM—Adductor  ir.andib 

e.\tei-nus  niedius 
CC — Constrictor  colli 


DM— Depressor  iiian(lii)ul, 
LS — Levator  scapulae 
siipoi-ficialis 


PS- -I^seudoteiiiporali 

superficiaHs 
TR     Trapezius 


Fig.   (3.      Dorsal    view    of    head    and    neck    musculature;     supeifu  ial    dejitli     at     left     and     fi 
at   right.    A.    Via;    15.    Urosaurus. 


ie|)tli 


258 


(JREAT  BASIN  N  ATT  HAL. L^ 


Vol.  35,  No.  3 


as    fan    shaped    in    I'tci    as    in    I'rosdurus 
(Figs.  6,  7,  10  ,11,  and  12). 

M.  Levator  scapular  profundus  has  a 
more  superficial  j)osition  in  ltd  than  in 
Vrosaurus  (Figs.  7,  12). 

Temporal   Mnsc  ulature 

.1/.  Ptery^oniandihularis  does  not  de- 
viate from  the  txpical  ionatiid  pattern 
(Figs.  4,  5). 

.1/.  Levator  angularis  oris  thffers  m  size 
in  Urosaurus  where  it  covers  over  half  the 
intratemporal  fossa  (Fig.  9).  In  Uta  it 
is  narrower,  covering  apjiroximately  one- 
third  of  the  fossa. 

.1/.  Adductor  mandihularis  externus  su- 
perficialis  is  similar  to  that  of  other  igua- 
nids   (Figs.  9,  10). 


M.  Adductor  itiaiidihuhiris  externus 
niedius  is  also  with  the  t\j)ical  iguanid 
pattern  (Figs.  6,  9,  10,  11)." 

U.  Adductor  mandihularis  etternus 
projundus  is  as  in  otht^r  iguanids  (Fig. 
12). 

M.  Pseudotemporalis  superficial  is  does 
not  deviate  from  other  iguanids  (Figs.  6, 
12). 

^L  Pseud()t('nip()r(dis  profundus  is  more 
ohscured  by  the  levator  pterygoideus  in 
Urosaurus   than   in   Uta    (Fig.    13). 

1/.  Adductor  mandihularis  posterior 
shows  some  \  ariations  in  the  two  genera, 
particularly  in  the  location  of  the  muscle 
with  reference  to  the  auditory  meatus 
( Fig.  13).  In  Urosaurus  it  is  located 
both  Aentral  and  anterior  to  the  meatus, 
whereas  in  Uta  the  nuiscle  is  found 
slishtlv    ventral    to    the   meatus. 


F,P     Episternoclcidoniiistnicleus         I.S      Icn 
I.P     Levator  scapulao  inofiindus  supcii 


Fig.   7.     Dorsal   view   of  liead   iiml 
right.  A.   IJla;   B.   Urosaurus. 


Sept.  1975 


FANGHELLA.  KT  AL:    Ll/ARD  ANATOMY 


259 


AI.  Levator  ptcrygoidrus  is  as  seen  in 
other  iguanids  (Fig.  14"). 

M.  Protractor  ptcrygoidrus  lias  a  nuu  h 
larger  insertion  on  the  medial  (rest  of  the 
{[uadrate  in  Urosaurus  than  in  Uta  (Figs. 
13,  14). 

DiSClTSSlON 

A  study  of  the  anterior  osteology  and 
myology  of  Uta  and  Urosaurus  reveals 
some  distinct  anatomical  differences  be- 
tween the  two  genera. 

Mittleman  (1942)  consider(Hl  the  rela- 
tionship between  Uta,  Urosaurus,  and  the 
iguanines  (Sauromalus,  Dipsosaurus,  and 
Ctenorsaura) .  He  regarded  Uta  and  Z7ro- 
saurus  as  distinct  genera.  Savage  (1958) 
outlined  the  iguanine  characteristics  and 
included  Crotaphytus  in  that  evolutionary 
line.  He  also  determined  some  structural 
differences  between  Uta  and  Urosaurus. 
The  differences   cited   by   Savage  include 


Urosaurus  possessing  a  ])ectf)ral  girdle  of 
the  urosanrine  l\pe:  lateral  xiphisternal 
ribs  present  and  no  supranasal  scales. 
Uta  is  (lislhutK  different  in  having  a 
pectoral  girdle  of  the  utiform  type,  no 
lateral  xiphisternal  ribs,  and  supranasal 
scales  separating  nasals  from  internasals. 
Avery  and  Tanner  (1964)  ])resent  several 
myological  differences  between  Sauromal- 
us  an(l  Crotaphytus  and  indicate  these  two 
genera  are  not  in  the  same  evolutionary 
line,  indicating  that  at  least  two  major 
subdivisions  exist  in  the  family  Iguanidae. 
Etheridge,  in  19(54,  also  examined  the 
iguanines  and  se])arated  Crotaphytus  from 
them  based  on  osteological  differences.  He 
states  that  osteological  comparison  sug- 
gests that  three  subgroups  of  sceloporines 
may  exist:  (1)  Holbrookia.  Callisaurus, 
and  Urna  possessing  the  scapular  fenestra; 
(2)  Uta,  Urosaurus,  Sator,  and  Sceloporus 
demonstrating  the  absence  of  the  scapular 


IE— Iiiteicostalis  e.xtciiii  SD-    Sf 

Fig.  8.     Dorsal   view   of   head 
rip;ht.    A.    Uta;    B.    I'msaurus. 


itus    (dorsal   pat 


SP-    Spinus   dorsi 


iicrU    musculature;     fourth    dei)t.l 


left    and    fifth    depth    at 


260 


GREAT  BASIN  NATURALIST 


Vol.  35,  No.  3       ! 


fenestra;  and  (3)  Pctrosaurus  possessing 
very  few  osteological  comparisons  to  the 
other  sceloporines. 

Presch  (1969)  reported  that  the  osteo- 
logical chararteristics  indicate  that  the 
horned  lizards  (Phrynosoma)  form  a 
highly  specialized  genus  within  the  sce- 
lojiorine  group  of  genera.  Phrynosoma  is 
distinguished  from  all  other  members  in 
having  a  large  sternal  fontanelle.  Petro- 
saurus.  he  states,  is  clearly  primitive  with 
its  moderately  sized  sternal  fontanelle  and 
four  sternal  ribs.  Of  the  two  groups  de- 
rived from  Petrosdurus.  the  least  altered 


are  Unjsaurus.  Via.  Sccloporus,  and  Sator. 
with  hooks  on  the  clavicle  and  a  covering 
o\er  the  anterolateral  ])rocesses  of  the 
frontal.  T'hus  Presch  places  Ufa  and  Uro- 
saurus  in  the  same  grou]i  but  as  distinct 
genera. 

Zug  (1971  )  studied  arterial  patterns  in 
many  iguanids  and  found  differences  be- 
tween Ufa  and  Urosaurus.  In  his  Figures 
10  and  15  he  illustrates  these  differences. 
Particularly  significant  is  the  representa- 
tion of  separate  phylogenetic  lines  for  these 
genera. 

Recentl}   Purdue  and  Carpenter  (1972) 


AM — Adductor  rnan(lit)ulaiis 

extornus  medius 
AS — Adductor  mandil)ularis 

e.xtemus  superficialis 
AU — Auditory  meatus 

Fig.  9.     Lateral    view    of    liciid 


(',(]     (Joiistrii  tor  coll 

i 

DM     Dciiicssor  ma 

Klil.ulai 

KI^      Epistciiioc  li'ido 

iiatdidi" 

IP      Intrnniiiulilnila 

IS 

posterior 

1    aiirl     neck    niuscul 

■iture; 

I..\      Levator   annularis 
us  I'H      I'lapezius 

superficial    depth.    A.    I'ta;     B.    Urosaur 


Sept.  1975 


rAN(;Hi-.i.i.,\.  i:t  al:  lizard  anatomy 


261 


have  studied  the  relation- 
orus.  Uta.  and  /  'rosauru^ 
h\  tlieir  dis])la\-  motions  <i 
tios  of  hi[)  and  shonldei-  ni 
tical  ev(^  nio\ cnieiil.  \\n 
LJta  and  L'rosanrns  .nc 
and   that    Urosaurus    is   i\v 


hips  of  Srr/op~ 
as  (Iclci-nnniMl 
tid  based  on  ra- 
)\('ni('nl  to  \(>i- 
\  sn,u,u(>st  that 
(HstiiKt  genera 
•i\('d    fi-oni   one 


group   of  Scrloporus,    wliile   L'/a   is   nujre 
closely  related  to  Pctrosaurus. 

Larsen  and  Tanner  (1975)  have  })re- 
sented  a  new  j^hylotreny  for  the  scelop- 
orines  hased  on  external  characteristics  in- 
cluding the  development  of  specialized 
scales  and  structure  of  the  gular  fold,  and 


(  ters 

ni(  hi(H 

1114     hip     anc 

Thev 

nid  Hat 

•  th.il  Seel  op 

1  and 

leri\ed 

h'oni  one  hnc 

intei-nal     (hi 

shoulder  rati 

0/7/V  is  adxan 

of  s(  ("lopofiiuvs.  iiK  Jnd 

I  Id.  and   /\ir(}\auf  i/s 

and  the  s,,nd  ii/.anjs. 

Iisdurus.     I  lolhroohid. 

comprise    a     separate 

Tliey    separate    the    primitive    species    of 

Sccloporus  from  that  genus  and  resurrect 

Cope's  genus  Lysoptychus  for  them.  They 

also    consider    lUd    and    Urosaurus    to   he 

closely  related,  with  Ufa  being  the  more 

primitive. 


ig  Sa/of\  Urosaurus, 
uhile  Phrynosoina 
irluding  (  ' iiid.  (d/- 
and  Cophosdurus. 
lin(>    of    evolution. 


AM-   -Adductor  iiianililuilaris 

e.xtenius  inedius 
AS — Adductor  mandihularis 

e.xternus  superficial  is 


DM-   Depressor  niandilDularis 
EP-    Episternocleidomastoideus 
IP — Interinandiliularis 
jiosterior 


LS-   Levatoi 

superficia 

OM— Oniohvoideus 

SH     Sternoliyoideu 


pulae 


Fig.    10.      Lateral    view    of    head 


let  k    musculature 


^t   depth.   A.    I'ta:    B.   rrosaurus. 


262 


GREAT  BASIN  NATHHAUST 


Vol.  35,  No.  3 


Ostcologv 

As  stated  eai'lier.  length-width  measure- 
ments of  hones  and  bone  shaj)Os  were 
utilized  to  analyze  the  osteological  rela- 
tionships between  the  two  genera.  The 
ratio  means  in  Tables  1  and  2  were  used 
to  make  these  relationships  clear.  Utilizing 
the  method  of  Avery  and  Tanner  (1971), 
one  can  assume  that  a  chfference  of  .40  or 
fewer  j)ercentage  points  (.20-. 60)  between 
means  of  the  same  bone  indicates  a  close 
relationship.  The  possession  of  bones  with 
similar  shape  is  also  an  indicator  of  close 
relationship. 


Based  on  comparisons  of  the  skulls  of 
iguanine  lizards,  Avery  and  Tanner 
(1971)  indicate  osteological  characters  of 
the  skulls  of  iguanid  lizards  to  be  stable 
within  generic  limits.  This  osteological  sta- 
bility is  also  demonstrated  by  the  skulls 
of  Uta  and   Urosaurus. 

Re^  iewing  Tables  1  and  2,  the  35  char- 
acteristics and  corresj)onding  mean  ratios 
indicate  24  structures  with  mean  ratios 
differing  bv  .40  or  more  percentage  points 
(.2()-.65).  ■ 

As  indicated  by  Etheridge  (1964),  the 
difference  between   the   skull   length  and 


AP — Adductor  niiindihul; 

externus  profundus 
LP — Levator  scapulae 

profundus 


LS    Levator   s: 
suprrfuialis 


Fig.    11.      Lateral    view    of    head 


ipulae 


dati 


PS  -  Pseudotoniporalis 
supcrfirialis 


d    df|)tli.    A.    ltd:    B.    rrosaunis. 


Sept.  1975 


FANGHKLLA.  ET  AL:    LIZARD  ANATOMY 


263 


width  ratio  in  Utd  and  Urosaurus  is  al- 
most negligible.  I  fowever,  oiu-  measure- 
ments of  the  basisphenoid  and  basioccipi- 
tal  bones  show  differences  of  1.05  and 
1.46  points  respectively,  along  with  the 
more  posterior  location  of  the  suture  be- 
tween the  two  structures. 

Considering  the  \entral  bone  structures 
(Fig.  1).  the  follwoing  mathematical  dif- 
ferences exist.  Both  the  jiterygoid  and  octo- 
pterygoid  bones  differ  by  more  than  .40 
j)oints  (Tables  1  and  2);  however,  the 
ectopterygoid  possessed  by  Urosaurus 
shows  anterior  wings  extending  to  the 
maxilla,   not   seen    in    Uta.    The   primary 


differences  found  between  the  vomer  and 
pahitine  bones  in  both  genera  is  mainly 
the  j)ositioning  of  their  common  suture. 
In  Urosaurus  the  suture  extends  antero- 
medially  from  the  inferior  orbital  foramen 
to  the  fenestra  exochioanalis,  whereas  in 
Uta  the  suture  is  found  extending  later- 
ally from  the  anterior  ])ortion  of  the  pyri- 
form  recess  to  the  maxilla. 

In  reference  to  the  nasal  capsule  (nasal, 
prefrontal,  lacrimal,  and  se{)tomaxilla), 
there  are  found  ratio  differences  in  the 
nasal  and  lacrimal  (slight  difference  in 
the  prefroiital)  and  practically  no  struc- 
tural   peculiarities.     The    premaxilla    and 


AM — Adductor  maiidilmlaris 

externus  medius 
EP — Episternocleidomastoideus 


LS — Levator  scapulae 
superficialis 


Fig.    12.     Lateral    view    of   head    and    neck    niuscula 


PM— Pharyngeal 
membrane 


thinl    depth.    A.    Uta;    B.    Lrosaurus. 


264 


(iUKAT  BASIN  NATURALIST 


Vol.  35,  No.  3 


maxilla  of  the  palate  coni])lex  show  a 
discrepancy  only  in  the  premaxilla.  The 
premaxillary  bon(>s  of  the  two  genera  dif- 
fer by  .n  points  \vith  the  premaxilla  of 
Uta  being  narrower  anteriorly  than  in 
JJrosaurus. 

The  parietal,  scpiamosal,  and  (juadrate 
(of  the  temporal  fenestra  portion  of  the 
maxillary  segment)  all  differ  in  size. 
shape,  and  ratio.  The  parietal  in  Uta  is 
much  more  rectangular  and  broad  than  in 
Urosaurus  and  co^■ers  more  of  the  pos- 
terior portion  of  the  braincase.  The  differ- 
ence between  the  squamosal  and  quadrate 


in  both  genera  is  mostly  in  ratio  (shape- 
size)    rather   than    in    location. 

On  the  dorsal  area  of  tlu>  skidl  a  striking 
difference  is  noticed  in  the  size  and  shape 
of  the  fenestra  exonarina.  In  Uta  the  fe- 
nestra is  oval  and  quite  similar  to  that  of 
SauroT?iaIus  o.  multiforaminatus  (Avery 
and  Tanner,  1964),  whereas  in  Urosaurus 
the  anterior  portion  of  the  fenestra  is  ex- 
panded anterolaterally. 

Turning  to  the  mandible,  we  see  several 
differences.  The  articular  differs  by  1.92 
points  in  I'ta  and  is  only  half  the  length 
seen   in    Urosaurus.    The   largest   mandib- 


AM-  -Adductor   mandihulnris 

posterior 
PP-  -Protractor  pterygoideus 


FT      Pseudotomporali^ 

profundus 
SI) — Spinus  dorsi 


SS--.Su[)iasrapuli 


Fig.    13.     Lateral   doi)th   of   hoad    and    neck   inus<  uhiturc;    fourth    <l(>ptii     A,    t'la;    B.    f'rosnurus. 


Sept.  1975 


lANGHELLA.  ET  AL:    LIZARD  ANATOMY 


265 


ular  difference  is  in  the  size  and  location 
of  the  surangular,  wliere  th(^  ratio  (hf- 
fcrential  is  .86  jioints  and  lh<>  [)osition  in 
Urosaurus  is  directly  Aciilral  lo  Uic  (oro- 
noid.  In  Uta,  however,  it  is  [i()st(>r()\(Mitral 
to  th(^  coronoid  with  onlx  an  anterior 
\^in^•  making  conlat  I  \Nith  the  (oronoid 
anteri()rl\'.  This  arrani^enicnt  of  the  snr- 
angnhu-  and  coronoid  I'onnd  in  Uta  is 
siniihir  to  Affihlyr/iy/u  /lus  ( risfatus\  Bra- 
chylophus  faciatus.  (lialarodon  niadagas- 
(■(ircTisis.  Conolophus  pallidus.  Clrnosaura 
pcctinatd.  Cyclura  nuicclryi.  Dipsosdurus 
(lorsalis.  Iguana  igiKnia.  Opiurus  scbac. 
and  Saurornalus  ohcsus   ( A^  ('r\'  and  Tan- 


ner,   1971  ).   None 
Urosaurus. 


lese  are   sinnlar  to 


:\lvol..-v 

The  anicr-ior  anatniny  in  I'la  wIkmi 
(()in|)ai-ed  with  dial  in  Urosaurus  sliows 
ohserx.ihie  ni\ nloLjic  <d  (hfferences.  In  con- 
sidering the  W.  onioliyoidens,  M.  sterno- 
h\-()id(>ns.  and  M.  slernothroidens,  one  can 
see  that  all  three  mnscles  were  distinctly 
separated  in  Uta  as  they  were  reported  for 
the  iguanids  Cyclura  uuchalis  and  Sauro- 
nudus     ohcsus     bv     Aver\'     and     Tanner 


(  1971  ).     where. 


resembles 


flight  I  \    that  of  Brachylophus  fasriafus. 


J'      Ia-vh 
'P      Proti 


tor  ptervgoideus 


SD-   Spiiius  (lorsi 
SK--Sonatus  iciorsal  part) 


Fig.    14.      Latei-al    \  unv    of    iicad    atid    iiort;    muscula 


liftli    (Irnt.li.    A.    rta; 


266 


GREAT  BASIN  NATLTRALIST 


Vol.  35,  No.  3 


The  dorsal  nius(  ulaturo  at  the  first 
depth  shows  a  distinct  difference  between 
IJta  and  Urosaurus.  The  M.  le\ator  scap- 
ulae superficial  is.  which  is  evident  in  all 
species  studied  h}-  Avery  and  Tanner 
(1971)  as  well  as  in  Uta.  wvis  overlaid 
by  the  trapezius  complex  in  Urosaurus. 
Another  distinction  of  the  ventral  muscu- 
lature is  that  seen  in  the  M.  episterno- 
cleidomastoideus.  In  Uta,  as  in  the  genera 
studied  by  Avery  and  Tanner,  the  M.  epi- 
sternocleidomastoideus  and  the  levator 
scapulae  profundus  are  found  anterior  to 
the  A  I.  levator  scapulae  superficialis, 
whereas  in  Urosaurus  the  jwsition  of  these 
two  muscles  is  posterior  to  M.  levator  scap- 
ulae superficialis. 

Lateral  musculature  shows  some  simi- 
larities between  Urosaurus,  Chalarodon 
madagascarensis ,  and  Oplurus  schac,  while 
the  similarity  of  Uta  to  Sauromalus  obesus 
and  Cyclura  nuchalis  (Avery  and  Tanner 
1971)  is  noticeable.  In  Urosaurus  the  M. 
episternocleidomastoideus  is  overlaid  by 
the  trapezius,  whereas  in  Lha  it  is  not 
covered.  The  M.  episternocleidomastoideus 
and  M.  levator  scajndae  superficialis  of 
Urosaurus  are  buried  beneath  the  second 
depth  of  muscle  tissue,  while  in  Uta  these 
muscles  are  mostly  superficial. 

The  orientation  of  the  M.  levator  scap- 
ulae superficialis  and  M.  le^'ator  scapulae 
])rofundus  at  the  third  depth  is  the  same  in 
Urosaurus  and  Chalarodon  rnadagascar- 
ensis,  while  these  muscles  in  Uta  resemble 
Sauromalus  obesus  and  Cyclura  nuchalis. 
In  Urosaurus  the  M.  le^■ator  scapulae  su- 
perficialis covers  the  M.  levator  scapulae 
profundus  ])osteriorly  as  seen  in  Chalar- 
odon rrmdagascarcnsis  (Avery  and  Tanner, 
1971 ) .  In  Sauromalus  obesus  the  common 
border  of  the  M.  levator  sca]:)ulae  pro- 
fundus and  M.  levator  sca])ulae  super- 
ficialis is  similar  to  that  in   Uta. 

The  temporal  musculature  of  Cyclura 
nuchalis  (Avery  and  Tanner,  1971),  M. 
protractor  ptervgoideus,  M.  ])seudotem- 
poralis  ])rofundus,  and  M.  levator  ptery- 
goidous  appears  to  be  similar  to  that  of 
Uta.  whereas  in  Urosaurus:  these  muscles 
are  seemingly  very  similai-  to  iliose  of 
Chalarodon. 


based  on  internal  and  external  characters. 
We  agree  with  this  analysis  of  Uta's  re- 
lationshi})  to  Urosaurus.  We  believe  that 
Uta  is  more  })rimitive  for  the  following 
reasons:  1.  The  ectopterygoid  of  Uta  is 
simple  ill  structure  and  shape,  while  that 
of  Urosaurus  is  greatly  expanded  and  more 
complex  in  shape.  2.  The  fenestra  exon- 
arina  of  Uta  is  similar  to  that  of  Sauro- 
malus. a  primitive  iguanine  lizard.  3. 
The  structure  of  the  surangular  and  coro- 
noid  bones  in  Uta  is  similar  to  that  seen 
in  the  more  primitive  iguanines  and  the 
Madagascar  iquanids.  4.  The  omohyoi- 
deus  muscle  complex  in  Uta  may  be  sub- 
divided into  three  distinct  muscle  bun- 
dles as  seen  in  the  primitive  Cyclura  and 
Sauromalus,  wheras  Urosaurus  resembles 
the  omohyoideus  configuration  seen  in  the 
specialized  Brachylophus.  5.  In  Uta  the 
levator  scapulae  superficialis,  levator  scap- 
ulae profundus,  and  the  episternocleido- 
mastoideus muscles  have  a  configuration 
similar  to  the  more  primitive  iguanines. 
In  Urosaurus  the  muscle  pattern  shows 
considerable  deviation.  6.  In  Uta  the  leva- 
tor scapulae  superficialis  is  superficial 
while  it  is  overlain  by  the  tra])ezius  com- 
plex in  Urosaurus,  indicating  to  us  a 
higher  degree  of  sjiecialization. 

The  similarity  of  these  two  genera  to 
the  Madagascar  iguanids  Chalarodon  and 
Oplurus  remains  a  confused  question.  If  ^ 
Oplurus  is  ancestral  to  the  iguanine  lizards  j 
as  indicated  by  Avery  and  Tanner  (1971), 
is  Clialarodon  also  ancestral  to  the  scelop- 
orine  lizards?  Ap]:)arently  not  since  both 
Uta  and  Urosaurus  share  some  character- 
istics with  Chalarodon  and  Oplurus  with- 
out establishing  a  consistent  pattern  of 
relationshij).  These  similarities  are  more 
likely  the  result  of  parallelism  than  a 
close  phylogenetic  relationship.  All  four 
genera  are  desert  animals  and  Chalarodon 
sui)erficially  resembles  both  Uta  and  Uro- 
saurus externally. 

Perhaps  future  comparisons  should  be 
inade  between  Phrynosoma  and  Petro- 
saurus  and  the  Madagascar  iguanids  to 
determine  the  phylogenies  of  the  main 
lines  of  iguanid  evolution  and  the  Mad- 
agascar iguanids. 


Phylog(Mieti(    Helationshijis 

Larson  and  Tainier  (  1975 )  consider 
Ita  to  be  more  primitive  than  Urosaurus 
and  both  more  primiti\e  than  Sceloporus. 


Conclusions  and  vSummary 

Data  (leri\(Ml  From  the  preceding  ob- 
servations strongly  indic-ate  that  Uta  and 
Urosaurus  are  (listinct  genera.  These  con- 


Sept.  197-5 


FANGIIELLA,  ET  AL:    I,T/AH1)  Ai\ATf):\IY 


267 


elusions  are  based  on  the  distinct  osteo- 
logical  and  nivological  (  liafacteristics 
found  in  the  head  and  thioat  anatomy. 

Osteological  differences  are  simunarized 
as  follows:  1.  Bones  showing  major  dif- 
ferences in  size  and  shajie  are  the  basi- 
sphenoid.  basioccipital,  pterygoid,  ecto- 
pterygoid.  premaxiUa,  articular,  and  sur- 
angular.  2.  The  ectoptervgoid  possesses 
an  extended  anterior  wing  to  the  maxilla 
in  Urosaurus  which  is  not  present  in  Uto. 

3.  The  parietal  in  Uta  is  much  more  rt>c- 
tangidar  and  broader  than  in  Urosaurus. 

4.  The  fenestra  exonarina  in  Uta  is  o^al, 
whereas  in  Urosdurus  the  anterior  portion 
is  expanded  anterolaterallw  5.  The  posi- 
tion of  the  surangular  in  Urosaurus  is  di- 
rectly ventral  to  the  coronoid,  but  in  Uta 
it  is  ])osteroventral  to  the  coronoid  with 
only  an  anterior  wing  making  a  narrow 
contact  with  the  coronoid. 

Several  noticeable  generic  my<jlogical 
differences  are  apparent:  1.  A  chstinct 
separation  of  the  M.  omohyoideus,  M. 
sternohyoideus,  anrl  M.  sternothryoideus 
is  seen  in  Uta,  whereas  in  Urosaurus  only 
the  M.  omohyoideus  and  M.  sternoh^■oi- 
deus  are  discernible.  2.  The  M.  levator 
scapulae  superficial  is  at  the  first  depth  is 
superficial  in  Uta  but  is  overlain  by  the 
tra])czius  com])lex  in  Urosaurus.  5.  In 
Uta  the  M.  episternocleidomastoideus  and 
the  M.  levator  scapulae  profundus  are 
anterior  to  the  M.  levator  scapulae  super- 
ficialis,  whereas  in  Urosaurus  the  ])osition 
of  the  first  two  muscles  is  posterior  to  the 
latter  muscle.  K  The  M.  protractor  ])tery- 
goideus,  M.  pseudotemporalis  profundus, 
and  M.  levator  pterygoideus  of  Uta  are 
similar  to  those  of  Cyclura  nuchalis.  while 
in  Urosaurus  these  muscles  are  similar  to 
Chalarodon. 

Because  Uta  and  Urosaurus  exhibit 
such  distinct  anatomical  differences,  the 
separate  generic  d(\signations  assigned  to 
them  are  considered  to  be  valid.  Phylogen- 
etically  Uta  is  considered  to  be  older  and 
more  primitive  than  Urosaurus  because 
of  (1)  the  simple  structure  of  the  ecto- 
ptervgoid in  Uta,  (2)  the  primitive  shape 
of  the  fenestra  exonarina,  (3)  the  common 
arrangement  of  the  surangular  and  coro- 
noid bones  in  Uta.  (4)  the  common  ar- 
rangement of  the  M.  omohyoideus  com- 
plex in  Uta  and  the  primitive  iguanines, 
and  (5)  the  common  configuration  of  the 
posterior  skull  and  anterior  shoulder  mus- 


lature   of   Ula   and    (he    pi-iniiti\('   igui 


linsl  I()(,K AFH-l 

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RoBLsoN,  W.  G..  AND  W.  W.  Tanner.  1962.  A 
comparative  study  of  the  species  of  the  genus 
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Sav.\ge,  J.  M.  1958,  The  iguanid  lizard  genera 
Urosaurus  and  Uta  with  remarks  on  related 
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Schmidt,  K.  P.  1953.  A  checklist  of  North 
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Saiitii.  H.  M.  194(>.  Handbook  of  lizards  of  the 
United  St.ites  and  Canada.  Comstock  Puhl. 
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S.MiTii.  H.  M..  AND  K.  H.  TvYLoR.  1950.  An 
annotated  (lieiklist  and  kev  to  the  reptiles 
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Stebbins,  R.  C.  1954.  Amphibians  and  reptiles 
of  Western  Nortii  America.  McGraw-Hill 
Book    Co.    5  36    p. 

Strjneger,  L..  AND  T.  Barbour.  1943.  A  check- 
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Van  Denbur(;h.  .1,  l'»J2,  The  rejjtiles  of  west- 
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DISTRIBUTION   AND  ABUNDANCE  OF    1111*:   BLACK-BILLED 
MAGPIE   [PICA  PICA)    IN  NORTH  AMERICA 


Carl  E.  Bock^  and  I. 


W.   I, 


Abstract. —  Analysis  of  Auduhnu    Society  Christmas    bird    mi 
variables  shows   the  degree  to  whidi   the  Bla(k-l)illed   Magpie   is 
America.    The  abundance  and  distiil)ution  of  this  species  apiicn 
barriers:    increasing  sunnner  temperatures  in   the  Southwest     ami 
precipitation  on  the  central  plains. 


tid    certain    environmental 
arid  regions  in  North 


d    b, 
sumnie 


iditv     and 


The  Black-billed  Magpie  ( Pica  pica ' 
is  one  of  the  most  conspicuous  ])asscri]io 
birds  on  the  western  plains  and  in  the 
Great  Basin.  In  this  paper  we  are  con- 
cerned ^^'ith  what  factors  determine  the 
abundance  ]iattern  and  southern  and  east- 
ern limits  of  distribution  in  this  species. 
Linsdale  (iji  Bent  1946:134)  noted  that 
■"a  rather  striking  relation  to  climate  ex- 
hibited by  this  bird  has  not  been  clearly 
explained"  or,  specifically,  that  Pica  pica 
in  the  New  World  seems  restricted  to  the 
"cold  type  steppe  dry  climate"  region 
characteristic  of  that  portion  of  the  ITiited 
States  north  of  approximately  35'^  latitude, 
west  of  about  100°  longitude,  and  east  of 
the  Sierra  Nevada-Cascade  Range.  With- 
in this  area  Pica  pica  is  widely  distributed, 
breeding  at  most  elevations  u])  to  10,500 
ft.  (Ligon,  1961). 

The  one  universal  characteristic  of  mag- 
pie habitat  is  an  association  of  thickets  or 
riparian  areas,  necessary  for  breeding  anrl 
roosting,  with  open  meadows,  grassland, 
or  sagebrush  fields  suitable  for  its  method 
of  foraging  (Linsdale,  1937;  (labrielson 
and  .lewett.  1940;  Grinnell  and  Miller, 
1944;  Jones,  I960;  Erpino,  1968).  With- 
out doubt  these  habitat  requirements  ex- 
plain the  absence  of  magpies  from  the 
closed  boreal  forests  in  the  north.  How- 
ever, this  sort  of  habitat  mosaic  occurs 
along  watercourses  in  the  Great  Plains  and 
Southwest,  so  that  it  is  not  immediately 
apparent  why  this  species  does  not  occup\ 
a  larger  part  of  the  Lhiited  States. 

We  have  analyzed  the  winter  abun- 
dance ])attern  of  Pica  pica  using  data  from 
the  annual  Audubon  Society  Christmas 
bird  counts.  Although  magpies  may  wan- 
der somewhat  in  winter  (Jewett  et  al., 
1953),  the  bulk  of  the  population  seems 
to  stay  within  the  breeding  range.  Christ- 
mas count  data  were  compared  with  cer- 
tain climatic  variables   taken   from   maps 


of  50-\c>ar  (  liniati(  iiicaiis  published  \)\  the 
U.S.  be|)arlnicMil  of  Agric  uhur(>  (1911). 
Results  pi-()\  ide  insight  nito  those'  eii- 
\'iroinneiital  tactors  which  directly  or  in- 
direct]\'  influence  the  abundance  and  dis- 
tribution  of   the  Black-billed  Magpie. 

Mi'.TilODS 

Each  (JhrisOnas  (ount  is  a  standardized 
one-day  census  conikicted  inside  a  pre- 
scribecl  circle  15  miles  in  diameter.  Hun- 
dreds of  such  counts  are  made  aimually 
in  North  America.  Data  gathered  include 
the  nuniher  of  each  sj)ecies  seen  and  the 
nund)CH'  of  ■■|)arty-hours"  of  fieldwork  as 
a  measure^  of  c ciisus  effort.  Published  re- 
^idts  of  the  cotints  provide  an  index  to 
c  ontinent-\\i(h'  patterns  of  bird  distribu- 
tion and  abundance  (Bock  and  Lepthien, 
1974;   Bystrak,   1974). 

Details  of  techniques  for  computerized 
data  storage,  retrieval,  and  analysis  have 
been  described  previously  (Bock  and  Lep- 
thien, 1974).  In  this  case  we  retrieved 
data  from  the  1969-70,  1970-71,  and  1971 
-72  Christmas  coiuits  (2,743  individual 
censuses),  sorle*!  thi-se  In-  blocks  of  five 
degrees  of  latitude  and  longitude,  and 
computed  mean  nundier  of  birds  ])er 
party-hour  lor  all  Kiunts  within  each 
l)lock  I  Eig.  I  ).  Similar  maps  of  climatic 
data  ucrc^  constructed  for  all  latitude- 
longitude  blocks  at  least  partiall}^  \^dthin 
thc^  L^nited  States,  adapted  from  the  maps 
in  the  U.S.  Department  of  Agriculture 
(1941). 

I'he  statistics  |)rogram  B:\ID-02R  (Dix- 
on. 1971  1  was  used  to  compute  stepwise 
regression  of  magpie^  mimbers  against  the 
series  of  (  limatic  \ariables  mapped. 

Rl'.Sl'LTS 

Eigiu'e  1  shows  the  winter  abiuidance 
[lattern  of  the  Black-billed  Magj)ie.  Christ- 


'Departmcnt    of    Etivironmental,    Population,    and    Organisniic   Biology,    Univcrsily   of   (Jol( 


Boulder  80302 


269 


270 


GREAT  BASIN  NATURALIST 


Vol.  35,  No.  3 


Fig.  1.  Winter  abundance  pattern  of  the 
Black-billed  Magpie,  based  on  Christmas  count 
data.  Open  blocks  =  no  birds  observed;  four  de- 
grees of  shading  represent  >  5.0,  3.0-4.9.  1.0-2.9, 
and  <  1 .0  birds  per  party-hour-,  respectively. 

mas  count  data  indicate  that  this  species 
is  restricted  to  the  region  described  by 
Linsdale  (19  37)  but  that  it  is  not  nni- 
formly  distributed  \yithin  that  area.  High- 
est densities  were  found  in  the  northern 
Great  Basin  and  on  the  northwestern 
plains  of  Montana,  Alberta,  and  Saskatch- 
ewan. Densities  appeared  lower  in  the 
southern  and  eastern  portions  of  the  range. 

Figure  1  suggests  that  those  same  fac- 
tors limiting  the  distribution  of  the  Black- 
billed  Magpie  also  may  be  influencing  its 
abundance  within  that  range.  Table  1 
shows  correlation  coefficients  between  bird 
abundance  and  various  climatic  factors. 
It  is  evident  that  magpie  densities  are 
negatively  correlated  with  a  variety  of 
temperature  and  moisture  variables. 
whether  or  not  one  includes  blocks  out- 
side the  range  of  the  species. 

Tables  2  and  3  show  the  results  of  step 
wise  multi[)le  regression  of  magpies  and 
the  climatic  \ariables  listed  in  Table  I. 
When  data  for  all  latitude-longitude  bhx  ks 
are  used,  only  two  variables  (.Jul}'  hu- 
midity and  maximum  temperature)  made 
a  meaningful  contribution  to  ihe  regres- 
sion equation,  but  these  a((i)nnte(l  lor  W- 
percent  of  the  \ariation  in  magpie  abun- 
dance.   Restricting  the  analysis  lo  llie  s[)(>- 


Table  1.  Correlation  coefficients  between 
Black-billed  Magpie  abundance  and  certain  en- 
vironmental valuables.  Bird  data  are  from  1969- 
70,  1970-71.  and  1971-72  Christmas  counts, 
grouped  by  blocks  of  latitude  and  longitude  (see 
I'ig.  1 ) ;  environmental  data  are  50-year  means 
from  USD  A  (1941). 


Variable 


X   aiuiual    t(>ni 

> 

Maximum    Im 

p 

Mininunn   ti'in 

> 

No.    frost -IVoo 

la 

AuTuial 

precipil,itii>ii 

Sunmier 

]iroci]iitation 

Winter 

l)recipitation 

X  Julv  humid 

tv 

Correlation 

coefficient 

Blocks     within 

,'\11  blocks 
(n  =  47) 

or  adjacent  to 

magpie  range 

(n  =  26) 

-.38* 
-.20 

-.39* 
-,43* 

-.41* 
-.40* 
-.36 
-.51* 

-.45* 

-.33 

-.45* 

-.31 

-.24 
-.59* 

-.12 
-.50* 

cies'  range  resulted  in  a  stepwise  regres- 
sion including  four  independent  variables 
and  accounting  for  59  percent  of  the  pat- 
tern of  magpie  density  (Table  3).  Frost- 
free  days  entered  as  the  best  predictor 
(negative)  within  the  range.  This,  how- 
ever, is  simply  another  parameter  of  tem- 
perature I'egime.  Table  ^  shows  that  the 
same  famih'  ol  \ariables  is  involved  with- 
in the  sj)ecies  range — namely,  an  inverse 
relationship  to  temperature  and  moisture 
(luring  the  warm  season  of  the  year. 


Tablk  2.  Stepwise  nuiltiple  regression  of 
eiglit  environmental  variables  against  winter 
alnuidance  of  Black-billed  Magpies.  Based  upon 
Christmas  count  data  and  climate  variables  for 
17  latitude-longitude  blocks  in  the  United  States 
(  see  text  and  Fig.  1 ) . 


Step  no.       Variable  entered 

R* 

R=** 

1  .luly  humiihty 

2  Maximum   temperature 

0.59 
0.74 

0.35 
0.54 

\V  =  iiiiillipU;    (oncl.iUnii    ,(.f-fririoiil 
I\-"    r=    1  (H'ltiriciil   (if  (Iclorniinntinii.   (^(| 

uv;il0Mt   t 
.1    .,1     rnrl 

percent 
■;tep. 

'i'\m  r.  ;.  Same  , 
upon  26  latitude-Ion 
States  within  or  adii 
range   (see  Fig.   1 ). 


Table  2.  except  based  only 
ude  blocks  in  the  United 
•ut    to   Black-billed   Magpie 


Step  no.         Variable  entered 


1  No.    frost-free   davs  0.51  0.26 

2  .Fulv  humidity  0.71  0.51 
5  Summer  precipitation  0.75  0.56 
4  Maximum    temperature  0.77  0.59 


Sept.  1975 


BOCK,  lepthii:n:  aia(;pii.  disikibtttion 


271 


Discussion  and  ('oxc.i.i mons 

Results  of  this  study  show  [\\o  dcgroo 
to  which  the  Black-billed  Magpie  is  a  bird 
of  cool  arid  climates  in  the  United  Slates. 
This  does  not  necessarily  mean  that  mag- 
pie distribution  and  abundance  actually 
are  determined  by  climate,  or  r\ou  h\  the 
influence  of  climate  on  food.  It  could  be 
that  the  species  is  restricted  by  babitat 
availability  or  by  the  appearance  of  close 
competitors  coincidental  with  changes  in 
climatic  regime.  We  can  ])ro\  ide  no  dc- 
finitiye  answer  to  this  dilennna.  but  a  \v\\ 
considered  speculations  are  in  order.  ])ar- 
ticularly  as  they  might  stinudate  more 
research. 

First,  there  is  no  clear  break  in  suitable 
habitat  in  the  United  States  wlncli  shotdd 
Imiit  magpie  distribution,  except  in  the 
West  where  the  closed  coniferous  forests 
constitute  a  sharp  boundary.  Riparian 
habitat  along  such  watercourses  as  the  Rio 
Grande  and  the  North  Platte.  South  Platte. 
Arkansas,  and  Colorado  rivers  all  support 
Black-billed  INIagpie  ])opulations;  yet. 
Pica  pica  become  scarce  and  disajipear 
along  these  rivers  when  they  reach  the  hot 
Southwest  or  the  more  humid  central 
plains.  Subtle  habitat  changes  may  occur, 
but  it  is  not  clear  wdiy  such  an  ojiportu- 
nistic  species  should  be  limited  by  them. 

Concerning  competitors,  it  is  obvious 
that  Pica  pica  is  replaced  by  the  closely 
related  P.  nuttalli  in  interior  California. 
Magpies  forage  opportunistically  on  in- 
vertebrates (especially  grasshoppers),  car- 
rion, and  various  other  items  (Linsdale, 
1937;  Verbeek,  1973).  It  is  very  difficult 
to  describe  the  foraging  niche  of  such  a 
species,  but  the  Common  Crow  (Corvus 
brachyrhynchos)  and  White-necked  Ra- 
ven (C.  cryptoleucus)  appear  geherall^ 
similar  in  food  and  habitat  requirements 
(Bent,  1946).  The  White-necked  Raven 
is  a  bird  of  the  Southwest.  The  Common 
Crow  is  distributed  all  across  the  United 
States,  but  is  especialh'  abundant  in  cen- 
tral and  eastern  regions  (Bystrak.  1974). 
While  these  two  species  of  (^onnis  do 
roughly  circumscribe  the  range  of  the 
Black-billed  Magpie,  we  would  be  rebu- 
tant  to  conclude,  without  more  field  evi- 
dence, that  they  are  invohed  in  a  com- 
petitive exclusion.  First,  the  White-necked 
Raven  is  uncommon  (Bystrak.  1974)  and 
more  typical  of  southwestern  grasslands 
than  the  actual  hot  desert  country  (Phil- 
lips et  al.,   1964)    marking  the  boundary 


)l  magpie  (bsl  fibul  ioi  i.  Second,  tlie  (oni- 
inou  (j-ow  is.  in  Die  I.  \^i(lel\■  s\  lupalfic 
uiib     ilie     P.lac  k   I. died     Mag[.ie.      hnisdale 


and 


lid 


(  l')-57  '  i-e|Mii-ls  ,,Mi\  (ic  lasi 
iiilefadioiis  belweeu  (i-o\\s  and  niagjiies. 
Vei-beek  (  I'iZ  )!  ohsecNcd  a  luimber  of  in- 
teractious  between  ci-o\\s  and  Pi((/  iiut- 
tdlli:  bc)\ve\c'f.  tb(>se  oc  c  urred  onh  .u-ounci 
llie  uesi  and  nia\  liaxc  imohed  a  response 
to  the  crows  as  potential  Jiest  predators. 
At  other  seasons  the  two  species  were 
loleccuit  of  each  other.  Finally,  Pica  pica 
is  Ilolarc  tic  in  distribution  and  in  Great 
Pjiitaiu  and  luirope  is  sympatric  with  four 
species  of  Corvus  with  generally  similar 
haliitat  recpnrements  (^Bannerman,  1953). 

^^  ('  would  like  to  conclude  by  returning 

to  the  cpie.iic f  climate.    It  seems  very 

likel^  Ihal  ihe  Black-billed  Magpie  (and/ 
or  their  in\  (>rtebrate  prey)  cannot  tolerate 
the  extreme  tem])eratures  of  a  warm  des- 
ert. Verbeek  (1972:571)  noted  that  Pica 
nuttalli  forage  in  summer  mainly  in  the 
morning,  partly  because  midday  heat  even 
in  California  stops  the  activity  of  inverte- 
brate prey,  and  ])artly  because  "this  same 
heat  sevcM'ly  limits  the  birds  in  their  feed- 
ing." In  the  Old  World  magpies  are  dis- 
tributcMl  throughout  Europe  and  into 
North  Africa  as  far  as  the  edge  of  the 
Sahara  (  Baimerman.  195  3  ) .  In  New  Mex- 
ico Pica  pica  breeds  only  in  the  northern 
third  of  the  state;  yet  in  the  cooler  winter 
season  birds  may  wander  down  the  Rio 
Grande  Valle\  considerable  distances 
(Ligon,  1961). 

While  the  eastern  distributional  limits 
of  the  Black-billed  Magpie  do  not  fit  \'S'ith 
striking  pin  siographic  or  habitat  changes, 
the\'  clo  coincide'  \\ith  major  changes  in 
climatic  regime  which  appear  to  have 
general  avifaunal  significance.  For  ex- 
am])le.  Salt  (1952)  concluded  that  the 
House  Finch  [Carpodacus  rncxicanus) 
does  not  breed  eastward  in  the  Great 
Plains  because  of  its  intolerance  of  high 
summer  humichty.  The  eastern  limits  of 
this  sj)ecies  iwv  \qy\  sinnlar  to  those  of 
the  magpi(\ 

The  eastward  disappearance  of  Pica  pica 
also  is  generally  coincidental  with  rather 
steep  zones  of  introgression  between  east- 
ern and  western  ])opulations  of  several 
bird  species.  The  most  familiar  of  these 
is  the  /one  of  ■'h\])ridization"  between  the 
western  [cafcr]  and  eastern  (auratus) 
subspecies  of  the  Common  Flicker  {Colap- 
les  auratus),  analyzed  by  Short   (1965). 


272 


(;ri:at  basin  xaii  hai 


Vol.  ^5,  No.  3 


Interestingly,  a  third  subspecies,  (\  a. 
chrysoides.  replaces  cafer  in  the  South- 
west, although  this  zone  is  somewhat  south 
of  the  limits  of  magpie  distribution.  .h)lu^ 
son  (1969:229)  reviewed  the  situation  in 
flickers  and  made  these  comments; 

One  of  llio  niosl  sliikiii^  ivn  .'lalions 
tliat  ranie  to  me  upon  exaniiiiiiig  Slioi't's 
generalized  maps  of  tlie  geof^iajihic  dis- 
tribution of  flickers  in  Nortii  America 
is  the  great  coincideiKc  of  l)llenot^•pic 
change  in  the  various  Imnis  willi  major 
climatic    boundaries    on    the    (onlinent. 

Rising  (1969),  in  fact,  studied  the  com- 
parative physiologies  of  Northern  Orioles 
{Icterus  galbula)  which,  in  the  same  part 
of  the  western  Great  Plains  as  the  flickers, 
undergo  marked  phenot^^pic  change.  He 
found  that  the  westerti  "Bullock's  Oriole" 
(/.  g.  huUocki)  is  better  adapted  to  hot  dry 
climate  than  is  the  eastern  "Baltimore 
Oriole"    (/.  g.  galbula). 

We  suggest  that  climatic  fac  tors  related 
to  temperature  and  humidity  may  limit 
the  abundance  and  distribution  of  the 
Black-billed  Magpie,  either  directly  or  by 
their  effects  upon  the  availability  of  in- 
vertebrate prey.  Future  research  on  this 
interesting  bird  coidd  profitably  include 
ecophysiological  studies  as  \\{A\  as  more 
fieldwork  on  the  relationships  of  this  spe- 
cies to  subtle  habitat  changes  or  the  ])res- 
ence  of  competitors,  especially  the  Com- 
mon Crow^ 

Acknowledgments. —  We  are  grateful 
to  the  University  of  Colorado  Computing 
Center  and  Taximetrics  Laboratory  for 
technical  assistance,  and  to  the  many  stu- 
dents who  have  helped  to  build  the  Christ- 
mas count  data  banks.  This  ])roject  was 
supported  by  grant  GB  368f)()  from  the 
National  Science  Foundation. 

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variation  in  flickers  (Colaptes)  by  Lester  L. 
Slioit.   Jr.    Wilson   Bull.   81:225-230. 

Jones.  R.  E.  1960.  Activities  of  the  magpie  dur- 
ing the  breeding  period  in  southern  Idaho. 
Northwest  Sci.   34:18-24. 

Eicon,  J.  S.  1961.  New  Mexico  birds  and 
where  to  find  them.  Univ.  New  Mexico 
Press.  .Mbuciuerque.    360  pp. 

EiNsnsTi:.  J.  M.  1937.  The  natural  history  of 
magpies.     Par.    Coast   Avifauna    no.    25. 

PlIIEEII'S.    \.    R..    J.    T.   M.\RSHAEE,   .\ND   G.   MoNSON. 

1964.     The  birds  of  Arizona.    TTniv.   Arizona 
Pi-ess.   Tucson. 

Rising.  J.  D.  1969.  A  comparison  of  metabo- 
lism and  e-vajiorative  watei-  loss  of  Baltimore 
and  Bullock  Orioles.  Comp.  BincJieuE  Phvsiol. 
31:915-925. 

Sait.  G  '\'\'  105  2.  Tlie  relation  of  metaboli'^m 
to  (limat(>  and  distribution  in  three  finches 
of  the  gtMius  Carpndacus.  Ecol.  Monogr.  22: 
121-152. 

Short.  E.  E..  Ti-.  1965.  Hvbiidization  in  the 
flickers  (Colaptes)  of  North  America.  Bull. 
Am.  Mus.  Nat.  Hist.  129:307-428. 

U.S.  Dep.\rtment  op  Agriculture.  1941.  Cli- 
mate and  man  (yearbook  of  agrii  ultuie). 
TJSDA.  Washington.  D.C. 

Verbeek,  N.  A.  M.  1972.  Daily  and  annual 
time  budget  of  the  Yellow-billed  Magpie. 
Auk    89:567-582. 

1973.  The  exploitation  svsteni  of  the 
Yellr.w  billed  Magpie.  TTniv.  Calif.  Publ. 
Zool.   99:1-58. 


NECTAR   COMPOSITION   OF   HAWKMOTH-VISITED   SPECIES 
OF   OENOTHERA    (ONAGRACEAE) 

Robert  E.   Stockliouse,  IF 

Abstract.-  Noctars  of  11  hawkmotlis-visited  taxa  of  Ocnolhcra  were  studied.  Qualitatively  the 
;ars  of  all  taxa  ucre  identical.  The  amount  of  nectar  prochjced  \)vv  night  among  the  taxa  was  quite 
•iable.  Potential  energy  availabh-  from  nectar  of  an  average  flowei-  of  O.  caespitosa  was  deter- 
ned    1,.    l.c    42.1     ,  ab.nes. 


There  is  litth'  iiifoi-inalioii  on  the 
amount  of  nectar  prochued.  its  composi- 
tion, or  nutritive  j)oteiitial  foi-  hawkmoth- 
visited  flow'ers.  (Gregor^',  Aliso  '5:357- 
419),  1963/64);  (Heinrich  and  Raven, 
Science  176:597-602,  1972);  (Handel  et 
al.,  Am.  Jour.  Bot.  ;59: 1030-1032,  1972); 
Baker  and  Baker,  Sttidies  of  nectar — con- 
stitution and  ])olhnator — plant  coevolu- 
tion.  Pages  100-140  in  Coevolution  of  Ani- 
mals and  Plants).  The  nectars  of  10 
hawkmoth- visited  taxa  of  OcJiothcra  were 
studied  to  determine  quantity  and  sugar 
composition  available  to  nocttirnal  visitors. 

Methods. —  Nectar  was  collected  \u 
capillary  tubes  (5  microliter)  in  the  field 
or  from  plants  grown  in  the  greenhouse. 
The  component  sugars  were  determined 
using  ])aper  chromatography.  Whatman 
No.  1  filter  paper  (5"  x  18")  was  spotted 
with  '5  microliters  of  nectar  from  each 
species  and  three  standard  sugars.  Each 
chromatogram  was  run  in  butanol,  etha- 
nol,  and  w'ater  (10:6:4  v/v/v)  for  44 
hours  until  the  solvent  had  nearly 
reached  the  end  of  the  paper.  The  chro- 
matograms  were  allowed  to  (h'v  and  wer(> 
spra\ed  ^^  ith  a  sugar  spra\'  ( (SO  n\\  of  95% 
ethanol,  10  m\  of  40%  TCA,  and  10  nd  of 
glacial  acetic  acid,  which  was  sattirated 
with  benzidine  dihydrochloride),' which 
made  the  sugars  visible.  The  j)ercentage 
sugar  was  determined  with  <i  Bausdi  c^- 
Eomb  low-range  hand  refrac  tomeler. 

Results  .\nd  Disci  ssion. —  The  nectar 
sugar  composition  of  eight  species  of  Oen- 
otlicra  is  presented  in  Table  1.  Qualita- 
tively the  eight  species  have  identical  nec- 
tar sugars,  consisting  of  glucose,  fructose, 
sucrose,  and  an  unknown  (prol)ably  ral- 
finose) .  Quantitative  measurements  were 
not  made,  although  it  appeared  from  the 
chromatograms  that  there  were  differ- 
ences in  the  quantity  of  the  sugars  pro- 
duced among  taxa. 

^Department  of  Biologj-,   Occitleiit.il   (College,   Los  Angeles,  Cnlifonii. 


The  amount  of  nectar  produced  per 
night  was  (h^termined  for  four  species 
{ Table  2 ) .  Oenothera  caespitosa  produced 
the  largest  volume  of  nectar,  averaging  35 
microliters  per  flower  (volumes  were  av- 
eraged for  subspecies  jonesii,  marginata, 
and  montana) .  Nectar  was  usually  within 
1  (lu  of  the  top  of  the  hypanthium  in 
().  caespitosa  when  the  flowers  opened. 
Oenothera  eximia  and  O.  muelleri  av- 
eraged 20  microliters  per  night.  Nectar 
of  O.  eximia  was  much  nearer  the  hypan- 
thial  opening  than  it  was  in  muelleri 
( Table  2  ) .  Proboscid  lengths  for  hawk- 
moths  visiting  O.  muelleri  must  be  very 
long  (at  least  1  3  cm)  if  they  are  to  gain 
access  to  the  nectar.  Oenothera  primiveris 
jiroduced  on  the  average  only  8  microliters 
of  nectar  j)er  night. 

Heinrich  (J.  Exp.  Biol.  55:223-239, 
1971  )  determined  that  the  3-gram  hawk- 
inoth,  Manduca  sexta^  expends  approxi- 
mately 1 1  calories  of  energy  per  minute 
while  hovering  and  somewhat  less  while 
flying.  Manduca  quinquemaculata  and 
Sphinx  chersis,  both  pollinators  of  O.  caes- 
pitosa  (unpublished  data),  are  approxi- 
mately- the  same  size  as  M.  sexta  and  prob- 
ably have  similar  energ;\'  requirements 
for'  flight. 


T.-\BLE  1.     Necta 
of  Oenothera. 


composition  of  eight  species 


Uiiknovvn 

Species 

Glucose 

Fructose 

Sucrose 

#1 

albicaulis 

+ 

+ 

+ 

4- 

caespitosa  ssp 

caespitosa 

+ 

+ 

+ 

4- 

jonesii 

+ 

+ 

+ 

4- 

marginata 

+ 

+ 

+ 

4- 

innijurea 

+ 

+ 

4- 

+ 

eximia 

+ 

+ 

4- 

4- 

macrosceles 

+ 

+ 

4- 

4- 

maysillesii 

+ 

-f 

+ 

4- 

muelleri 

+ 

+ 

4- 

4- 

priiniveris 

+ 

+ 

+ 

+ 

psammophila 

+ 

+ 

4- 

4- 

273 


274 


GREAT  BASIN  NATUHALLST 


Vol.  35,  No.  3 


Taiu.e  2.      Sunimai 
species  of  Oenothera. 


it    of    nertai 


(ludMl       p 


ight     by     six     hawkinoth-visited 


Average 
%   sugar 

Nectar 

volunu>s 

Average 

height  of 

nectar  in 

hypanthium 

(cm) 

Average 
length  of 

Species 

Average  in 
microliters 

Range   in 
microliters 

hypanthium 
(cm) 

caospitosa   

cximia  

32.5 

35.0 

26.0 

35 
20 

20 
8 

18-69 
5-32 

17-23 
4-9 

6.2 
4.0 

3.0 

3.0 

9.4 
5.5 

29.5 

16.0 

priniiveris 

34.0 

5.0 

xylocarpa  

33.5 

The  amount  of  jiotetitinl  energy  avail- 
able from  an  average  flower  of  O.  caes- 
pitosa  was  calculated.  \\\e  flowers  av- 
eraged 35  microliters  of  nectar  ])er  night, 
of  which  a]i[)roximately  32.5%  was  sugar. 
[Of  the  35  microliters  approximately  25.5 
(73%)  were  available  when  the  flower 
opened  at  sunset,  6  additional  had  been 
produced  by  9:15  p.m.,  the  remainder  by 
8  a.m.  the  following  morning.}  Assuming 
there  are  approximatel^'  3.7  calories  per 
mg  of  glucose  (Fleinrich  and  Raven. 
Science  176:  597-602,  1972)  there  are 
42.1  calories  per  flower  per  night  avail- 
able for  hawkmoths  [35  mg  nectar 
flower  X  0.325  (sugar  concentration)  x  3.7 


calories  mg  sugar  =  42.1  calories  per 
flower.]  Even  as  a  rough  approximation, 
it  is  clear  that  each  flower  is  a  large  po- 
tential energy  source  for  hawkmoths. 
E^■en  in  small  populations  with  only  20- 
50  flowers  open  on  a  given  night,  42  cal- 
ories per  flower  would  offer  a  large  energy 
reward  for  the  hawkmoth  pollinators. 

Acknowledgments. —  I  wish  to  thank 
Steve  and  Susan  Chaplin  and  Pat  Wells 
for  helpful  comments  on  this  manu- 
script. This  study  was  supported  in  part 
by  a  grant-in-aid  of  research  from  the 
Scientific  Research  Society  of  North 
America. 


A  REVISION  OF  THE  NEARCTIC  SPECIES  OF  CLINOHELEA 
KIEFF1<:R  (DIPrERA:  CERAIOPOGONIDAE) 

William  L.  Giogaii.  Jr.'  and  Willis  W.  Wirtlr 


Abstract. —  The  seven  species  of  Clino/ielea  known  to  inhabit  North  America  are  described  and 
illustrated,  and  a  key  is  provided  for  identification.  Two  species  groups  are  recognized:  the  unimacu- 
lata  group  and  the  bimaculatd  group.  Clinohclca  longitheca  and  C.  pscudonubifera  are  new.  Clino- 
Iielea   nebiilosn    (Malloch)    is    a    synonym   of   C.   rurrici   (Coquilh^tt)    new  synonymy. 


ClijioJielca  Kioffer  is  a  fairh 
lis  of  ceratopogoiiids.  worldwid 


mall  t^eii- 
in  (listri- 
i)iition.  Little  is  known  of  their  bioloo^•; 
hut  the  adult  females  are  predaceous  ou 
other  insects,  and  the  larvae  are  aquatic. 
The  fiAe  ])reviously  known  North  Amer- 
ican species  were  described  bv  Loew 
(1861),  Adams  (1903),  Coquillett  (1905). 
Malloch  (1915),  and  Wirth  (1952).  Most 
of  these  species  were  originally  described 
in  the  genus  Ccratopoiron  INleigen  and 
later  transferred  to  Palpomyia  Meigen, 
.Johannseniella  Williston,  or  J nliannscno- 
rnyia  Malloch.  Although  .lohannsen 
(1943)  correctly  placed  these  species  in 
Clinohelea,  the  North  American  species 
have  needed  comprehensive  revision  and 
a  good  key  for  identification. 

In  the  present  paper  seven  Nearctic 
species  of  CUnohclea,  two  of  which  are 
new,  are  described  and  illustrated.  Two 
species  groups  are  recognized  and  given 
the  names  of  the  oldest  named  species  in 
their  group.  All  of  the  types  of  CUnohelca 
species  from  North  America  have  been 
examined,  as  well  as  examples  of  17  spe- 
cies from  other  parts  of  the  world.  All 
specimens  examined  unless  otherwise 
noted  are  part  of  the  collection  of  the  Na- 
tional Museum  of  Natural  History 
(USNM)   in  Washington.    In  the  Jists  of 


speciuKMis  examined,  slide-moimted  spe- 
( imens  are  denoted  (S),  and  pinned  spe- 
( imens  as  (P).  The  types  of  our  new  spe- 
cies will  be  deposited  in  the  USNM. 

Measurements  and  other  data  are  based 
on  slide-moiuited  specimens  and  are  re- 
corded in  the  manner  of  Chan  and  LeRoux 
(  1965 ) .  When  j^ossible,  10  females  of  each 
s[)ecies  were  critically  measured.  The 
data  are  presented  in  the  following  man- 
ner: mean  value  (mininumi  value  -  maxi- 
mum value,  n  =  number  of  measure- 
ments), except  in  the  case  of  new  species, 
where  the  actual  values  are  given  for  the 
holotype,  and  the  mean,  minimum-maxi- 
mum, and  number  of  measurements  are 
given  in  the  variation  section.  Numerical 
characters  for  female  Nearctic  Clinohelea 
are  presented  in  Table  1. 

For  general  terminology  of  Ceratopogo- 
nidae  see  Wirth  (1952)  and  Chan  and  Le- 
Roux (1965).  The  following  special 
terms  are  used  in  the  descriptions  of  fe- 
males. Wing  length  is  measured  from 
the  basal  arculus  to  the  wing  tip.  Antennal 
proportions  (AP)  are  the  relative  lengths 
of  each  flagellomere;  antennal  ratio  (AR) 
is  the  length  of  the  proximal  8  flagello- 
nieres,  divided  into  the  length  of  the  distal 
5  flagellomeres.  Palpal  ratio  (PR)  is  the 
length  of  the   3rd  palpal  segment  divided 


T.\BLE     1.- 

-  Numeiical 

charac  te 

rs    of    female    Nearctic 

Clinohelea 

(minimum-maximum   values) . 

Wing 
length 

(mm) 

Wing 

breadth            Costal 
(inm)               ratio 

Palpal 
ratio 

Antennal             Femoral  spines 

Species 

ratio            Fore      Mid      Hind 

Unimacui,.\ta  Group 

curriei   2.53-3.23 

nubifera    2.32-2.42 

pseudonubifera     ....  2.03-2.19 

BlMACLILATA    GrOUP 

bimaculata    1.65-2.68 

dimidiata     2.32-2.74 

usingeri  2.50-2.74 

longitheca   1.97-2.00 


0.81-0.94 
0.71-0.77 
n.65-0.69 

0.50-0.74 

0.68-0.87 

0.70-0.81 

0.61 


0.81-0.87 
0.85-0.86 
0.82-0.83 

0.82-0.86 
0.76-0.82 
0.80-0.81 
0.82-0.84 


3.20-4.50 
4.00-4.36 
2.86-3,00 

2.60-140 
3.17-3.80 
3.17-3.67 
2.89-3.11 


1.50-1.66 
1.41-1.46 
1.35-1.38 

1.17-1.36 
1.25-1.40 
1.29-1.35 
1.17-1.36 


0-3 
0 
0 

0 
0 
0 
0 


0-2  1-3 
0-3  2-4 
0-1         0-1 


'Department    of   Enlomologv,    University    of   Marvlanii.    College   Park.   Maryland  20742. 
^Systematic    Entomology    laboratory.    IIBIII.    .\p.iA     He     Sfry  .     USD.\.     i/o     U.S.     National     Mi 
20560. 


0-3 
0 
0 
0 


Washington,     D.C. 


275 


276 


GREAT  BASIN  NATLHiAMST 


Vol.  35,  No.  3 


by  its  greatest  breadth.  'J  erniiiiology  deal- 
ing \\'ith  male  genitalia  follows  that  of 
Snodgrass  (1957)  and  Chan  and  LeRonx 
(1965).  All  female  genitalia  and  sper- 
mathecae  have  been  (h-awn  to  the  same 
scale.  Types  ha\e  been  illustrated  when- 
ever possible. 

We  are  especially  indebted  to  Mrs. 
Ethel  L.  Grogan  for  preparation  of  the 
illustrations.  Thanks  are  also  extended  to 
the  following  jiersons  and  their  institu- 
tions for  the  loan  of  type  and  other  ma- 
terial or  information  regarding  s])ecimens 
in  their  collections:  Donald  W.  Webb, 
Illinois  Natural  History  Survev,  Urbana 
(INHS);  George  W.  Byers,  Snow  Ento- 
mological INIusemn,  University  of  Kansas. 
Lawrence  (KU);  Wilford  .1.  Hanson,  Utah 
State  University,  Logan  (LTSLT) ;  and 
.Tanice  C.  Scott,  Museum  of  Comparative 
Zoology.  Cambridge,  ?slassachusetts 
(MCZ). 

Gemis  Clijiohclca  Kieffer 

C/ino/irIra  Kieffer.  I'd  7:  205.  Tvi)e-speries.  Crrn 
topogon  variegatus  WiTincrtz,  by  original 
designation. 

Di.\GNOsis. —  Moderately  large,  shining 
ceratopogonids,  usually  with  infuscated 
wings;  body  nearly  bare,  rather  slender; 
plenron  usually  with  transverse  sihery 
band.  Eyes  bare;  widely  separated.  An- 
tenna slender;  flagellomeres  1-8  long,  fla- 
gellomeres  9-13  elongate  in  female;  fla- 
gellomeres 11-13  elongate  in  male;  plume 
sparse  in  male.  Palpus  slender;  3rd  seg- 
ment slender,  lacking  a  pit.  Female  man- 
dible with  coarse  teeth.  Mesonotum  mod- 
erately   robust,    without    himieral    pits,    a 


short  anterior  tubercle  sometimes  present. 
Femora  slender,  occasionally  with  up  to 
four  spines;  4th  tarsomere  of  at  least  mid 
and  hind  legs  deeply  bilobed,  each  lobe 
ending  in  a  stout  blunt  spine  and  smaller 
spines;  fore  5th  tarsomere  greatly  swollen 
in  both  sexes;  fore  claws  equal,  mid  and 
hind  very  unequal  in  female;  all  claws 
equal  in  male.  Wing  long,  without  macro- 
trichia;  costa  extending  to  0.75  but  not 
more  than  0.90  of  wdng  length;  two  radial 
cells  present,  2nd  much  longer  than  1st; 
no  intercalary  fork;  medial  fork  broadly 
sessile.  Female  abdomen  without  eversible 
glands  or  gland  rods;  genital  sclerotization 
small,  simple;  two  well-developed  sper- 
mathecae.  Male  genitalia  with  9th  ster- 
luun  short,  broad;  9th  tergum  tapered 
\vitli  large  cerci;  basimere  and  telomere 
relatively  long  and  slender;  aedeagus  with 
low  anterior  arch,  distal  portion  broad, 
underlying  membrane  extending  beyond 
tip;  clasjiettes  usually  divided,  each  por- 
tion slender  withan  elongated  bulbous  tip. 

Iaimaturf.  stages. —  Larvae  are  aquat- 
ic. Wirth  (1951)  described  the  pupa  of 
C.  hirnaculata.  which  he  reared  from  the 
sandy  margin  of  a  small  stream  in  Vir- 
ginia. This  is  apparently  the  only  Nearc- 
tic  sjiecies  that  has  been  described  in  an 
innnature  stage. 

Adult  habits. —  Adults  can  be  found 
on  ^  egetation  bordering  water,  and  Gro- 
gan has  taken  them  at  flowers  and  from  a 
small  grove  of  trees  in  Utah.  Downes 
(1960,^1971)  stated  that  adult  females 
are  [iredaceons  on  other  small  insects  that 
are  captured  in  flight,  but  did  not  give 
specific  examples. 


Key  to  the  Nearc  tic    Species  ui  Clinoliclca  (])rimarily  Females) 

1.  b'ore  5th  tarsomere  solid  bioxMi;   wing    with    two    sjiots.    one    centered    over 

1st  radial  cell,  second  iicvir  tip  of  (osta    {  hirudcuhitd   group) 2 

bore  5lh  tarsomere  with  pale  b.iiid:   \^ill^  with   1   spot  (entered  over  1st  ra- 
dial     (('II     or     eiitii-('l\       iiiliisc  ,il('(l     ^  liuinuicnUitd  group)    5 

2.  Spermathe(ae    large,   elongated,   ellipsoid   longitJwca  n.  sp. 

Spermathecae    small,    spheroid    to    ovoid 3 

3.  Legs  pre(loniiii.nitl\    \cllow.  dislid  one-Fouiili   of  hind    fetnur  brown 

hirriaculata  (Loew) 

Legs  prcHJonnn.niilx    hfouinsh.  disi.d  one-hidf  ol    hind   fenuu'  brown  4 

4.  Tibiae  entirely   biown;  basnl  arms  of  nnde   a(>(leagus    separated 

dimidiata  (Adams) 

Tibiae  yellowish  in  midportion.  basal  an(lapi(al  portions  brown;  basal  arms 

of  male  aede.igus  inta(  t  usingeri  V^irXh. 


Sept.  1975 


(JHOCAN.  WIRTH:   CERATOPOC.OIVIDAE 


277 


5.    Wing  with  narrow  dark  infuscation  extending    from    apex    of    costa    to    tip 

nubifera   (Coquillett) 

Wing  without  narrow  dark  infuscation  at  tij)  . 6 

().    Fore  6th  tarsoniere  with  pale  hand  twice  as  long  as  width  of  tarsomere  

curriei  (Coquillett) 

Fore  5th  tarsoniere  with  pale  hanrl  nnich   shorter  than   width  of  tarso-   .... 

mere pseudonubifera  n.  sp. 

Unimaculata  Group  Clinohclca  curriei  (Coquillett) 

Wing  with  infuscation  usually  centered  \   ^S-     ■<     ^) 

over   1st  radial  cell  or  entirely  infuscated.  Ccratopogon  curriei  Coquillett,   1905:    62   (female; 

Fore   5th  tarsomere  with  pale   band.    At  British  Columbia) 

least     hind     femur     wdth     spines     usually  Palpomyia    curriei     (Coquillett);     Malloch     1914: 

present.    Species  examined   in  this  group  219  (combination,  description;  key) 

X  r            i\T     4.1      A           •          /^           •              7    J  Llinonelea   curriei    (Coquillett);   Johannsen,    1943: 

not  from  North  America:  C.  umnmculata  733   ^combination);  Wirth,  1965:    136   (distri- 

(Macquart),  Europe.  bution) 


Fig.    1.     Clinohelea    curriei    ( Coquillett]    female:    a.    antenna;    b.    leg  pattern;   c.  variations  in  hind 
leg  pattern;    d,   wing;   e.   genitalia. 


278 


GREAT  BASIN  NATURALIST 


Vol.  35,  No.  3 


Palpomyia  nebulosa  Malloch.  1915:  ^22  (female; 
Miciiignn).     new  synonymy 

Clinohelea  nebulosa  (Malloch);  Johannsen,  1943: 
783  (combination);  Wirth,  1965:  136  (distri- 
bution) 

Diagnosis. —  Distinguished  from  all 
other  Nearctic  Clinohelea  by  the  following 
combination  of  characters:  fore  5th  tarso- 
niere  with  pale  band  longer  than  width  of 
5th  tarsomere;  wing  with  infnscation  cen- 
tered over  1st  radial  cell  or  entirely  in- 
fuscated. 

Female.—  Wing  length  2.78  (2.53- 
3.32,  n  =  10)  mm;  breadth  0.85  (0.81- 
0.94,  n  -   10)  mm. 

Head:  Brown.  Antenna  (Fig.  la)  slen- 
der; pedicel  yellow  to  pale  brow^i;  basal 
flagellomere  with  j)roximal  two-thirds 
j)ale,  distal  one-third  brown;  remaining 
flagellomeres  brown;  AP  22-10-10-10-10- 
10-11-12-30-28-29-29-31  (n  =  10);  AR 
1.57  (1.50-1.(56,  n  =  10).  Palpus  brown; 
3rd  segment  longer  than  5th;  PR  4.03 
(3.20-4.50,  n  =  10).  Mandible  like  that 
of  C.  bimaculata  (Fig.  4c). 

Thorax:  Mesonotum,  scutellum,  post- 
scutellum  dark  browTL  Fegs  (Fig.  lb) 
yellow;  apex  of  fore  tibia,  mid  femorotibial 
area,  fore  distal  3  tarsomeres,  mid  and 
hind  4th  and  5th  tarsomeres  brown;  hind 
leg  pattern  variable,  most  common  form 
(Fig  lb)  with  distal  one-sixth  of  femur 
and  tibia  brown,  other  hind  leg  patterns  as 
in  Figure  Ic;  fore  5th  tarsomere  with  pale 
band  longer  than  width  of  tarsomere;  0-3 
fore,  0-2  mid,  and  1-3  hind  femoral  spines. 
Wing  (Fig.  Id)  usually  with  infuscated 
area  centered  over  1st  radial  cell  or  en- 
tirely infuscated;  veins  brown;  CR  0.84 
(0.81-0.87,  n  =   10).    Halter  pale. 

Ahdorrien:  Brown.  Genitalia  as  in 
Figure  le  with  a  ])air  of  slender,  pos- 
teriorly directed  sclerotized  arms  arising 
anteriorly  from  a  lighter  sclerotized  area. 
Spermathecae  small,  spheroid  to  ovoid, 
subequal   to  unequal    with   short   nee  k^. 

Male. —  Similar  to  female  with  follow- 
ing differences:  smaller;  antennal  pedicel 
flark  brf)wn.  flagellum  brown;  legs  more 
diffused  with  brown;  0-1  fore,  0-1  mid, 
and  0-2  hind  f(>moral  spines.  Genitalia 
form  and  shape  like  that  of  C.  bimaculata 
'Fig.  4f ) ;  aedeagus  as  in  Figure  6a. 

Distribution. —  Alaska  and  California 
to  Newfoundland  and  Florida  (locality 
records  plotted   in  Figure  3). 


Types. —  Holotype  female  of  C.  curriei, 
Kaslo,  British  Columbia,  17  .Tune  1903,  R. 
P.  Currie  (Type  no.  8361,  USNM);  holo- 
tyj)e  female  of  C.  nebulosa,  Grand  Junc- 
tion, Van  Buren  Co.,  Michigan,  15  July 
1914,  C.  A.  Hart  (INHS). 

Specimens  examined. —  82  slides,  218 
pinned  specimens  from: 

ALASKA:  Anchorage  (Aldrich) ;  Matanuska 
(Chamberlin).  CALIFORNIA:  Eldorado  Co.,  Lu- 
ther Pass  (Schlinger.  Univ.  Calif.  Davis).  CON- 
NECTICUT: Fairfield  Co..  Redding  (Melander); 
Tolland  Co..  Storrs  (Melander).  DELAWARE: 
New  Castle  Co.,  Delaware  City.  FLORIDA:  Ala- 
chua Co..  Gainesville  (Wirth).  IDAHO:  Ada  Co., 
Boise  (INHS);  Nez  Perce  Co.,  Sweetwater  (Aid- 
rich).  INDIANA:  Porter  Co.,  Mineral  Springs 
(INHS).  IOWA:  Hancock  Co.,  Pilot  Knob  St. 
Park  (Gaud).  MAINE:  Hancock  Co.,  Bar  Harbor 
(Johnson).  MASSACHUSETTS:  Franklin  Co., 
Row."  (Cohcr);  Middlese.x  Co.,  Bedford  (Wirth); 
Concord  (Wirth);  Suffolk  Co..  Boston  (Melander). 
MICHIGAN:  Cheboygan  Co.  (Dreisbach),  Doug- 
las Lake  (Williams) ;  Clare  Co.  (Dreisbach) ; 
Iron  Co.  (Dreisbach);  Lake  Co.  (Dreisbach); 
Livingston  Co.,  George  Reserve  (Sabroskj-,  Steys- 
kal);  Manistee  Co.  (Dreisbach);  Midland  Co. 
(Dreisbach);  Nottawa  (Sabrosky.  Dreisbach); 
Missaukee  Co.  (Dreisbach);  Osceola  Co.  (Dreis- 
bach); Roscommon  Co.  (Dreisbach);  Van  Buren 
Co.,  Grand  Junction  (Hart,  holotype  of  nebulosa); 
Wexford  Co..  (Dreisbach) .  MINNESOTA:  Ram- 
sey Co.  (Wall).  NEBRASKA:  Cherry  Co..  Hack- 
berry  Lake  (Wirth),  Pelican  Lake  (Wirth). 
NEW  HAMPSHIRE:  Grafton  Co.,  Stinson  Lake 
(Wirth).  NEW  YORK:  Chautauqua  Co.,  S.  Day- 
ton (Wirth);  Erie  Co..  East  Aurora  (Van  Duzee), 
East  Concord  Bog  (Wirth);  Franklin  Co..  Adiron- 
dacks  (Melander);  Lew^is  Co..  Brantingham  Lake 
(Wirth).  Letchworth  St.  Park  (Wirth),  Whet- 
stone Gulf  (Wirth);  Monroe  Co.,  Braddock  Bay 
(Wirth);  Orleans  Co..  Albion  (Wirth);  St.  Law- 
rence Co..  Cranberry  Lake  (Wirth);  Suffolk  Co., 
Cold  Spring  Harbor  (Melander);  Tompkins  Co., 
Ringwood  Reserve  (Wirth).  OHIO:  Summit  Co. 
(Lipovsky,  KU).  UTAH:  Cache  Co..  Hvrum 
(Grogan).  VERMONT:  Caledonia  Co..  Lyndon 
(Melander).  VIRGINIA:  Alexandria  (Wirth); 
Fairfax  Co..  Dead  Run  (Wirth).  WISCONSIN: 
Polk  Co.  (Baker,  paratype  of  nebulosa,  INHS). 
WEST  VIRGINIA:  Pocahontas  Co..  Cranberry 
Glades  (Wirth.  Sabrosky).  BRITISEt  COLUM- 
BIA: Kaslo  (Currie.  holotype  of  curriei).  Quebec: 
Meach  Lake  (Wirth).  NEWFOUNDLAND: 
Squire's  Mem.  Park  (Alexander).  NOVA 
SCOTIA:  Baddeck  (Fairchild).  ONTARIO:  Al- 
gonquiti  Park  (Wirth);  Kemptville  (Wirth);  Ot- 
tawa (Melander.  Wirth);  Toronto  (Van  Duzee); 
Waubamick    (Melander). 

Disci TssioN. —  The  Palaearctic  species, 
C.  unimaculata  (Macquart)  closely  re- 
sembles C.  curriei.  However,  the  hind  tibia 
is  pale  except  for  the  narrow  base  and 
apex;  the  apices  of  the  fore  and  mid  fe- 
mora are  conspicuously  dark,  narrowly 
on  the  fore  leg  but  more  broadly  on  the 
mid    leg;   atifl    the  hind   femur  lacks  any 


Sept.  1975 


CROCAN,  WIHTH:    CKKATOPOCON  IDAi: 


279 


trace  of  infuscation  except  tlie  conspicuous 
a[)ical  (lark  band. 

Clitiohclcd  nuhifcra   ( Coquillett) 
(Fig.  2a.  c.  e.  g;  fib) 

Ceratopogon  iiubifer  Coquillett.   1905:    f)l    li'eniale; 

Florida) 
Palpomyia   nubifera    (Coquillett);   Mallocb.    1914: 

217  (combination;  key) 
Clinohelea      nubifera       (Coquillett);       .lohannsen, 

1943:     783    (combination);    Wn-tli,    1965:     136 

(distribution) 

Diagnosis. —  Distinguished  from  all 
other  Nearctic  Clinohelea  by  the  following 
combination  of  characters:   wing  with  in- 


fuscation centered  o^■er  1st  radial  cell  and 
a  narrow  infuscated  band  extending  from 
apex  of  costa  to  wing  tip;  and  fore  5th 
tarsomere   with   pale  band. 

Female.—  Wing  length  2.35  (2.32- 
2.42,  n  =  3)  mm;  breadth  0.74  (0.71- 
0.77,  n  =  3)  mm. 

Head:  Vertex  and  proboscis  brown, 
frontoclypeus  lighter  brown  to  yellowish. 
Antenna  (Fig.  2a)  slender;  pedicel  yellow 
to  light  brown;  proximal  5-8  flagellomeres 
j)ale  on  basal  portions,  distal  portions  light 
brown;  distal  5  flagellomeres  brown;  AP 
26-12-1 1-1 1-1 1-1 1-1 1-12-32-29-30-30-30  (n 
=  3);  AR  1.43  (1.41-1.46,  n  =  3).  Palpus 


Fig.  2.     Female    Clinohelea:    a,c,e,g,    C.    nubifera    (Coqudlett); 
,b,  antennae;  c,d,  leg  patterns;  e,f,  wings;  g,h,  spermathecae. 


d.f.h.    C.    pscudonubifera    n.    sp. 


280 


GREAT  BASIN   NATURALIST 


Vol.  35,  No.  3 


brown;  3rcl  segment  nearly  twice  as  long 
as  5th;  PR  4.12  (4.()()-4.36"  n  =  3).  Man- 
dible like  that  of  C.  himaculata  (Fig.  4c). 

Thorax:  Mesonotuni,  scutellnm,  post- 
scutellimi  dark  brown.  Legs  (Fig.  2c) 
yellow;  proximal  jiortions  of  mid  and  hind 
coxae,  distal  i)ortion  of  hind  femur,  apices 
of  fore  and  mid  femora,  all  of  hind  tibia, 
proximal  one-half  of  fore  and  mid  tibiae, 
and  4th  and  5th  tarsomeres  brown;  fore 
5tli  tarsomere  with  })ale  band  shorter  than 
width  of  tarsomere;  0-2  mid  and  2-4  hind 
femoral  sj)ines.  Wing  (Fig.  2e)  veins 
brown;  infuscated  area  centered  over  1st 
radial  cell,  and  narrow  infuscated  band 
extending  from  apex  of  costa  to  tip;  CR 
0.86  (0.85-0.86.  n  -  3).  I4alter  stem 
brownish;  knob  pale. 

Abdomen:  Brown.  Sjiermathecae  (Fig. 
2g)  small,  ovoid.  sid)equal  with  short 
]iecks. 

Male. —  Unknown.  A  female  specimen 
from  Santa  Rosa  Co.,  Florida,  had  male 
genitalia  moimted  with  her  on  a  slide. 
These  male  genitalia  were  remounted  in 
an  attempt  to  examine  them  in  detail.  The 
overall  shape  and  form  of  the  genitalia 
are  like  that  of  C.  himaculata  (Fig.  4f ) ; 
aedeagus  as  in  Figure  6b. 


Dlstkibution. —  Florida,  New  York 
(locality  records  plotted  in  Figure  3). 

Type. — Holotype,  female,  Jacksonville, 
Florida,  Mrs.  A.  T.  Slosson  (Type  no. 
8357,  USNM,  pinned). 

Specimens  examined. —  From  the  fol- 
lowing localities: 

FLORIDA:  Alachua  Co.,  Gainesville  (Blan- 
ton),  2  females  (S);  Jacksonville  (Slosson,  holo- 
type female.  P);  Highland  Co.,  Sebring  (Wirth), 
1  female  (P);  Santa  Rosa  Co.,  Blackwater  River 
(Fairchikl),  1  female  (S).  NEW  YORK:  Suffolk 
Co.,  Cold  Spring  Harbor  (Melander),  1  female 
(P). 


Clinoheh'a  pseudoimhifera  Grogan 
and  Wirth,  n.  sp. 

(Fig.  2b,  d,  f,  h;  6c) 

Clinohelea   species    1;   Wirth,    1951:    321    (females; 
Virginia). 

Diagnosis. —  Most  closely  related  to 
(\  uulnfera.  and  can  be  distinguished  from 
all  other  Nearctic  Cliiiohdea  by  the  fol- 
lowing combination  of  characters:  fore  5th 
tarsomere  with  very  short,  pale  band,  legs 
mostly  yellow  wdth  hind  tibia  and  distal 
five-sixths  of  hind  femur  brow^n,  and  wing 


nubiTera 
C.  pseudonubifera  n 


Fig.  3.     North   American   localitv   records  for  Clino/irlra  of  tiic  iiniiuaculatn  group. 


Sept.  1975 


r,RO(;.\X.  WIHTII:  ceratopogonidae 


281 


with  iiifuscatioii  ((Mitci-cd  nxcr  1st  r-adial 
cell. 

Female  iioloiyim:. —  \^illl^  Iciit^tli  2.0^ 
mm;  breadth  ().(y5  mm. 

Head:  Vertex  and  [jroboscis  brown; 
frontoclypeiis  lighter  brown.  Antenna 
(Fig.  2b)  slender;  pethcel  yelloNv.  j^roximal 
4  flagellomeres  pale,  distal  9  flagellomeres 
brown;  AP  17-9-9-8-9-9-9-10-21-21-21-21- 
26;  AR  \.W.  Pal|)ns  brown;  ^rd  segment 
about  as  long  as  1th;  I^R  100.  Mandible 
Hke  that  of  ('.  hiffit/culafa   (Fig.  4c). 

lljorai:  Mcsoiiotnm,  scutellum,  j)ost- 
scutellum  dark  lirown.  Legs  (Fig.  2d) 
yellow;  ])roximal  jiortions  of  coxae,  (hs- 
tal  five-sixths  of  hind  femur,  distal  one- 
fifth  of  mid  fenuir,  mid  and  hind  tibiae, 
and  4th  and  '5th  tarsomeres  of  tarsi  browu; 
fore  tibia  very  light  brown;  fore  5th  tarso- 
mere  with  ver^'  short,  pale  baud;  mid  and 
hind  femora  with  1  spine.  Wing  (Fig. 
2f)  with  dark  infuscation  centered  over 
1st  radial  cell;  \eins  brown;  CR  0.8  3.  Hal- 
ter pale  with  dark  brown  spot  on  knob. 

Abdomen:  Rrown.  Spermathecae  (Fig. 
2h)  small,  o^•oid.  subequal,  with  short 
necks. 

Male  .allotype. —  Similar  to  female 
holot^"pe  with  the  following  differences: 
smaller;  antennal  ])edicel  dark  brown, 
flagellum  brown;  femora  lacking  spines. 
Genitalia  sha{)e  and  form  like  that  of 
C.  himacidata  (Fig.  4f);  aedeagus  as  in 
Figure  6c. 

Etymology. —  The  iiame  pseudotinhi- 
fera  refers  to  the  resembhnue  to  ('.  nuhi- 
fera. 

Varl\tion. —  The  following  characters 
were  recorded  for  the  single  female  to])o- 
t^•pe:  wing  length  2.19  mm;  breadth  0.69 
mm.  AR  7.35.  PR  2.86.  CR  0.82  The 
general  coloration  of  all  of  the  paratypes 
is  like  that  of  the  holotype.  Femoral 
spines  ranged  from  0-1  mid.  and  0-1  hind. 

Distribution.  -Ontario  to  North  (>aro- 
lina  (locality  records  plotted  in  Figure  3). 

Types. —  Female  holotyjje,  male  allo- 
type, 1  female  j)aratype.  Snow  I  [ill. 
Worcester  Co.,  Maryland,  2  June  1968. 
W.  H.  Anderson,  light  tra])  (Tvpe  no. 
66495,  USNM).  Other  paratyj)es,  4 
pinned  females  as  follows:  NORTH 
CAROLINA:  Macon  Co.,  Highlands.  15 
.Tune  1957,  .1.  R.  Vockeroth.  1  female 
(Canada  Nat.  Coll.).  ONIARIO:  Ottawa, 


Mer  Rleue.  23  .June  1952  G.  E.  Shewell, 

1  female  (CNC).  VIRGINIA:  Fairfax  Co., 
Falls  Church,  4  July  1950,  W.  W.  Wirth, 

2  females. 

Discussion. —  Wirth  (1951)  in  refer- 
ence to  the  two  female  specimens  from 
Falls  Church,  Virginia,  stated  that  they 
were  close  to  C.  nubifera  and  C.  dimidiata 
but  declined  to  name  them  at  the  time. 


Bimaculata  Group 

\\'ing  with  two  nifuscated  areas,  one 
(entered  o\er  1st  radial  cell,  the  other 
near  tij)  of  costa.  Fore  5th  tarsomere 
uniformly  brown.  Femora  usually  lack- 
ing spines,  or  if  present,  only  on  hind 
femur.  Species  examined  in  this  group 
not  from  North  America:  C.  barrettoihane 
and  Duret,  Brazil;  horacioi  Lane,  Brazil; 
rieirai  Lane.  Brazil;  nigripes  Macfie,  Bra- 
zil; pachydactyla  Kieffer,  Singapore; 
nibriceps  Kieffer,  Paraguay;  saltanensis 
Lane  and  Duret,  Argentina;  townesi  Lane, 
Brazil;    toivnseudi   Lane,   Brazil. 

(linohelea  bimaculata  (Loew) 
(Fig.  4,  6d) 

Ceratopogon  bimaculatus  l.oew,  1861:  311  (fe- 
male; Wasliington.  D.C.). 

Johaiinsrniella  bitnaculain  (Loew);  Malloch,  1914: 
226    (combination;    description;    kejO- 

J  ohannsenomyia  bimaculata  (Loew);  Malloch, 
1915:   352  (combination;  l^ey). 

Clinohelea  bimaculata  (Loew);  Kieffer,  1917:  317 
(combination;  key;  fig.  tarsus);  Wirth,  1951: 
321  (description  and  fig.  pupa);  Johannsen, 
1952:  164  (key,  fig.  tarsus);  Wirth,  1965:  136 
(distribution). 

Diagnosis. —  Distinguished  from  all 
other  Nearctic  Clinofielea  by  the  following 
combination  of  characters:  legs  mainly 
yellow  with  a  dark  subapical  band  on  the 
hind  femur,  two-spotted  wings,  and  an- 
tennal pedicel  yellow. 

Female.—  Wing  length  2.06  (1.65-2.68, 
n  10)    nun;    breadth    0.63    (0.50-0.74, 

n   =  10 )  mm. 

Head:  Frontovertex  brown;  j)roboscis 
and  palpus  pale  yellow.  Antenna  (Fig. 
la )  slender;  pedicel  yellow,  proximal  8 
flagellomeres  brown,  distal  5  flagellomeres 
lighter  brown;  AP  20-11-10-10-10-10-11- 
12-24-23-23-23-25  (n  =  10);  AR  1.24 
(1.17-1.36,  n  =  10).  Palpus  with  3rd  seg- 
ment slightly  longer  than  5th;  PR  2.87 
(2.60-3.40,  n  =   10).    Mandible  (Fig.  4c) 


282 


GREAT  BASIN  NATURALIST 


Vol.  35,  No.  3 


Fig.  4.     Clinohelea   bimaculata    (Loew):    a-e,   female;    f,    male   genitalia;    g-i,   pupa;    a.   antenna;   b, 
wing;  f,  mandible;  d,  leg  pattern;  e.  genitalia;   g.   operculum;  li.  respiratory  organ;  i.  anal  segment. 


heavily  sclerotized;  inner  margin  with  six 
to  nine  large  coarse  teeth;  outer  margin 
with  four  or  five  small  teeth. 

Thorax:  Mesonotum,  scutellum,  post- 
scutellum  brown.  Legs  (Fig.  4d)  yellow; 
proximal  portion  of  hind  coxa,  subapical 
band  on  hind  femur,  and  1th  and  ^th  tar- 
someres  brown;  hind  f(Mnur  witii  0-3 
spines.  Wing  (Fig.  lb  i  veins  brown;  in- 
fuscations  centered  ov(n-  1st  raihal  cell  and 
just  before  tip  of  costa;  occasionally,  in- 
fuscated  areas  joined  posteriorly;  CR  0.84 
(0.82-0.86,  n  =  10).  Halter' stem  pale 
yellowish;  knob  white. 


Abdomen:  Brown  to  reddish  brown. 
Genitalia  as  in  Figure  4e  wdth  a  pair  of 
slender,  sclerotized,  anteriorly  directed 
arms  arising  from  shorter,  thicker  arms. 
Spermathecae  small,  spheroid  to  ovoid, 
subequal  to  unequal,  with  short  necks. 

Mali.. —  Similar  to  female  with  the 
following  differences:  smaller;  antennal 
pedicel  brown;  hind  femur  lacking  spines. 
Genitalia  as  in  Figure  4f.  Ninth  sternum 
about  three  times  broader  than  long,  base 
shghtly  curved  with  a  caudomedial  exca- 
vation; 9th  tergum  tapered  distally  to  a 
rounded  tip.  cerci  short,  not  reaching  apex 


Se])t.  [97  1 


CKOCAIV.  WIRTH:    CllHATOPOGONIDAE 


283 


of  basimeres.  Basiniorc  sli^htl^  curved. 
2.5  times  longer  tliaii  broad;  telomere 
slightly  longer  than  iiasimer(\  luiNcd. 
tapered  distally  ^^ith  poiiiti^d.  hooked  lip. 
Aedeagus  (Fig.  ()d  i  hea\  il\  selerolized, 
triangular,  aliout  as  broad  a>>  long:  basal 
arm  recurved  about  'M)  degret^s.  hea\ily 
sclerotized;  (hstal  portion  with  iilinit 
pointed  ti]i;  underl^  ing  membrane  ex- 
tending beyond  ti]).  rounded  with  a  dark 
spot.  Clasj)ettes  di^ided;  basal  arm 
heavily  sclerotized,  recm'\ed;  (hstal  portion 
more  lightly  sclerotizecb  ti])  (>longate.  bid- 
bous. 

PrPA. —  Length  3.5  nmi;  color  light 
brown.  0])ercuhmi  (Fig.  4g)  narrow;  0.9 
times  as  broad  as  long  with  a  ]iair  of 
rounded  tubercles  bearing  long  seta;  sur- 
face with  fine  tubercles,  those  on  lateral 
margin  sharp  and  setose.  Respiratory 
horn  (Fig.  4h )  moderately  long  and  slen- 
der, about  iWo  times  longer  than  broad 
with  10  aj)ical  spiracular  paj^Uae.  Anal 
segment  (Fig.  4i)  about  twice  as  long  as 
broad;  surface  covered  with  fine  tubercles; 
apicolateral  processes  about  one-third  of 
total  length,  with  subapical  fine  tubercles; 
tips  heavih'  sclerotized  and  sharply  ])oint- 
ed. 

Distribution. —  Michigan  and  Texas 
to  New  Hampshire  and  Florida  (locality 
records  plotted  in  Figure  7  ) . 

Type. —  Holotype,  female,  Washington, 
D.C.,  Osten-Sacken  coll.  (Type  no.  10379, 
MCZ). 

Specimens  examined. —  177  slides,  158 
pinned  specimens  from: 

ALABAMA:  Mobile  Co..  Mobile  ( Blantou, 
Cannon).  CONNECTICtJT:  Litchfield  Co..  Lake 
Waramaug  (Melander).  DLSTRICT  OF  COLUM- 
BIA: Washington  (Coquillett ) .  FL0RII3A:  Ala- 
chua Co.,  Gainesville  (Blanton.  Wirth);  Baker 
Co.,  Olustee  (Blanton) ;  Bay  Co..  Panama  City 
Beach  (McElvey);  Calhoun  Co..  Blountstown 
(Blanton);  Collier  Co..  Collier  Seminole  St.  Park 
(Wirth),  Ochopee  (Blanton);  Escambia  Co.,  Bratt 
(Blanton);  Glades  Co..  Palmdale  (Irons);  Gulf 
Co.,  2  mi.  N  Beacon  Hill  (Blanton).  Wevva- 
hitchka  (Blanton);  Hardee  Co.,  Ona  (Irons); 
Highlands  Co.,  Archbold  Biol.  Sta.  (Wirth).  Lake 
Placid  (Layne),  Sebring  (Wirth);  Indian  River 
Co.,  Fellsmere  (Wirthj.  Vero  Beach  (Wirth); 
Duval  Co..  Jacksonville  (Knight) ;  Jefferson  Co., 
Monticello  (^'V^^itcomb) ;  Lake  Co.,  Leesburg 
(Braddock);  Leon  Co.,  3  mi.  N  Tallahassee  (Blan- 
ton); Liberty  Co..  Torreya  St.  Park  (Blanton, 
Fairchild.  Weems,  Wirth);  Marion  Co.,  Juniper 
Springs  (Wirth);  Orange  Co..  Lake  Magnolia 
Park  (Irons),  Rock  Springs  (Wirth);  Palm  Beacli 
Co..  W.  Palm  Beach  (Hardy,  KU);  Putnam  Co., 
Lon's     Lake     (Blanton);     Sarasota     Co.,     Myakka 


i^iver  St.  Park  (Wirth);  Suwanee  Co.,  Suw,'anee 
Springs  (Beamer,  KU);  Wakulla  Co.,  Ocklocko- 
iiee  River  St.  Park  (Wirth);  Walton  Co.  (Butler). 
(iKORGTA:  Charlton  Co.,  Okefenokee  Swamp 
i  Beamer.  KU);  .Mitchell  Co.,  Newton  (Pratt); 
iiiomas  Co..  ThomasviUe  (Palmer).  ILLINOIS: 
llenrv  Co..  Algonquin  (INHS);  Champaign  Co., 
Urbana  (Malloc.h.  INHS);  Piatt  Co.,  Monticello 
i.Malloch,  INHS);  Pulaski  Co.,  Pulaski  (Malloch, 
INHS).  INDIANA:  Tippecanoe  Co.,  Lafayette 
(Aldrich.  Melander).  LOUISIANA:  East  Baton 
Rouge  Parish.  Baton  Rouge  (Wirth).  MARY- 
LAND: Anne  Arundel  Co..  Mayo  (Wirth);  Cal- 
v(Mt  (ji..  (;hesapeake  Beach  (Shannon.  Knab); 
(^harh's  C>o.,  Nomomonee  (Wirth);  Frederick  Co., 
Thurmont  (Steyskal);  Montgomery  Co.,  Glen 
Echo  (Malloch);  Prince  Georges  Co..  Beltsville 
(Malloch);  Worcester  Co..  Snow  Hill  (Wirth). 
MASSACHUSETTS:  Middlese.x  Co.,  Bedford 
(Wirth).  Concord  (Wirth).  MICHIGAN:  Lapeer 
Co..  Deerfield  (Stevskal);  Livingston  Co.,  George 
Reserve  (Stevskal);  Midland  Co..  (Dreisbach); 
Wayne  Co.,  Detroit  (Steyskal).  NEBRASKA: 
Nemaha  Co.,  Peru  (Harmston).  NEW  HAMP- 
SHIRE: Grafton  Co.,  Stinson  Lake  (Wirth). 
NEW  YORK:  Franklin  Co..  Adirondacks  (Melan- 
der). NORTH  CAROLINA:  Durham  Co.,  Nel- 
son (Beamer.  KLT);  Onslow  Co..  Jacksonville  (Bo- 
hart.  USU).  SOUTH  CAROLINA:  Georgetown 
Co..  Hobcaw  House  (Henry).  TENNESSEE: 
Lake  Co.,  Reelfoot  Lake  (Snow).  TEXAS:  Col- 
lin Co.,  Piano  (Tucker);  Kerr  Co..  Hunt  (Wirth), 
Kerrville  (Bottimer).  VIRGINIA:  Alexandria 
(Wirth);  Fairfax  Co.,  Falls  Church.  Montgomery 
Co..  Blacksburg  (Messersmith).  WEST  VIR- 
GINIA: Pocahontas  Co.,  Cranberry  Glades 
(Wirth.  Sabroskv) ;  Taylor  Co.,  Grafton  (Steys- 
kal). 

CUnohclcd  dimidiata  (Adams) 
(Fig.  5a,  e,  g;  6e) 

Ceratopogon  diinidiatus  Adams,  1903:  27  (female; 
Arizona). 

Johannseniella  dmndiata  (Adams) ;  Malloch:  226 
(combination;   key). 

] ohannsenomyia  dimidiata  (Adams) ;  Malloch, 
1915:332   (combination;  key). 

Clinohelea  dimidiata  (Adams);  Johannsen,  1943: 
783  (combination);  Wirth,  1965:  136  (distri- 
bution). 

Diagnosis. —  Distinguished  from  all 
other  Nearctic  ClinoJielea  by  the  two- 
spotted  wings  and  dark  brown  tibiae; 
males  with  basal  arms  of  aedeagus  sep- 
arated. 

Female.— Wing  length  2.54  (2.32-2.74, 
n  =  5)  mm;  breadth  0.76  (0.68-0.87,  n  = 
4)  nmi. 

Head:  Rrown.  Antenna  (Fig.  5a)  slen- 
der; brown,  proximal  two-thirds  of  basal 
flagellomere  pale;  AP  17-10-9-9-9-10-10- 
11-22-21-23-23-25  (n  =  4);  AR  1.35 
(1.25-1.40,  n  -  4).  Palpus  with  3rd  seg- 
ment slightly  longer  than  5th;  PR  3.43 
(3.17-3.80,  n  =  3).  Mandible  like  that  of 
C.  biinaculata  (Fig.  4c). 


284 


GREAT  BASIN  NATUKALIST 


Vol.  35,  No.  3 


c:ocii3Gr]c 


Fig.  -5.     Fenial 
thcca  ri.  sp.;  ;i-c.  ;: 


(Uinnhclca:    a.e.p.    C.    dimidiata    (Adams);  b.d.h.  C.  usingeri  Wirth;  r.f.i.  C.  longi- 
tciinac;  d-f.  \v^  pattcfns;   g.  genitalia;   h.   i.  sponnathecae. 


Thorax:  Mesoiiotuni,  scutelluni,  post- 
scutellum  dark  brown.  Legs  (Fig.  5ej 
brown;  yellow  on  fore  coxa,  tli.stal  one- 
fourth  of  mid  nnd  Iniid  coxae,  trochanters, 
most  of  fore  tcimu-.  proximal  five-sixths 
of  mid  femur,  proximal  half  of  hind  fe- 
mur, and  mid  and  hind  1st  and  IwA  tarso- 
meres.  Wing  like  that  of  ('.  hima(  uhita 
(Fig.  4bj.    Halter  stem   pale;   knol)  white. 

Abdomen:  Brown.  Genitalia  as  in  h'igure 
5g   with   three   small    pairs   of   anteriorly 


directed,  lightly  scleroti/.ed  arms.  Sper- 
mathecae  small,  ovoid,  subequal  wdth 
short  necks. 

M\Li..  -Siiuilar  to  the  female  with  the 
following  differences:  smaller;  flagellum 
(Milir-el\  brown;  legs  more  diffused  vsdth 
l)i(>\Mi.  Genitalia  shape  and  form  like  that 
of  (\  hunacnUttd  (Fig.  4f ) ;  aedeagus  as  in 
j'igure  (ie  with  basal  arms  apparently  sep- 
arated, and  an  anteriorlv  directed  point 
on  the  anterior  membrane. 


Sept.  197-5 


;k()(,\:\.  wiktii:  ci-.matoi' 


285 


Distribution. —  Aj-i/oiin,  !\'o\v  Mexico. 
Utah  (locality  recoi-ds  plotted  in  l''i,mire 
7). 

IVPES. —  Female  lee  lotApe,  )  leniale 
paralectotyi^es,  (iraiid  C^aiiNon,  Coconino 
Co.,  Arizona,  C.  F.  Adams,  (KU),  here 
designated. 

Specimens  ex.\mined. —  From  the  fol- 
lowing localities: 

ARIZONA:  Apachf  Co..  Spniigeiville 
(Wirth).  I  male  (P);  Cocomno  Co.,  Grand  Can- 
yon (Adams,  types).  4  females  (KII)  (P).  NEW 
"MEXICO:  Taos  Co.,  Rio  Grande  (Wirth),  3  fe- 
males (P).  UTAH:  Cache  Co.,  Hyrum  (Grogan). 
I  females  (S);  Box  Elder  Co..  Brigham  City 
(Hardy,  Stains.  USU).  I  male,  1  female  (SJ. 
1  female  (P);  Weber  Co..  Huntsville  (Hardy, 
USU).  1  female  (P);  Wasatch  Co..  Heber  City 
(Dreisbach).   I   female  (P). 

Discussion. —  The  syntype  series  was 
labeled  "G.  Zmii  R..  Ariz.,  7-27."  which 
Adams  ])ublished  as  "Grand  Canon,  Ari- 
zona" for  the  type  locality. 


Clinohelca  usingeri  Wirth 
(Fig.  2b,  d.  h;  6f) 

Clinohelea    usingeri    Wirth,    1952:    209     (female; 
Cahfornia);  Wirth.  1965:   136  (distribution). 

Diagnosis. —    Distinguished     from    all 


other  N(Nn-(  li(  (Tuiohelca  by  the  two- 
spotled  winn  and  the  legs  mainly  yellow 
with  distal  li, df  of  hind  femur  and  apices 
of  tihiae  hrowii:  males  with  basal  anus 
of  aed(>agns   intact. 

Fe.m.\li:.—  Wing  length  2.65  (2.50-2.74, 
11  -  4)  mm;  breadth  0.76  (0.70-0.81,  n 
3)  mm. 

Head:  Brown.  Antenna  (Fig.  2b)  mod- 
erately slender;  basal  one-half  of  proximal 
flagellomere  lighter  brown  than  remain- 
der of  flagellum;  AP  17-10-9-9-9-9-9-10- 
23-22-22-22-20  n  =  3);  AR  1.33  ri.29- 
1.35,  n  --  3).  Palpus  with  3rd  segment 
longer   than    5th;    PR   3.33    (3.17-3.67,   n 

3  ).  Mandible  like  that  of  C.  bimaculata 
(Fig.  4c). 

Thorax:  Mesonotum,  scutellum,  post- 
scutellum  dark  brown.  Legs  (Fig.  2d) 
yellow;  proximal  one-third  of  fore  coxa, 
most  of  mid  and  hind  coxae,  distal  one- 
half  of  hind  femur,  femorotibial  areas  of 
mid  and  hind  fore  legs,  apices  of  tibiae, 
and  distal  4  tarsomeres  brown.  Wing  like 
that  of  C.  bimaculata  (Fig.  4d).  Halter 
})ale  to  whitish. 

Abdomen:  Brown.  Spermathecae  (Fig. 
2h)  small,  spheroid,  subequal  with  short 
necks. 


Fig.   6.     Aedeagi    of   male    Clinohelea-.    a,    C.    curriei;    b,    C.    nubifera;    c,    C.    pseudonubifera;    d,    C. 
bimaculata;    e.   C.   dimidiata;    f.   C.   usingeri. 


286 


(,REAT  BASIN   NATURALIST 


Vol.  35,  No.  3 


Male. —  Similar  to  female  with  the  fol- 
lowing differences:  smaller;  flatrellum 
entirely  brown;  femora  and  tibiae  entirely 
brown.  Genitalia  shape  and  form  like  that 
of  C.  bimaculata  (Fig.  4f ) ;  aedeagus  as  in 
Figure  6f. 

Distribution. —  Arizona,  (California 
(locality   records   j)lotted  in   Figure   7). 

Type. —  Holoty]:)e,  female,  Black  Lake 
Canyon.  San  Luis  Obispo  Co.,  California. 
22  August  1943,  W.  W.  Wirth  (Type  no. 
59949,  USNM). 

Specimens  examined. —  From  the  fol- 
lowing localities: 

ARIZONA:  Cochise  Co.,  Sunnvside  Canyon 
(Hardy,  KU).  1  male.  3  females.  CALIFORNIA: 
San  Diego  Co..  Live  Oak  Park  (Melander)  1 
female  (P);  San  Luis  Obispo  Co.,  Black  Lake 
Canyon  (Wirth,  type  series).  1  male.  2  females 
(P),  3  females   (S). 

Discussion. —  Present  records  indicate 
that  C.  dimidiata  is  an  inhabitant  of  the 
Great  Basin  and  the  LT]:)per  Colorado  Pla- 
teau, while  C.  usingeri  is  an  inhabitant  of 
the  Mojave  and  Sonoran  deserts.  Further 
collecting  is  necessary  to  determine  wheth- 
er the  geographic  ranges  of  these  two 
closely  related  s])ecies  oyerlap,  or  if  they 
are  separated  by  altitude. 


Cliuohclca  longitheca  Grogan 
and  Wirth  n.  sp. 

(Fig.  5c,  f,  i) 

Diagnosis. —  Distinguished  from  all 
other  Nearctic  CUnohelea  by  the  very 
large,  unequal,  elongate,  ellipsoid  sper- 
mathecae,  the  legs  mainly  yellow  with 
hind  tibia  and  distal  fourth  of  hind  femur 
brown,  and  the  two-spotted  vyings. 

Female  holotype. —  Wing  length  2.00 
mm;  breadth  0.61  mm. 

Head:  Brown;  frontoclypeus  lighter 
brown.  Antenna  (Fig.  5c)  slender, 
brown;  AP  17-11-10-10-10-10-10-11-21- 
21-21-20-20;  AR  1.17.  Palpus  brown;  3rd 
segment  longer  than  5th;  PR  3.11.  Man- 
dible like  that  of  C.  bimaculata  (Fig.  4c). 

Thorax:  Mesonotum,  scutellum,  post- 
scutellum  dark  brownish  black.  Legs  (Fig. 
5f)  yellow;  most  of  mid  and  hind  coxae, 
distal  one-fourth  of  hind  femur,  hind  tib- 
ia, and  distal  3  tarsomeres  brown;  distal 
one-fourth  of  fore  tibia  light  brown.  Wing 
like  that  of  C.  bimaculata  (Fig.  4d)  with 
an  infuscation  over  1st  radial  cell  and  just 
before   tip   of   costa.    Halter  light   brown. 

Abdomen:  Brown.  Spermathecae  (Fig. 
5i)  very  large,  unequal,  elongate  ellipsoid. 

Male. —  Lin  known. 


C.   bimaculata 
C.  dimidiata 
C.   usi  ngeri 
C.    lonqitheca 


Fig.   7.     North   American   locality  records  for  Clinohrlra  of  the  himaculata  group. 


Sei)t.  1975 


CROGAN.  WIKTH:   C1:HAT0P0G0NIDAE 


287 


Etymology. —  The  name  longitheca  is 
derived  from  the  Latiii  longus  (long)  and 
t/ieca  (sac)  and  refers  to  the  very  large, 
elongate,  ellipsoid  sj^ermathecae  that  are 
characteristic  of  this  sj)ecies. 

Variation. —  The  following  characters 
were  recorded  for  the  single  female  to])o- 
t^•])e:  wing  length  1.97  mm;  hreadth  ().()! 
mm.  AR'i.36.  PR  2.89.  CR  0.84.  I'he 
general  coloration  is  like  that  of  the  liolo- 
type. 

Distribution. —  Florida  (type  locality 
plotted  in  Figure  7  ) . 

Types. —  Female  holotype,  1  female 
})aratype  (S),  A.  &  M.  Riological  Station, 
Blackwater  River  State  Forest,  Santa  Rosa 
Co.,  Florida,  21  May  1971,  G.  B.  Fairchild, 
hlack  light  trap  (T\pe  no.  66496,  USNM). 

Literature  Cited 

Ad.-mnis.  C.  F.  1903.  Dipterological  contributions. 
Kansas  Univ.  Sci.  Bull.  2:21-47. 

Chan,  K.  L.,  and  E.  J.  LeRoux.  1965.  Descrip- 
tion of  Forcipomyia  (Neoforcipomyia)  saun- 
dersi  n.  sp.  and  redescription  of  Forcipomyia 
(Neoforci])07nyia)  ccjues  (Johannson)  (Dip- 
tera:  Ceratopogonidae)  with  an  account  of 
the  digestive  and  reproductive  systems.  Phy- 
toprotection  46: 74- 1 04. 

CoQUiLLETT.  D.  W.  1905.  New  Nematocerous 
Diptera  from  North  America.  Jour  New 
York  Ent.  Soc.  13:56-69. 

DowNEs,  J.  A.  1960.  Feeding  and  mating,  and 
their     interrelationshiji     in     the     insectivorous 


(>eratoi)ogoninae  (Diptera).  Verh.  XI  Int. 
Kongr.    Ent.    Vienna    1:618. 

.      1971.     The     ecology     of    blood    sucking 

Diptera:  an  evolutionary  perspective.  Pages 
232-258  in  A.  M.  Fallis,  Ecology  and  physi- 
ology of  parasites,  a  symposium.  Univ. 
Toronto  Press. 

JoHANNSEN.  O.  A.  1943.  A  generic  synopsis  of 
the  Ceratopogonidae  (Heleidae)  of  the  Ameri- 
cas, a  bibliography,  and  a  list  of  the  North 
American  species.  Ann.  Ent.  Soc.  Amer.  36: 
763-791. 

.  1952.  Guide  to  the  insects  of  Connecti- 
cut. Part  6.  The  Diptera  or  true  flies. 
Fasc.  5.  Midges  and  gnats.  Heleidae  (Cera- 
topogonidae). Bull.  Conn.  St.  Geol.  Nat. 
Hist.  Surv.  80:    149-175. 

KiEFFER,  .1.  J.  1917.  Chironomides  d'Amerique 
conserves  au  Musee  National  Hongrois  de 
Budapest.  Budapest  Magyar  Nemzeti  Muz., 
Ann.  Hist.  Nat.  15:292-364. 

LoKW,  H.  1861.  Diptera  Americae  septentriona- 
lis  indigena.  Centuria  prima.  Berlin  Ent. 
Ztschr.  5:307-359. 

Mali.och,  J.  R.  1914.  Notes  on  North  Ameri- 
can Diptera.  Bull.  Illinois  St.  Lab.  Nat.  Hist. 
10:213-243. 

.      1915.     Tlie   Chironomidae   or   midges   of 

Illinois.  Bull.  Illinois  St.  Lab.  Nat.  Hist.  10: 
275-543. 

Snodgrass,  R.  E.  1957.  A  revised  interpreta- 
tion of  the  external  reproductive  organs  of 
male  insects.    Smithson.  Misc.  Colls.  135:1-60. 

WiRTii,  W.  W.  1951.  New  species  and  records 
of  Virginia  Heleidae.  Proc.  Ent.  Soc.  Wash- 
ington 53:313-326. 

.      ]  952.     The      Heleidae      of      California. 

Univ.    California    Publ.    Ent.    9:95-266. 

.  1965.  Family  Ceratopogonidae  (Helei- 
dae). Pages  121-142  in  A.  Stone  et  al.,  A 
catalog  of  the  Diptera  of  America  north  of 
Mexico.  U.S.  Dept.  Agr.  Handbook  276.  1696 
pp. 


BASIDOMYCETKS  THAI    DECAY  JUNIPERS  IN  ARIZONA' 


R.  L.  Gilheitsoir  and  .) .  F.  Liiulsey- 


Abstract. —  Twenty-seven  spin  ies  dl  \v(i 
in  Arizona.  A  key  to  the  spei  i(>s.  iies(  i-i|)t  ii 
vide.l.  Pymfonirs  dcmidnjjii.  Daralra  junipri 
licartwood    in    living    tiecs.      H yphodi'niut    il< 


rnuing 

an.l    Ii 

/.  Pruia 

ticoUi  ( 


:et('s  are  reported  to  decay  junipers 
;s  of  nnrroscopic  characters  are  pro- 
id  I'hrllinus  texanus  cause  decay  of 
I.iuds.  is  described  as  new. 


Species  of  the  genus  .liniipcru\  of  th(^ 
Cupressaceae  are  among  the  most  (oii- 
spicLious  members  of  the  flora  of  Arizona. 
Eive  species  occur  in  the  state:  Junipcrus 
(Icppeana  Steud.  ( aUigator  juniper),  Juni- 
pcrus scopulorum  Sarg.  (Rocky  Mountain 
juniper),  Juniperus  monospernu/  ( En- 
gehn. )  Sarg.  (one-seed  juniper),  ,/z//?//;(77/\ 
ostcospcrrna  (Torr.)  Little  (Utah  juniper), 
and  Juniperus  communis  L.  (common 
jimiper).  The  first  four  sj)ecies  listed  at- 
tain tree  size  and,  with  pinyon  (Pinus 
cdulis  Engelm.),  occupy  a  large  area  of 
Arizona  in  the  pinyon- juiii|)(n-  woodland 
vegetation  type  from  kiOO  to  7,500  feet 
in  elevation  ( little.  100(S  ) .  I  lo\^ever,  juni- 
|)ers  occiu'  over  a  wider  eknational  range, 
from  the  u]iper  desert  grass  vegetation 
type  at  3,000  feet  to  the  ponderosa  })ine 
forest  at  approximately  8,500  feet.  The 
amuial  precijiitation  in  the  piii\(>ii-|iuiiper 
woodland  is  only  12-20  int  hes.  most  of 
which  occurs  in  the  summer  rainy  season 
in  .July  and  August  and  during  a  winter 
[)eriod  in  December  or  .lainiar\  .  During 
the  remainder  of  the  year  it  is  usually 
dry  in  the  zones  where  the  junipers  are 
found,  and  conditions  are  unfavorable  for 
the  development  of  wood-rotting  fungi. 

A  relatively  small  number  f)f  wood-rot- 
ting fungi  have  been  foinid  on  junipers. 
This  is  probably  due  to  the  pres(>iK(^  of 
fungistatic  compounds  in  juniper  wood. 
Species  of  oaks  and  pines,  connnonly  as- 
sociated with  junipers  and  expost'd  to  the 
same  environmental  ( oiiditions.  are  sid) 
strata  for  many  wood-rotting  fungi,  .luni- 
pers  are  widely  used  in  the  Southwest  for 
fence  posts  because  of  their  high  resis 
tance  to  decay  and  their  ahiuidancc  in 
most  areas.  .Junipers  are  aUd  (oninionix 
used  for  fireplace  wood. 

Hearti-dl  in  ]i\ing  .Arizona  juni[i('fs  is 
maiidy  attributed  to  loin-  spfu  ics  in  llic 
Polyporaceae.  These  are  PyrofoiiK-^  dcini- 
dofjii.  Dacdalea  juniperina.  /'on a  ri//i<j\a. 

^University    of    Arizona    .\KriculliMvil     KxpcniiuMil    SliiiKni 
-Department   of   Plant    Patliolopy,    L'liivoTsiiv    of   .\ri/niiii. 


and  PlicUiiius  texanus.  Fungi  that  decay 
dead  hraiuhes  on  living  trees  include 
Rxidiopsis  calcea,  LachneUa  alhoviolascens, 
and  Dendrothele  incrustans.  A  number  of 
wood-rotting  basidiomycetes  are  found  on 
dead,  fallen  junipers,  with  GloeopJiyllum 
tntJyeiuu.  Poria  ferox.  Coriolellus  sepium, 
<ni(l  Piuuis  fu/ridus  appearing  to  play 
major  roles.  Poria  apacheriensis  is  com- 
monh'  found  on  roots  and  the  root  crown 
region  of  dead  standing  junipers  in  south- 
eastern Arizona  and  may  cause  a  root-rot 
in  li\  ing  trees. 

Microscopic  characters  were  determined 
from  freehand  sections  or  crushed  tissue 
in  4  percent  KOH  and  phloxine  and  also 
in  Melzer's  reagent.  Capitalized  color 
names  are  from  Ridgway    (1912). 

The  oidy  ])revious  publication  dealing 
s|)e(  ifi(  ally  with  fungi  causing  decay  in 
iinii|)ers  is  that  of  Hedgcock  and  Long 
(  1912).  They  discussed  Fomes  juniperinus 
(von  Schrenk)  Sacc.  et  Syd.  and  Fomes 
earlei  (Murr. )  Sacc.  et  D.  Sacc.  (now  con- 
sidered synonyms  of  Pyrofomes  demi- 
doffii)  and  Fomes  texanus^  included  as 
Phellinus  texanus  in  this  paper. 

I  lost  indices  listing  decay  fungi  on  juni- 
pers in  the  Southwest  include  those  by 
Gilbertsoii.  Martin,  and  Lindsey  (1974) 
for  Arizona  and  the  U.S.  Department  of 
Agriculture  host  index  (Anonymous, 
HH)()i.  Shaw  (1973)  has  provided  a  host 
index  for  fungi  of  the  Pacific  Northwest. 
He  lists  Fomes  juniperinus  {=  Pyrofomes 
deniidofjii  ]  on  Juniperus  communis  in 
.Montana  and  Polyporus  hirtus  Quel,  on 
Juniju-rus  seopulorum  in  British  Columbia. 
TlH>se  are  the  only  host-fungus  relation- 
shi|)s  gi\en  by  Shaw  that  are  not  recorded 
for  tli(>  Southwest.  The  USDA  host  index 
lists  Fomes  roseus  (Alb.  et  Schw.  ex  Fr.) 
(  ke.  and  (  Oniophora  corrugis  Burt  on  juni- 
per in  .Arizona,  but  we  have  not  seen 
\(Mu  her  sjx'c  imens  to  support  the  records. 

uinal     .\ill<lo    .No.    2-KT. 


288 


Sept.  1975  (.ii.Hii?  r>().N.  ijxdm.'i  :  dkcay  fungi  289 

Key  to  1^'iiiigi  That  I)(Hay  .Iiiiiipers  in  Arizona 

1.  Hviiienojihore    ( oiisistiiii^    of    imited   tubes 2 

Hymenophore  Miiooth.  hydiiac  (H)us.   oi-  lamellate  —    16 

2(1).     Tissue    brown,    beconniiu    pernianetitlv    blackeiuMi    in    KOH  solution   .... 

_ Phellinus  texanus 

Tissue  white  or  pale  colored,  not  permanently    blackening   in    KOH   so- 
lution  3 

3(2).    Basidiocarps  centrally  stipitate Polyporus  arcularius 

Rasidioc  arps  s(^ssile  or  resujiinate 4 

4(3).    Basidiocar])s  with  brick  red  (onte.xt  tissue Pyrofomes  demidoffii 

Basidiocarps  with  context    not    brick  red 5 

5(4).    Basidiospores  thick  walled,  dextrinoid  in  Melzer's  reagent  

Fames  fraxinophilus 

Basidiospores  thin  w^allcd.  not  dextrinoid  in  Mel/.er's  reagent 6 

6(5).    Basidiocarps   always  resu])inate 7 

Basidiocarps  sessile  or  reflexed  11 

7(6).    Pore  surface  often  rose  jiiid^;  h^■phae  siniple-sej)tate.  no  clanip  comiec- 

tions  ])resent  ..  Poria  tarda 

Pore  surface  white  to  pale  buff;  generative  hyphae  with  clamp  connec- 
tions  ; 8 

8(7).    Basidiospores   subglobose   to    broadi^'  ellii)soid Poria  apachcriensis 

Basidiospores   cylindric    to    allantoid 9 

9(8).    Basidiospores  allantoid;  cystidia  present  .-. Poria  rimosa 

Basidiospores  cylinrlric;  c^■stidia  not  jiresent 10 

10(9).    Basidiospores  4-5.5  x  1.5-2  /an  .   Poria  sinuosa 

Basidif)S]X)res  <S  10  x  V3.5  /an  ;.... Poria  ferox 

11(6).    Pores  1-4  ]ier  mm,  circular  to  angular 12 

Pores  daedaloid  or  tubes  splitting  to  form  a  lamellate  hymenophore  14 

12(11).  Pore   surface   purplish:   c\stidia   abundant.   <ipically   incrusted   

-- Ilirschioporus  versatilis 

Pore  surface  cream  colored;  cystidia  not  |)resent 13 

1  3(12).  Basidiospores  12-13  x  5-6.5., /'.m;  pores   1-2   per  nnii 

....  Coriolcllus  hctcroniorphus 

Basidiospores  10-14  x  15-4.5  /an;  [jores  2-4  per  nun       .  Coriolcllus  sepium 

14(11).  Pores    large,    daedaloid;    tissue    (lale  buff  Daedalea  juniperina 

i^)res  r(>gnlai'.  tubes  splitting  to  Fur  in   a    lamellate   hymenophore;    tissue 

brown  15 

15(14) .  UppcM-  surface^  usually  distin(tl\-   zoiiate  with   bright  yellowish  or  red- 
dish brown  zones;  hymenophon^   strongl\-  lamellate  

(ilocophyllurri  sacpiarium 
Up[)er  surlai  e  .i/onatc  or   indistin(  tl\      /onale.     dull     brown;     liNineno- 

pliore  ponod  to  lamellate  Cjlorophylliun   trabeurn 

1()(1).     Basidiocarps  stipit.ite;   b\  nicnoplidro  r.!(liall\    lamellate  .._.  Panus  fulridus 

Basidiocar])s     sessile     or     r(>supiiiate;    hymenophore   smooth   to   hydna- 

ceous  17 


290  CHI-AT  BASIN  NATIRALIST  Vol.  35,  No.  3 

17(  1()).  Basidiocarps  c  ii|mlat('.  up  to  2  nun    wide LachncUa  alhoviolascens 

Basidiocarps   ccsupiiialc  18 

18(17K  HynuMiophnrc  smooth      ... 19 

H\nuMiophor(>  liMlnac  (M)ns -  25 

19(  18).  Basidia  \erticall\    .septate - ....-  Exidiopsis  calcca 

Basirlia  nonsoptatt'  20 

20(  19).  Hvnieiiial  surtacc  oin  accous;  hasidiosporcs    thick    walled,   dextrinoid   in 

Melzers  reai^ciit  ...  Coniophora  erctnop/nla 

H3'menial  suita((>  not  olivaceous;  basidiospores    thin     walled,    negative 

or    amyloid    in    Melzer's    reagent     21 

21  (20) .  Basidios]X)res    amyloid    in    Melzer's  reagent;  ac'anth()hy})hi(Ha  and  glo- 

eocystidia  present 22 

Basidios])ores  negative  in  Melzer's  reagent;    acanthohyphidia    and    glo- 

eocystidia  not  j)resent  23 

22(21  ).  Basidiospores  ().i-8  x  4-5  //m;  liymenial       surface       becoming      bluish 

gray ..  Alcurodiscus  lividocoeruleus 

Basidiospores  9.5-11.5  x  5.5-7  /-m;  h^^^lenoph()re  remaining  pale  buff  .... 

Alcurodiscus     cerrusatus 

2^(21  ).  Basidios])ores    globose    to   subglobose;   dendroh\  ])hidia   i)resent  

Dendrothele  incrustans 

Basidiospores  cylindric  to  ellij)Soid;   dendroh\phidia  absent  24 

24(23).  Cystidia  subulate,  thin  walled,  (S-8    /an  wide;  aggregates  of  golden  crys- 
talline material  present  ..  Hyphoderrna  pallidum 

Cystidia    cylindric.    with    a    rooted  base,  thick  walled,  10-12  /im  wide; 

golden   crystalhne   material   absent  Hyphoderrna  descrticola 

25(18).  tlymenophorc  pa|)illate.  papillae  with  an  amber-tolored,  beadlike  drop- 
let at  the  aj)ex  .. Dacryobolus  sudans 

Hymenophore   distinctly    hydnaceous  with  cylindric  or  flattened  teeth 26 

26 (25 J.  Sagittate  cysitha   present Hyphodontia  arguta 

Sagittate  cysti(ha  not  present  Hyphodontia  spathulata 

J'lxiDiopsis    CALCKA     ( Pers. )     \V(dls,    My-  basidia  to  55  /xm  long;  basidiospores  (Fig. 

cologia  5  5(4):   548.  19f)l.  Ic)  cylindric,  curved  to  slightly  allantoid, 

777r/,7;Wr/ r«/rr«  Pers..  Mv.oi.  Km.  I:  I -.1  182:2.  smooth,    hyahue,    germinating    by    repe- 
tition,  negative   in   Melzer's   reagent,    14- 

Basidiocarps  resupinate,  thin.  hard,  arid-  o()  ^  (^.^  .,-^^1 

wax^■.  at  first  de^eloi)ing  as  small  patches.  ^  Exidiopsis  calcea  decays  dead  branches 

then  coalescing  to  become  widely  ef  used.  ,,^-  ^^,^/^^^^^  ^^d  shrubs  and  occurs  from 

cracking   laterally   to   expose   the   substra-  „^,.  Sonoran  Desert  to  high-elevation  coni- 

tum;    hymenial   surface   white   to  grayish  ,•.,.   ,,,^.,.^j^      j^   ^^  associated  with  a   white 

white,  smooth,  shining;   margin  minutel\  .^  . 

fimbriate.     sf)rnetim(>s     abrupt;     subicidar  .,        ,  .  t-    n     /-      r-  i  n     -^ 

hvphae     of     tw.)      tvpes.      some      narrow.  \  "u.  her  specimen:    E.   R    Canfield,   / 1- 

branched     and    s,mious     1    1   ')    „ni    (ham  '''•  "'"   ''H".^''!"!"  juniper.  Scotia  Canyon, 

aseptate,     others     branched,     with     damp  Hu.i.hu.  a   \  Its..  Cochise  Co.,  AZ  (ARIZ) . 
connections,  3-5  "in  diain  (  b"ig.  la),  tlu^se 

giving  rise   to   basidia;    basidia    (Fig.    Ibi  Fachxi.lla      alboviolascens      (Alb.      et 

with    a    basal    clamp    connection,    hypo-  Sc  hw.   ex   Fr.  i    Fr.,   Sum.   Veg.   Scand.   p. 

basidia   subglobose   to   ellipsf)id,   becoming  ^65.      1849. 

longitudinallv    septate    and    foiUMelled    at  /,^,^-.^,  alhoriolasccus  Alb.  et  Schw.  e.x  Fr.,  Syst. 
maturity.   12-1)  x  r)-20  /uii.  the  tour  (^pi-  Myc.  1:  'ifj.  1822. 


Sept.  1^)75 


CII.HI'.RTSON.  I.INDSI:Y:    DF.CAY  IM'NOI 


291 


Fig.  1.  Rxidiopsis  calcca  (ERG  71-131),  a, 
subinilar   hyphae;    h.    hasidia;    c.   basidiospores. 

Basidiocai'ps  cupulate.  gregarif)iis.  de- 
\plopiiig  directly  from  the  substratiiiii 
with  a  small  patch  of  basal  white  my- 
( (diiim.  separate  or  crowded  together,  to 
1  mm  diam.  sessile;  outer  surface  co\  ered 
with  a  white,  wooly-matted  layer  of  hy- 
phae; hymenial  surface  pale  pinkish 
brown,  smooth,  obscured  on  (h'ied  spe- 
cimens by  the  inrolled  margin;  hyphal 
system  dimitic,  generative  hyphae  of  the 
outer  layer  (Fig.  2a  )  thick  walled,  hyaline, 
with  occasional  clamp  coimections.  6-7 
/un  diam,  some  with  an  extremely  narrow. 
caj)illarv  lumen;  skeletal  hyphae  (Fig.  2b) 
of  outer  layer  thick  walled,  aseptate,  to  7 
,"m  diam;  subhymenial  hyphae  moderately 
thick  walled,  with  abundant  clamp  con- 
nections, 2-4  /ail  diam;  basidia  (Fig.  2c ) 
broadly  clavate,  with  thickened  walls 
i()-<)()  i,m  long  and  13-l(i  /an  at  the  apex, 
four-sterigmate,  the  sterigmata  2-3  /im  in 
basal  (ham;  basidiospores  (Fig.  2d) 
broadly  ellipsoid,  hyaline,  negati\e  in 
.Melzer\s  reagent,  14-18  x  8.5- 10  /an. 

This  fimgus  is 
branches  of  a  mu 
(les,>rl  shrubs  nnd 
brown  rot. 

Voucher  spe(  imei 


i.r 


Ml  (in  d(Vi 
s(iiilb\\('st(n- 
i.ilcil     with 


RLC;  10193.  on  o 


seed  juniper.  Black  Oak  Cemetery,  (^anelr 
R.L.    Santa   Cruz  Co..   A/,    f  ART/ ) . 

■I    Cnl- 


(  ioNIOI'lIOHA      l.KI.MOl' 

III.A      1 

.inds. 

bcrts..    iM\cotaxon    1: 

-SI).     1 

0  7'). 

Basifhocarj)s     Irai^il 

'.     eas 

\\     se[ 

occurring  in  small   pa 

tclics  ( 

r  of  111- 

cm;    hymenal    surbn 

'    srno< 

dh.    \n 

Brownish  Olixc  from 

inass('( 

basid 

ited. 


as  they  mature;  hymenial  layer  soft  and 
flo(  (  os(>  oyer  a  white  arachnoid  subiculum; 
margin  with  fine,  white  mycelial  strands 
radiating  from  subiculum;  subicular  hy- 
phae simple-septate,  hyaline,  often  lightly 
incrusted,  some  (Fig.  3a)  thin  walled,  2-5 
/an  diam,  others  thin  to  thick  walled  (Fig. 
3b),  to  10  /im  diam;  cystidia  none;  basidia 
(Fig.  3c)  utriform  to  clavate,  usually  sin- 
uous, 40-(S()  X  6-10  /im,  four-sterigmate,  the 
sterigmata  to  7  /an  long;  basidiospores 
(Fig.  3d)  brownish  olive  in  mass,  pale 
yellow  in  KOH,  cyanophilous,  dextrinoid 
in  Melzer's  reagent,  thick  walled,  ellipsoid 
to  subglobose,  9-11  x  5-8.5  /xm,  with  an 
apical  germ  })ore  and  prominent  peglike 
apiculus. 

Coniophora  cremophila  has  also  been 
found  on  several  Sonoran  Desert  trees  and 
shrubs  and  is  associated  with  a  brown  rot. 

Voucher  specimen:  RLG  7400.  on  one- 
seed  juniper.  Gallinas  Mts.,  Lhicoln  Co., 
NM   (ARIZ). 

Aij:i!rodiscus  CKRUssATUs  (Bres.)  Hoelin. 
et  Litsch.,  K.  Acad.  Wiss.  Wien  Math.- 
Nal.   Kl.  Sitzungsb.  116:    807.   1907. 

Corticium    cerussalum    Bres.,    Fung.    Trid.    II.    37. 

1892. 


il    I. 


Fig    2.     Lac/tnrlla         alboviolascens  (RLG 

101931,   a.    grnerative   tiyphae;    b.   skeletal  hypha; 
.    hasidia;    d.    basidiospores. 


292 


(iHi:.\T  BASIN  NATlKALIsT 


Vol.  35,  No.  3 


Biisidiocarps  in  small  |)<it(li(vs.  IxHoniiiii!, 
(•(tiifliuMit.  cffiisCHl.  (  i-ackitiLi  (l('('|)l^  witli 
age;  li\  iiicnial  siirlaoM  ream  lodull  \\liit('; 
liyj)lial  N\si(Mii  UKiiioinit  i( .  coinposcd  of 
contorted,  thill  walled  li\  [iliae  ( Fig.  4a) 
2-4  /'111  diaiii.  with  .ihiiiidaiit  (lamp  con- 
nections; gloeocystidia  (  I'ig.  I  hi  emhed- 
ded.  cylinch'ic.  sinuous,  with  i-(4ra(ti\(> 
oily  ((Hiteiits.  some  moiiililorm.  10-70  x 
10-1  ^  //in;  weakh  positixc  in  sulphoheir/.- 
aldehyd(>;  acalltlloph^  ses  (  Mg.  l-c )  api- 
cally  thick  walled,  thin  walled  at  the  base, 
cylindric.  30-55  x  >l  /nn;  with  a  basal 
clamj)  connection;  mature  hasidia  (Fig. 
4d)  four-sterigmatc.  5-9  x  45-hO  /an; 
basidiospores  ( Fig.  4e)  broadly  cylindric 
to  ellii)soid.  thin  walled,  smooth,  amyloid 
in  -Melzer's  reagent,  inostly  collajised  or 
fragmenterl.   9-11    x   5-7   /an. 

AIriir()(lis(  us  ct-russa/us  grows  on  dead 
branch(>s  of  other  shruhs  and  tre(>s  in  the 
Southwest.  It  is  associated  with  a  white 
rot. 

Vcnu  htu"  s])ecimen:  RFG  8551.  on  one- 
seed  junij^er.  Cliiricahna  Nat.  Monnment. 
Chiricahna  Mts.,  Cochise  Co..  AZ  fARIZ). 

ALF.riu)i)isc:rs    li\  nj()f:()KKrM:is    (  Karst. ) 
Femke.  Can.  .T.  Rot.   12:  252.  1964. 

Corliciuni     UvidocoerulcuTn     Karst.,     Not.     Siilsk. 
Faun,  et  IHor.  Fonn.  Forli.  9:    570.    1868. 

Rasidiocarj)s  resupinate.  originating  as 
small,  separate  ])atclies,  then  becoming 
confluent  and  widely  effused,  w-axy;  mar- 
gin abrupt  and  fertile;  hymenial  surface 
cream  colored  or  Pale  Pinkish  Buff  to  blu- 
ish gra}'  (Light  Neutral  Gray  to  Dark 
Plumbeus),  often  rimose  with  age;  hyphal 
system  monomitic;  subicular  hyphae  (Fig. 
5a)  thin  walled,  with  clamp  connections, 
3-6//mdiam;  gloeocvstidia  (Fig.  5b)  abun- 
dant, positive  in  sulphobenzaldehyde,  em- 
bedded or  slightly  projecting,  some  moni- 
liform  or  mammillate,  20-90  x  6-F3  /i.m; 
acanthophyses  (Fig.  5c )  abundant,  acule- 
ate o\(>r  the  terminal  [)ortioii.  thin  to 
thick  walled  at  the  ajKw.  with  a  basal 
clamp  connection.  15-^)0  x  ^)-7  /an;  hasidia 
(Fig.  5d)  clavate.  -10-50  x  10-11  /an.  ste- 
rigmata  not  seen;  basidiospores  (  |'"ig.  5e) 
broadly  ( ylindrir  to  ellipsoid.  Inaline. 
smooth,  amyloid  in  Mel/er's  reagent,  thin 
walled  and  collapsing  readily.  f)-S  x  ').5-5 
/'.m. 

This  fungus  occurs  thronghonl  the 
Rocky  Mountain  region  on  niaii\  (onifers. 
It  is  associated   with  ;i   \\hil('  i-ot. 


\^)n(her  specimens:  RLG  10777,  on  al- 
ligator iunij)er,  Turkey  Creek,  Chiricahna 
\lts.,  Cochise  Co.,  AZ;  ERC  71-234,  on 
alligator  juniper.  General  Hitchcock  Pic- 
nic Area,  Santa  Catalina  Mts.,  Pima  Co., 
AZ   (ARIZ). 


Di 


xDHorin-.LK       ijxcRusTANs        ( Lcmke) 
Hike.    !\>rso()nia     ^:     MM^     1965. 


■Ucurocorliciuin    incrus/ans    Lcmke,    Can.    J.    Bot. 


42:    7' 
Basidi 


')f)4. 


|)s  small,  usually  in  patches 
tip  to  5  mm  wide,  sometimes  confluent; 
margin  abru])t,  fertile;  hymenial  surface 
smooth,  \^hite  to  Cartridge  Buff  or  cine- 
reous, rimose  on  drying;  subicular  hj'phae 
(  Fig.  ()a  )  slender,  with  frequent  branch- 
ing, with  inconspicuous  clamp  connections, 
1-2.5   /an   diani;    cystidia   absent;   hyaline 


oias_ 


Fig.  3.  Coniophora  eremophila  (RLG  7400), 
a.  narrow  subicular  hyphae;  b,  broacl  subicular 
li>pha;  c.  basidia;  d.  basidiospores. 

Fig.  4.  Aleurodiscus  cerrusatus  (RLG  8551), 
a,  subicular  hyphae;  b,  gloeocj'Stidia;  c,  acantho- 
physes; d,  basidia;  e.  basidiospores. 

Fig.  5.  Aleurodiscus  Uvidocoeruleus  (RLG 
10777),  a,  subicular  hypha;  b,  gloeocystidia;  c, 
ai  anthophyses;    d,    immatui-e  -basidia;    e,    basidio- 


Sept.  1075 


(;ir,Bi:HTs()%'. 


NDSl.^  :    ni.CAl    I'UNGI 


293 


(londrohy|)lii(lia  i  Fig.  ()1)  i  ahiitidaiil  iti  h^ 
menial  region,  ultimate  hiaiic  lies  |  /,iii 
(liam  or  less;  hasidia  i  I''ig.  fxl  i  (la\at(>  lo 
cvlindric.  some  snmiIIcii  al  the  hase.  de- 
veloping from  imbedded  hasidioles  (Fig. 
6c),  30-60  X  8-10  /-ni.  four-storigniate,  with 
a  basal  clamp  connection;  basidiospores 
(Fig.  6e)  hyaline,  smooth,  negative  in 
Melzer's  reagent,  globose  to  suhglohose. 
9-11  X  8-9.5  /-m.  sometimes  adhofing  in 
groups  of  two  to  four. 

Dendrotlicle  'uKrustdiis  aj)parently  util- 
izes bark  tissue  as  a  substratum.  It  also 
occurs  on  bark  of  se^•eral  other  southern 
Arizona  trees.  inchuHng  oaks  and  Arizona 
madroiK*  [Arhutiis  arlznnica  ( Cjray ) 
Sarg.).  Lendse  (  l<)6  1i  reports  this  fungus 
(as  Aleurocorticium  inr rustans }  from  sev- 
eral localities  in  the  Pacific  Northwest, 
but  it  has  not  been  rejiorted  previously 
from  the  Southwest. 

Voucher  spe<im«Mi:   P.1X.  1  ()()2().  on  hark 


living 


Ch 


illigator  juniper,  Turkey  Creek, 
1  Mts..  Cochise  Co..  AZ  (ARIZ). 


^smm 


Fig.  6.  Dendrothele  incruslans  (RLG  10020). 
a,  subicular  hyphae;  h.  (Iciidrohypliidia;  c,  basi 
dioles;  d,  mature  hasidia;  e.  Ijasidiospoies.  some 
in  pairs  and  tetrads. 

Fig.  7.  Dacryobolits  sudans;  (RLG  0^1<).  9944. 
and  10322).  a,  subicular  liypliae;  b.  fragment  of 
thick-walled  hypha;  c.  cvstidia:  d.  hasidia;  e. 
basidiospores. 


1). 


\(^HYOH()H 


^^•DANS   (Fr.)   Fr.,  Summa. 
404.  1849. 


Veg.  Scan. 

Hydnum  sudans  Fr.,  Syst.  Myc.   1:   425.   1821. 

Basidiocar])s  resupinate,  adnate,  effused 
up  to  6  cm;  hymenial  surface  papillose, 
Cream  Color  to  Warm  Buff;  papillae 
hemispherical  and  wartlike  to  conical, 
each  terminated  by  a  drop  of  amber, 
viscous  licpiid  that  dries  to  form  a  beadlike 
apex;  pa])illae  also  appearing  as  small 
(raters  where  the  exudate  has  broken 
away;  subicular  hyphae  (Fig.  7a)  thin  to 
thick  walled,  with  clamp  connections, 
fragments  of  thick  walled  hyphae  (Fig. 
7b)  appearmg  aseptate,  1.5-5  /mi  diam; 
cvstidia  (Fig.  7c)  thin  walled,  nonseptate 
or  with  clamps  and  simple  septa,  clustered 
at  apices  of  papillae,  70-90  x  3-6  /xm; 
hasidia  (Fig.  7d)  narrowly  clavate,  20-25 
X  3-5.5  /an,  four-sterigmate,  with  a  basal 
(lamp  connection;  basidiospores  (Fig.  7e) 
narrowly  allantoid,  hyaline,  smooth,  neg- 
ative in  Melzer's  reagent,  5-7  x  1-1.5  /j,m. 

Docryoholus  xudans  is  associated  with  a 
distinctive  brown  cubical  pocket  rot. 

Voucher  s])ecimen:  RLG  9944,  on  one- 
seed  juniper.  Sycamore  Canyon,  Atascosa 
Mts.,  Santa  Cruz  Co.,  AZ  (ARIZ). 


jESERTicoL.^     Gilberts,     et 


HyPlIODI.IiMA 

Linds..  sp.  no\ . 

Basidiocarpus  effusus.  mollis;  hy:nenium  laevis, 
inibaliiuis,  cystidiis;  hyphis  systematis  mono- 
mitico;  hyphis  suhiculis  tenuitunicatis,  3-5  nm 
diam;  hyphis  fibulatis;  cystidia  nimierosa, 
subulata,  90-125  x  6-13  /im,  base  ramosa,  radi- 
cata;  hasidia  clavata,  constrictione  media,  20- 
30  X  6-7  ,am.  4  stcrigmatibus;  basidiosporae 
hyalinae,  laeves,  non  amyloideae,  6-7  x  3-4 
/an.  HOLOTYPUS:  in  ligno  Juniperus  dep- 
peana  Steud.,  Gardner  Canyon.  Santa  Rita 
Mts..  Santa  Cruz  County,  AZ,  USA,  leg.  R.  L. 
Gilhertsoii.  no.  10921;  in  herb.  Nat.  Fungus 
Collections.   Beltsville,  MD,  USA    (BPI). 

Basidiocarps  effused,  in  small,  confluent 
patches,  soft,  easily  separated;  hymenial 
surface  smooth.  Light  Buff  to  Pale  Pinkish 
Buff,  becoming  rimose  on  drying,  cystid- 
i.ilc  under  a  M)X  lens,  cystidia  appearing 
pale  golflen  or  amher  due  to  apical  incrus- 
tation; hyjihal  system  monomitic;  subi- 
<  ular  hyphae  (Fig.  8a)  thin  to  moderately 
iln(  k  walled.  ( losely  septate,  with  clamp 
(oniK^c  lion^  al  all  septa,  3-5  /i,m  diam; 
(\slidia  ( l*'ig.  8b)  abundant,  subulate, 
thin    to    moderateh'    thick    walled,    many 


294 


GREAT  BASIN  N ATLtr,a^li_sT 


Vol.  35,  No.  3 


Voiuher  s|)ccimeu:  RLG  10921,  on  al- 
ligator junij)er,  (jardner  Canyon,  Santa 
Rita  Mts.,  Santa  Cruz  Co.,  AZ  (TYPE). 


\^;J^=^ 


Fig.  8.     Hyphoderma  deserticola  (RLG 

10921),  a,  subicular  hyphae;  b,  cystidia;  c,  basidia; 
d.  basidiospores. 

with  a  branched,  rooted  base,  90-125  x 
6-13  /.im,  most  apically  incrusted  with 
fine,  pale  golden  crystals  that  dissolve 
readily  in  KOH;  basidia  (Fig.  8c)  clavate, 
with  a  median  constriction,  20-30  x  6-7 
/im,  four-sterigmate,  occurring  in  candela- 
brums,  with  a  basal  clamp  connection; 
basidiospores  (Fig.  8d )  hyaline,  smooth, 
cylindric-ellipsoid,  negative  in  Melzer's 
reagent.  6-7  x  3-4  /vm. 

Basidiocarps  of  Hyphodcrnia  deserticola 
are  macroscopically  and  microscopically 
similar  to  those  of  H.  argillaccuryi  (Bres. ) 
Donk.  Cystidia  of  H.  deserticola  are  more 
thick  walled  and  less  swollen  at  the  base 
than  those  of  H.  argillaceurn  and  many  are 
branched  and  rooted  at  the  base.  The  ap- 
ical incrustation  on  the  cystidia  is  best 
seen  in  water  mounts  and  dissolves  rapidly 
in  KOH.  Basidiospores  of  H.  argillaceurn 
are  wider  (4-5  ,um)  than  those  of  //.  des- 
erticola, appearing  more  ellipsoid  than  cy- 
lindric.  Hyphoderma        macedonicurri 

(Bres.)  Donk,  as  described  by  Eriksson 
and  Ryvarden  (1975),  is  also  similar  but 
has  thin-walled,  nonrooted  cystidia  and 
narrower  spores  (2-2.5  n.m)  than  H.  des- 
erticola. 


Hyphodkhma    p.\llidum 
Fungus  27:   15.  1957. 


(Bres.)     Donk, 


Corticium    pallidum    Bres.,    Fung.    Trid.    2:     59. 
1898. 

Basidiocarps  effused  up  to  10  cm,  soft, 
thin;  hymenial  surface  Tilluel-Buff  to 
Pale  Vinaceous-Faw^n,  smooth,  speckled 
with  golden  brown  under  a  30X  lens; 
margin  thinning  out,  fertile;  hyphal  sys- 
tem monomitic;  subicular  hyphae  (Fig. 
9a )  with  clamp  connections,  2.5-4  /xm 
diam;  spherical  to  irregularly  shaped 
masses  of  golden  brown  crystalline  ma- 
terial (Fig.  9d)  abundant  throughout  su- 
bicidum,  to  30  /an  wide;  cystidia  (Fig.  9b) 
narrowly  subulate,  thin  walled,  not  in- 
crusted,  40-50  /an  long  and  6-8  /<.m  wide 
at  the  swollen  base;  basidia  (Fig.  9c) 
clavate,  four-sterigmate,  25-35  x  6-7  /xm, 
with  a  basal  clamp  connection;  basidio- 
spores (Fig.  9e)  hyaline,  smooth,  slightly 
curved,  cylindric,  negative  in  Melzer's  re- 
agent, 8-11  x  3-4  jiva. 

Hyphoderma  pallidum  is  associated  with 
a  white  rot  of  conifer  slash  in  the  South- 
west. 

Voucher  sj)ecimen;  RLG  10778,  on  al- 
ligator juniper,  Turkev  Creek,  Chiricahua 
Mts.,    Cochise   Co.,   AZ    (ARIZ). 

Hyphodontia    arguta    (Fr.)    J.    Erikss., 
Symb.  Rot.  Upsal.  16:  1.  p.  104.  1958. 

Hydum  argutum  Fr.,  Syst.  Myc.   1:   424.   1821. 

Basidiocarj)s  becoming  widely  effused, 
hymenial  surface  hydnaceous.  Light  Buff 
to  Pinkish  Buff,  the  teeth  usually  crowded, 
cylindric  and  tapering  at  the  apex,  simple 
or  branched,  up  to  3  mm  long,  the  apices 
finely  tomentose  or  fimbriate;  subiculum 
cream  colored  to  pale  buff,  very  soft  and 
thin;  hyphal  system  monomitic;  subicu- 
lar hyphae  (Fig.  10a)  thin  walled,  with 
frequent  branching,  with  abundant  clamp 
connections,  2-3  n.m  diam;  cystidia  of  two 
types,  some  (Fig.  10c)  sagittate,  apically 
incrusted,  15-30  /tm  long,  projecting  up  to 
18  //.m.  5-6  nm  diam  at  the  swollen  base, 
slender,  stalkHke  part  1.5-2  /-.m  diam, 
incrusted  aj)ex  about  2.5  /.cm  diam,  other 
cysticha  (Fig.  10b)  not  incrusted,  with 
slight  terminal  or  intercalary  swellings, 
f)ften  with  clam[)  connections,  thin  walled, 


Sept.  1975 


GILBKKTSON.  LINDSF.Y:    UKCAY  FUNGI 


295 


40-50  X  3-6  /irn  and  projecting  to  25  /i.m; 
basidia  (Fig.  lOd)  clavate,  with  a  median 
constriction,  four-sterigmate,  10-21  x  4-5 
/im;  basidiospores  (Fig.  lOe)  ovoid  to  el- 
lipsoid, hyaline,  smooth,  negative  in  Mel- 
zer's    reagent,    4-6    x    2.5-4    /im. 

Hyphodontia  arguta  has  been  foiuid  on 
both  conifers  and  hardwoods  in  the  South 
west  and  is  associated  with  a  white  rot. 

Voucher  specimen:  RLG  8325,  on  al- 
ligator juniper,  Carr  Camon,  Huachuca 
Mts.,  Cochise  Co..  AZ    (ARIZ). 

Hyphodonti.a  sp.\THiiL.'\T.'\  (Sclirad.  ex 
Fr.)  Parm.,  Conspect.  Svst.  Cort.  p.  123. 
1968. 

Hrdnum  spathulaturn   Schrad. 
1:   423.    1821. 

Basidiocarps  becoming  widely  effused; 
hymenial      surface     hydnaceous,      cream 


e.\  Fr.,  Syst.  Myc. 


Fig.  9.  Hyphoderma  pallidum  (RLG  10778). 
a,  subicular  hyphae;  b,  cystidia;  c,  basidia;  d, 
massos  of  aniori)hous  niattM-ial  from  .subiruluni;  e. 
basidiospores. 

Fig.  10.  Hyphodontia  arguta  (RLG  8  325).  a. 
subicular  hyhae;  h.  iioiiini  rusted  cystidia;  c,  sa- 
gittate, incinstcd  ivslidia;  d.  basidia;  e.  basidio- 
spores. 

Fig.  11.  Hyphodontia  spathulata  (RLG  9795, 
9949,  and  10131),  a,  subicular  hypliae;  b,  capi- 
tate  cystidia;    c.    l)asidia;    d.    basidiospores. 


(olorefl  to  Light  Buff,  the  teeth  cyHndric 
to  fhittened  and  tending  to  be  confluent, 
forming  elongated  ridges;  hyphal  system 
monomitic;  subicular  hyphae  (Fig.  11a) 
thin  to  moderately  thick  walled,  with 
abiuidaiit  clamp  coimections,  with  fre- 
{juont  branching  and  irregular  swellings 
and  constrictions,  2-4  ,um  diam;  cystidia 
(Fig.  lib)  capitate,  25-30  x  3-4  ^um, 
slightly  projecting,  also  hyphoid  end  cells 
tufted  at  ajjices  of  teeth,  these  lightly  in- 
crusted;  basidia  (Fig.  lie)  clavate  with  a 
median  constriction,  15-17  x  5-6  jjjn,  four- 
sterigmate;  basidiospores  (Fig.  lid)  ovoid 
to  subglobose,  hyaline,  negative  in  Mel- 
zer's  reagent,  smooth,  4-5  x  3-4  /xm. 

Hyphodontia  spathulata  is  associated 
with  a  white  rot  of  both  conifers  and  hard- 
woods in  the  Southwest. 

Voucher  specimen:  RLG  10131,  on  al- 
ligator juniper,  Parker  Can^'on,  Huachuca 
Mts.,  Santa  Cruz  Co.,  AZ  (ARIZ). 

Phellinus  texanus  (Murr.)  Gilberts, 
et  Canf.,  Mycologia  65:    1304.   1972. 

Pyropolyporus    texanus    Murr.,    N.    Amer.    FI.    9: 
104.   1908. 

Basidiocarps  perennial,  sessile,  ungulate, 
up  to  15  cm  wide;  upper  surface  at  first 
pale  brown,  matted-tomentose,  becoming 
blackened  and  deeply  rimose  with  age  and 
weathering,  sulcate;  margin  rounded,  pale 
brown  and  tomentose;  pore  surface  pale 
brown  (Buckthorn  Brown  to  Mummy 
Brown),  smooth,  the  pores  4-6  per  mm; 
dissepiments  finely  tomentose,  entire; 
context  hard  and  woody,  yellowish  brown, 
appearing  mottled  with  streaks  of  paler 
tissue;  tid)e  layers  stratified.  Buckthorn 
Brown,  becoming  stuffed  with  light-col- 
ored mycelium;  sections  permanently 
darkening  in  KOH  solution;  some  contex- 
tual hy])hae  thin  walled  and  hyaline  to 
yellowish,  with  occasional  septa  and  rare 
branching,  2-5  ,am  diam  (Fig.  12a),  others 
thick  walled,  yellowdsh  brown,  aseptate, 
with  rare  branching,  3-5  fim  diam  (Fig. 
1 2b ) ;  hyjihae  in  the  pale  colored  areas 
with  a  parallel  arrangement,  easily  sepa- 
rated, hyphae  in  the  darker  areas  densely 
interwoven,  contorted,  difficult  to  separ- 
ate; tramal  hyphae  interwoven,  pale  yel- 
lowish, with  slightly  thickened  walls  and 
orcasional  septa,  2-4  ^m  diam;  setae  (Fig. 
12d)  few,  slightly  thick  walled  to  thick 
walled,  ^^ith  an  inflated  base  to  10  ^m 
diam  and  a  slender  apical  portion,  25-60 


296 


GRKAT  BASIN  NATl'RALIST 


Vol.  35,  No.  3 


Fig.  12.  Phr/limjs  texamis  (RLG  6959),  a, 
thin-wallcd  (ontcxtual  li\'i)hae;  1).  tliick-wallcd 
contextual  liyphac;  c  tlnii-walled  cystidioid  hy- 
phal   ends;   d.   setae;    e.    hasidia;    f.    hasidiospores. 

/an  long;  setae  apparently  developing 
from  hyaline,  thin  walled  cystidioid  hy- 
phal  ends  (Fig.  12c)  similar  in  shaj)e  and 
size  to  the  setae  and  conmion  in  all  sec- 
tions; basidia  (Fig.  12e)  broadly  clavate 
to  ovoid,  fonr-sterigmate,  17-25  x  8.5-12 
/-m,  the  sterigmata  slender,  to  4  /.an  long; 
basidiospores  (Fig.  12f)  subglobose,  hya- 
line, strongly  dextrinoid  in  Melzer's  re- 
agent and  with  distinctly  thickened  walls 
when  mature,  7-0  x  6.5-9  /an. 

Phellifjus  te.irnius  occurs  on  many  So- 
noran  Desert  trees  and  shrid^s  including 
cacti,  and  causes  a  white  heartrot  of  li\  ing 
plants. 

Voucher  sj)ecimens;  HIXt  ()M5*^K  on  al- 
ligator juniper,  Sponsellor  Lake  Rd.,  Nava- 
jo Co..  AZ;  RLG  7406,  on  one-seed  juni- 
per, Gallitias  Mts.,  Lincoln  Co.,  NM; 
RLG  8042,  on  Rocky  Mountain  juniper. 
Doc  Long  Picnic  Area,  Saiuha  Mts., 
Bernahllo  Co.,  NM;  P.  D.  Keener  74.  on 
alligator  juniper,  rVIingus  Mt..  Ya^a])ai 
Co..  AZ;  K.  J.  Martin  481  on  alligator 
juniper,  Rucker  Campground,  Chiricahua 
Mts..  Cochise  Co.,  AZ  (ARIZ);  W.  H. 
Long,   327.  TX   (TYPE)    (NY). 


CORIOLELLL'S  HKTI.HOAU  )KPI  H 

et  Sing..  Ann.  M\c.   ^9:   ()(). 


,  (  Fr. 

1941 


Bond. 


Daedalea    heteromorpha    Fr., 
1821. 


Syst.    Myc.    1:    340. 


Basidiocarps  animal,  sessile,  (d'fused-re- 
flexed.  or  resupinate;  uj)per  surface  on  re- 
flexed  specimens  cream  colored  to  pale 
brownish,  glabrous  to  finely  tomentose; 
pore  surface  Light  Buff  to  Pinkish  Buff; 
pores  regular  to  slightly  daedaloid.  1-2 
I)er  mm,  or  in  some  specimens  most  1a-  (nw 
1  mm  diam;  dissepiments  thi(  k  iuul  to 
mentose  at  first,  becotning  thin  and  split 


ting  with  age;  margin  thinning  out  or 
abrupt,  tomentose;  context  \Qry  thin,  pale 
buff;  tube  layer  concolorous  \^'ith  context; 
liyphal  system  dimitic;  generative  hyphae 
(Fig.  13a)  hyaline,  thin  to  thick  walled, 
with  abundant  clamj:)  connections,  3-4  /im 
diam;  skeletal  hyphae  (Fig.  13b)  hyaline, 
thick  walled,  aseptate,  with  occasional 
branching,  3-7  /an  diam;  cystidia  none; 
basidia  (Fig.  13c)  clavate,  four-sterigmate, 
30-40  X  9-11  /an,  with  a  basal  clamp  con- 
nection; basidiospores  (Fig.  13d)  hyaline, 
broadly  cylindric.  narrowed  and  curved 
near  the  apiculus.  smooth,  negative  in 
Melzer's   reagent,    10-13   x   5-7   /mi. 

(^oriolcUus  hctcromorphus  causes  a 
brown  cubical  rot.  usually  of  conifers. 
Thin,  white  mycelial  felts  develop  in  the 
shrinkage  cracks  of  the  decayed  wood. 
Resuj)inate  specimens  with  large  pores 
may  be  confused  with  Daedalea  juni- 
pcrina.  The  later  can  be  distinguished  by 
its  much  smaller  spores  and  basidia. 

Voucher  specimens:  RLG  9306,  9945, 
and  9952,  on  one-seed  juniper,  Sycamore 
Canvon,  Atascosa  Mts.,  Santa  Cruz  Co., 
AZ   (ARIZ). 

CoRioLELLus  SEPIUM  (Berk.)  Murr.,  Tor- 
rey  Bot.  Club  Bull..  32:   481.   1905. 

Trat)irlrs   sepiutn    Berk..    London    J.    Bot.    6:    322. 
1847. 

Basidiocarjis  annual,  sessile,  effused-re- 
flexed,  or  occasionally  resupinate,  pilei 
solitary  or  imbricate,  dimidiate  to  elon- 
gate, to  25  X  4  X  2  cm;  surface  of  the 
pileus  Light  Buff  to  Mummy  Brown  or 
blackening   with   age,   glabrous   or   finely 


Fig.   13.     Coiiolellus      heteromorphus       (RLG 
9952).   a.   generative  hyphae;   b,  skeletal   hyphae; 

(.   I)asidia;  d.  basidiospores. 


Sept.  1975 


GILBERTSON.  LINDSKY:    DECAY  FUNGI 


297 


D.m:i)aei.,> 

Fl.  9:    12- 


.riTNiPERiNA  Murr.,  N.   Amer. 
1908. 


Fig.  14.  Coriolellus  sepium  (RLG  10129).  a, 
contextual  generative  liyphae;  b.  contextual  skele- 
tal   hyphae;    c.    basidia;    d.    l)asidiospoies. 

tomentose,  azonate,  shallowl\'  sukate, 
smooth  or  slightly  rugose;  tiuirgiti  con- 
coloroiis,  rouiidecl,  fertile  or  narrowly 
sterile  below;  pore  surface  Cinnaniou 
Buff  to  Buckthorn  Brown,  the  pores  cir- 
cular to  angular,  2-3  per  mm,  the  dissep- 
iments thick,  entire,  eventually  becoming 
lacerate;  context  ivory,  azonate,  corky, 
to  4  mm  thick;  the  tube  layer  concolor- 
ous  and  continuous  with  context,  to  1  cm 
thick;  odor  sometimes  fragrant;  hyphal 
system  dimitic;  contextual  generative  hy- 
[)hae  (Fig.  14a)  hyaline  in  KOH,  mostly 
thin  walled,  rareh'  branched,  with  abun- 
dant clamp  connections,  2.5-5.5  /ini  diami; 
contextual  skeletal  hyphae  (Fig.  14b) 
thick  walled,  aseptate,  3-5.5  n.m  diam; 
tramal  hyphae  similar;  cystidia  none; 
basidia  (Fig.  14c)  clavate,  four-sterigmate, 
20-25  X  8-10  /-,m;  basidiospores  (Fig.  14d) 
hyaline,  negati^■e  in  Melzer's  reagent,  cy- 
lindric,  9.5-14  x  3-4  /an. 

Coriolellus  scpiu/n  is  associated  with  a 
brown  cubital  rot  with  thin,  arachnoid 
mycelial  felts. 

Voucher  sjx'cinuMis:  RL(  i  72()l.()n  al- 
ligator juniper,  Rucker  Canytjii.  (diirica- 
hua  Mts.,  Cochise  Co.,  AZ;^RLG  10045, 
on  juniper  post,  Washington  Camp.  Pata- 
gonia Mts..  Santa  Cruz  Co..  A/;  RI.G 
10048,  on  alligator  JLuiipcr.  Montezuma 
Pass,  Huachuca  Mts..  (^odiise  Co..  AZ; 
RI/i  10198,  on  juniper  post,  Sunn\'side, 
Cochise  Co.,  A/;  mXr  10129,  on  alligator 
juniper,  Parker  Canvon  Lake,  C>)chise 
Co.,  AZ;  RLG  108  38,  on  alligator  juniper, 
Indian  Creek,  Animas  Mts.,  Hidalgo  Co., 
NM;  ERC  70-L  on  alliuator  juniper.  Rust- 
ler Park,  Chiri(ahii,i  Mtv.  Co.  Ium"  Co. 
AZ;  ERC  71-129.  on  alligator  jumper-. 
Scotia  Canyon,  Ilii,i(lui(a  Mts..  {d.jiisc 
Co.,  AZ  (ARIZ ) 


Basidiocarps  annual  or  perennial,  ef- 
fused-reflexed  or  resupinate;  pilei  solitary 
or  imbricate,  f)fteii  laterally  fused,  to  5 
X  10  x  9  cm;  surface  of  the  pileus  weath- 
ering Light  Buff  to  Cinnamon  Buff,  to 
gray  or  blackish,  indistinctly  zonate,  be- 
coming glabrous  and  incrusted;  pore  sur- 
face Light  Buff,  rough,  pores  large,  dae- 
daloid,  often  more  than  1  mm  diam;  dis- 
sepiments thick,  entire,  splitting  with  age 
to  form  lamellae  or  spines;  context  yel- 
lowish ivory,  soft-corky,  faintly  zonate,  to 
2  cm  thick;  tube  layers  concolorous  and 
( ontinuous  with  context,  to  6  cm  long; 
hvjihal  system  dimitic;  contextual  gener- 
ative hyphae  (Fig.  15a)  thin  walled,  with 
rare  branching,  with  abundant  clamp  con- 
nections, 3-fi  //m  diam;  contextual  skeletal 
hvphae  (Fig.  15b)  hyaline  in  KOH,  thick 
walled,  with  rare  branching,  aseptate,  3-7 
//m  diam;  tramal  hyphae  similar;  cystidia 
none;  basidia  (Fig.  15c)  clavate,  20-25  x 
6-7     nin.     four-sterigmate;     basidiospores 


K.    !■ 


Dacdalca  juniper ina  (JPL  328,  RLG 
IDfjOh.  a.  contextual  generative  liyphae:  b,  con- 
icxiM.il  skeletal  hvphae;  c.  basidia;  d.  basidiospores. 
Fig.  16.  Fomes  fraxinophilus  (ERC  71-25), 
a.  subicular  generative  hyphae;  b,  subicular  skele- 
tal hyphae;  c.  basidia;  d,  basidiospores. 


298 


GREAT  BASIN  NATURALIST 


Vol.  35,  No.  3 


(Fig.  15d)  hyaline,  negati-ve  in  Melzer's 
reagent,  smooth,  cylinclric,  6.5-9  x  2.5- 
3.5  /.(.m. 

Daedaica  jiniipcriiia  is  associated  with 
a  brown  cubical  heartrot  of  living  juni- 
pers. Thick,  buff  colored  mycelial  felts 
develop  in  shrinkage  cracks  of  decayed 
wood. 

Voucher  specimens:  RLG  6940,  on  one- 
seed  juniper.  Salt  River  Canyon,  Gila  Co., 
AZ;  RLG  9945  and  ERC  7^1-26,  on  one- 
seed  juniper,  Sycamore  Canyon,  Atascosa 
Mts.,  Santa  Cruz  Co.,  AZ;  P.  D.  Keener, 
on  alligator  juniper,  Mingus  Mt.,  Yavapai 
Co.,  AZ;  RLG  10604,  on  Utah  juniper, 
Seegmuller  Mt.,  Mohave  Co.,  AZ;  ,IPL 
328,  on  alligator  juniper,  Rucker  Cannon, 
Chiricahua  Mts.,  Cochise  Co.,  AZ;  KJM 
335,  on  one-seed  juniper.  Canyon  del  Oro, 
Santa  Catalina  Mts.,  Pinal  Co.,  AZ 
(ARIZ). 

FoMEs  FRAXiNOPHiLus  (Pk.)  Cke.,  Gre- 
villea  15:  51.  1886. 

Polyporus   fraxinophilus   Pk..    Bot.   Gaz.    7:    43-44. 
■l882. 

Basidiocarps  perennial,  resupinate,  ef- 
fused to  6  cm;  pore  surface  Pale  Ochrac- 
eous  Buff  to  Pinkish  Buff,  the  pores  3-4 
per  mm;  dissepiments  thick,  minutely  to- 
mentose;  margin  narrowly  sterile,  0.5 
mm  wide.  Light  Buff;  subiculum  ])ale 
buff,  to  1.5  mm  thick;  hyphal  system 
dimitic;  generative  hyphae  (Fig.  16a)  in 
subiculum  thin  walled,  2-3  /mi  diam,  with 
inconspicuous  clamp  connections,  these 
more  apparent  in  marginal  tissue;  subi- 
cular  skeletal  hyphae  (Fig.  16b)  hyaline, 
moderately  thick  walled  to  thick  walled, 
with  occasional  branching,  aseptate  or 
rarely  with  simple  septa,  2.5-5  /'.m  diam; 
tramal  hyphae  similar  to  subicular  skele- 
tal hyphae  but  mosth'  2-3  /mi  diam;  cys- 
tidia  none;  basidia  (Fig.  16c)  broadly 
clavate,  four-sterigmate,  20-25  x  9-11  /'m; 
basidiospores  (Fig.  16d)  broadly  ellipsoid 
to  subglobose,  truncate  at  the  apex,  hya- 
line, dextrinoid  in  Melzer's  reagent,  9-10.5 
x  6.5-8  jim,  thick  walled  at  maturity  with 
a  germ  pore  at  the  truncate  apex. 

Fomes  fraxinophilus  causes  a  wbit(» 
heartrot  of  living  trees  and  is  common  in 
southern  Arizona  on  Frnxinus  rclufina 
Torr.  (Arizona  ash).  Small  resupinate 
basidiocarps  are  also  commonly  found  on 
ash.  Clamp  corniections  of  F.  frarinoph- 
ilus  are  abundant   and   conspicuous   in  (h- 


karyotic   cultures   of   the   fungus   but   are 
difficult  to  discern  in  basidiocarp  tissue. 

Voucher  specimens:  ERC  71-25,  on 
one-seed  juniper.  Sycamore  Canyon,  Atas- 
cosa Mts.,  Santa  Cruz  Co.,  AZ;  J.  L.  Lowe 
9091,  on  one-seed  juniper,  Silver  Creek 
Rd.,  Chiricahua  Mts.,  Cochise  Co.,  AZ 
(ARIZ). 

Gloeophyllum    saepiarium     (Wulf.    ex 
Fr.)    Karst.,  Finl.  Hattsv.  2:    80.    1879. 

Daedalea   sepiaria   Wulf.    e.v.    Fr.,    Syst.    Myc.    1: 
333.  1821. 

Basidiocarps  annual,  effused-reflexed, 
sessile,  or  occasionally  resupinate;  upper 
surface  Warm  Sepia  to  Bister  or  blacken- 
ing, hirsute  to  coarsely  strigose,  concen- 
trically zonate;  lower  surface  Sudan 
Brown  to  Amber  Brown,  pores  present  at 
the  margin  of  some  specimens,  but  tubes 
typically  splitting  to  form  a  radially 
lamellate  hymenophore;  context  Honey 
Yellow  to  Clay  Color,  darkening  to  Sepia, 
azonate,  up  to  4  mm  thick;  hyphal  system 
dimitic;  contextual  generative  hyphae 
(Fig.  17a)  hyaline,  thin  walled,  with 
abundant  clamp  connections,  3-5  /im  diam; 
contextual  skeletal  hyphae  (Fig.  17b) 
thick  walled,  pale  yellowish  brown,  asep- 
tate, with  occasional  branching,  2.5-5  /im 
diam;  cystidia  (Fig.  17c)  thin  to  thick 
walled,  cylindric,  not  incrusted,  to  85  /xm 
long  and  2.5-6  /.i.m  diam;  basidia  (Fig. 
17d)  narrowly  clavate,  with  a  greatly 
elongated  base,  60-80  x  7-8  /mi;  basidio- 
spores hyaline,  smooth,  cylindric,  slightly 
curved,  negati\e  in  Melzer's  reagent,  8.5- 
11  X  3-4  /mi. 

G/oeophyiluni  saepiarium  causes  a 
brown  cubical  rot  of  conifers  and  hard- 
woods in  Arizona  but  is  rarely  found  on 
junijier. 

Voucher  sj)ecimen:  RLG  10918,  on  al- 
ligator juniper,  Gardner  Canyon,  Santa 
Rita  Mts.,  Santa  Cruz  Co.,  AZ  (ARIZ). 

Gloeophyllum  trabeum    (Pers.  ex  Fr.) 
Murr.,  N.  Amer.  Fl.  9:   129.  1908. 

Daedalea  Irabea  Pers.  ex  Fr.,  Syst.  Myc.   1:   335. 
1821. 

Basidiocarps  annual,  sessile,  effused-re- 
flexed or  occasionally  resupinate  in  early 
stages  of  do\elopment;  pilei  dimidiate  to 
elongated,  often  imbricate  and  confluent; 
upper  surface  yellowish  -  brown,  weath- 
ering to  tan  or  grayish,  tomentose  and  be- 


Sept.  1975 


GII.BF.RTSON.    LTiyDSK^':     DKCAY    FUNGI 


299 


Fig.  17.  Gloephyllurn  sacpiariutn  ( RIXi 
10918),  a,  contextual  generative  hyphae;  b.  con- 
textual skeletal  hyphae;  c,  cystidia;  d.  basidia;  e, 
basidiospores. 

Fig.   18.     Gloephyllurn  Irabeum  ( RLCi 

10128),  a,  contextual  generative  hyphae;  b.  con- 
textual skeletal  hypliae;  c,  cystidia;  d,  basidia;  e, 
basidiospores. 

coming  glabrous.  f;iiiitl\  zoiiate;  lower 
surface  i)ale  brownish,  poroid  at  first  and 
in  some  specimens  remaining  poroid  witli 
])ores  1-^  j)er  nmi.  radially  elongated,  in 
others  becoming  ratlially  lamellate  l)\ 
splitting  of  dissepiments;  (ontext  pale 
brown,  soft-felt},  a/ojiate,  up  to  12  nnn 
thick;  tube  layer  or  lamellae  ]iale  brown, 
up  to  4  mm  thick;  hyphal  system  diniitic; 
contextual  generative  hyjihae  (Fig.  IHa) 
hyaline,  thin  walled,  with  abundant  (lam]) 
connections,  2-15  nn\  diani;  contextual 
skeletal  hy])hae  (Fig.  IHb)  pale  \ellowish 
brown,  thick  walled,  aseptate,  with  rare 
branching,  3-5.5  /im  diam;  tramal  hyphae 
similar;  cystidia  (Fig.  18c)  cylindric,  thin 
walled,  35-50  x  3-5  /an;  basidia  (Fig.  18d) 
clavate,  foi.u--sterigmate,  30-45  x  5-7  //.m; 
basidiospores  (Fig.  18e)  tylindric  .  h\  aline, 
smooth,  negative  in  Mc'l/cr's  icagent, 
7.5-9  X   3-15   nm. 

Glocophyllum      Iralx-iini      is     a^so(  iated 


with  <i  ')r-o\\n  cubical  rot.  Mycelial  felts 
do  iiol  d('\('lo[)  in  the  decayed  wood. 
Cj.  trdbciun  is  the  most  common  wood- 
rotting  fiuigus  on  dciid  fallen  jinhpers  in 
southern  Arizona  and  is  also  found  as  a 
wootl-rotting  fungus  on  houses  in  Tucson. 
Voucher  specimens:  RLG  10128,  on  al- 
ligator junifier.  Parker  Lake,  Huachuca 
Mts.,  Codns,"  Co..  AZ;  RLG  10132,  on  al- 
ligator juniper.  Sunnyside,  Huachuca 
Mts.,  Cochise  Co.,  AZ;  RLG  1(J204,  on  al- 
ligator- juniper.  Scotia  Canyon,  Huachuca 
Alts.,  Cochise  Co.,  AZ  (ARIZ). 


I  luiscmoiH) 
zeki,  .1.  .lap 


IS     \  IIHSATILL^ 

Rot.  20:   288. 


(Rerk.)    Ima- 
1945. 

r's  J.   Bot.    1:    150. 


Tramctes  rerscitilis  Ber'k.,  Hookc 
1842. 

Rasidioc  ar[)s  resupinate  to  effused-re- 
flexed,  annual;  {)ore  surface  purplish 
when  fresh,  dull  purplish  brown  on  age 
and  drying,  the  pores  circular  to  angular, 
1-2  per  mm;  hyphal  system  dimitic;  su- 
bicular  generative  hyphae  (Fig.  19a)  thin 
walled,  2-3.5  /mi  diam,  with  inconspicu- 
ous clamp  connections;  tramal  hyphae 
similar;  contextual  skeletal  hyphae  (Fig. 
19b)  hyaline,  thick  walled,  aseptate  or 
with  rare  clamp  connections,  wdth  rare 
branching,  2.5-5  /im  diam;  cystidia  (Fig. 
19c)  abundant,  fusoid,  capita tely  incrus- 
ted,  20-30  x  3-5  /im;  basidia  (Fig.  19d) 
clavate,  four-sterigmate,  14-17  x  5-6  /im; 
basidiospores  (Fig.  19e)  cylindric,  hyaline, 
negative  in  Melzer's  reagent,  5.5-8  x  2-2.5 
/an. 

Ilirsrhioponis  versa  tills  causes  a  white 
pocket  rot  and  is  apparently  rare  in  the 
Southwest. 

A^oucher  siiecimens;  W.  FI.  Long  and 
C.  G.  Hedocock.  F.   P.  98/6.  on  one-seed 


Fig.  19.  Hirschiopnnis  vcrsatiUs  ('.ILL  11454). 
,1.  coiil'.'xtual  geiieiative  hyphae;  b,  contextual 
skeletal  hyphae;  c,  capitatelv  incrusted  cystidia; 
d.  basidia;  e.  basidiospores. 


300 


(;Ki:Ar  uasin  inatuuallsi' 


Vol.  35,  No.  3 


juniper,  Magdaleiia,  NM,  Type  ol  7m- 
rnetes  ruhricosa  Bres.  (BPI);  G.  G.  Hedg- 
cock,  F.  P.  18496,  on  one-seed  juniper. 
Gila  Nat.  Forest,  NM,  determined  as 
T.  ruhricosa  by  Bresadola  (BPI);  (i.  G. 
Hedgcock  and  W.  H.  Long.  F.  P.  10410. 
on  one-seed  juni])er,  Alogollon.  NM 
(BPI);  W.  H.  Long.  12704,  on  one-seed 
junij)er.  Winona,  (jxonino  (>»..  AZ 
ARIZ). 

PoLYPORUs    ARCuLARH's    Batscli    ex    Fr., 
Syst.  Myc.   L    342.   1821. 

Basidiocarps  annual,  centrally  stii)itate; 
pilei  circular,  solitary,  uj)  to  2.5  cm  diam 
and  0.3  cm  thick;  surface  of  the  pileus 
straw  colored  to  dark  brown,  a/.onate, 
glabrous,  smooth  to  rugose;  margin  ciliate. 
acute,  sterile  below;  stripe  central,  con- 
colorous  with  pileus,  glabrous,  up  to  3.5 
cm  long  and  0.4  cm  thick;  pore  surface 
cream  colored  to  buff,  dull,  rough,  the 
pores  large,  he.xagonal,  radially  aligned, 
1-2  per  mm,  the  dissepiments'  thin,  be- 
coming lacerate;  context  whitish  to  buff, 
azonate,  tough,  less  than  1  nun  thick;  tube 
layer  concolorous  and  continuous  with 
context,  up  to  2  mm  thick;  hyphal  sys- 
tem dimitic;  generative  hyphae  (Fig.  20b) 
hyaline  in  KOH,  thin  walled,  often 
branched,  with  abundant  clamp  connec- 
tions, 2.5-5  ,am  diam;  skeletal  hvphae 
(Fig.  20c,  Fig.  20d)  thick  walled,' asep- 
tate,  with  occasional  branching,  2-11  nxn 
diam;  tramal  hyphae  similar,  not  readily 
separable;  hyphae  on  pileus  surface  (Fig. 
20a)  slender,  thin  walled,  with  clamp 
connections,  1-1.5  /-.m  diam;  basidia   (Fig. 


Fig.  20.     Polyporus  arcularim    i.IPI,   278   and 

HLG  790^ j,  a.  livjihae  from  ])il('us  siirfact';  h. 
generative  hvpliac;  <.  broad  skeletal  livi)hao;  d. 
narrow     skeletal     lispliar.     ,..     Iwisidui;     I.     b.isidin 

S|l<)|cs 


20e)  four-sterigmate,  clavate,  25-35  x  5-6 
/an;  cystidia  none;  basidiospores  (Fig. 
2()f)  h^'aline,  negative  in  Melzer's  re- 
agent, smooth,  cylindric,  straight  or  slight- 
ly curved,  7-9  x  2.5-3.5  /tm. 

Poly  par  us  circular  ius  causes  a  white  rot 
and  is  especially  common  on  dead  oak 
wood  throughout  the  oak  woodland  vege- 
tation zone  in  southern  Arizona.  It  is  oc- 
casionall}'  found  on  other  substrata.  This 
report  is  based  on  a  field  observation  of 
P.  arcularius  on  one-seed  juniper  at 
Sunnyside,  Iluachuca  Mts.,  Cochise  Co., 
AZ  by  RLG. 

PoRiA  APACHKRiENsis  Gilberts,  et  Canf., 
Mycologia  65:    1117.  1973. 

Basidiocarps  annual,  effused  up  to  10 
cm.  adnate,  soft-fibrous;  pore  surface 
white  to  Cream  Color  or  Light  Buff; 
sterile  margin  tomentose;  pores  circular 
to  daedaloici,  up  to  1  nxn  diam,  mostly  2-3 
per  mm;  tube  layer  soft-fibrous,  white  to 
cream  colored,  up  to  2  mm  thick;  subi- 
culiun  soft,  white,  less  than  0.5  mm  thick; 
hyphal  system  monomitic;  subicular  hy- 
phae (Fig.  21  a  j  with  abundant  clamp  con- 
nections, thin  to  moderately  thick  walled, 
with  occasional  branching,  2-4  //.m  diam; 
tramal  hyphae  similar,  incrusted  at  dis- 
sei:)iment  edges;  cystidia  thin  walled,  of 
two  types,  some  acicular  or  cylindric 
(Fig.  21b),  smooth  to  lightly  incrusted, 
45-60  X  2.5-5  ,um,  others  capitate  (Fig. 
21c),  3-5  /un  diam  with  swollen  apex  up 
to  8  /im  diam,  40-50  /tm  long;  basidia 
(Fig.  21  d)  with  a  median  constriction, 
four-steriginate,  18-30  x  6-7  /xm;  basidio- 
spores (Fig.  21  e)  hyaline,  smooth,  nega- 
\\\i'  in  MeTzer's  reagent,  broadly  ellipsoid 
to  subglobose.  5-6.5  x  1.5-5.5  //in. 

Poria  apachcricusis  occurs  on  a  number 
of  Sonoran  Desert  plants,  including  the 
Saguaro  cactus,  Carnegiea  gigantea  (En- 
gelin. )  Britt.  et  Rose  (Lindsey  and  Gilbert- 
son.  1975),  and  causes  a  white  rot.  It 
I'ruitb  aroinid  the  base  of  recently  killed 
junipers  at  the  ground  line,  suggesting  it 
may  cause  a  root  rot  in  Hving  trees. 

Vou.her    spc(ini(Mis:     RLG    10047    and 
l"',KC     LSI.    (in    alligator    juniper,    Monte- 
/.inna    l*<rss.    (>)chise   Co.,    AZ    (ARIZ). 
PoHi A  1  I  iu).\  Long  et  Baxter,  Papers  Mich. 
Acad.  S(i.  25:   1  19.  1()40. 

Basidiix  ;ir[is  perennial,  becoming  wide- 
\\  fffuscil;  [lore  surface  (cartridge  Buff  to 
l^ndsish   P.utr  nr  I'ale  Orange  Yellow,  the 


Sr,.l 


501 


N 


u 

Fig.  21.  Porid  iiimchericfisis  iJPL  407  and 
409).  a.  sul)i(ular  liyphae;  b.  cvliiidiic  rvstidia;  c. 
capitate  cystidia;  d.  basidia;  e.  basidiospoies. 


pores  circular  to  .lugular.  4-()  per  tnni; 
margin  abrupt,  fertile  or  narrowly  sterile, 
blackening  with  age;  subiculum  less  than 
1  nnn  thick,  cream  colored  to  jjale  buff; 
tube  layers  stratified,  single  layers  u\)  to 
1  mm  thick;  hyphal  system  dimitic;  su- 
bicular  generative  hyphae  (Fig.  22a)  in- 
conspicuous, thin  walled,  with  clamp  con- 
nections. 2-3  ii.m  diam;  subicular  skeletal 
hyphae  (Fig.  22b)  hyaline,  thick  walled, 
aspetate,  with  rare  branching,  5-5  //.m 
diam;  tramal  hyphae  similar;  fusoid  cys- 
tidioles  (Fig.  22c )  present,  barely  pro- 
jecting, 11-18  .X  5.5-7  /an;  basidia  (Fig. 
22d),  broadly  clayate,  four-sterigmate. 
14-18  X  8-9  /(.m;  basidiospores  (Fig.  22e ) 
cylindric,  subfusiform,  hyaline,  smooth, 
negative  in  Melzer's  reagent.  8-10  x  5-4 
,um. 

Poria  ferox  causes  a  brown  c  ubical  rot 
with  consjHcuous  cream  colored  to  pale 
buff  mycelial  felts  in  the  shrinkage 
cracks. 

Voucher  specimens:  RLG  10126.  on  al- 
ligator iuniper.  Parker  CauAon,  Huacliuca 
Mts.,  Santa  Cruz  Co..  AZ;  RLG  10817 
and  10822,  on  alligator  juniper,  Indian 
Creek,  Animas  Mts..  Hidalgo  Count^■. 
NM;  RLG  10127.  on  alligator  juniper. 
Parker  Can^•oll  Lake.  Cochisf  Ca)..  AZ 
( ARIZ ) . 

PoRi.\  RiMos.y  ATurr..  Mycologia  12:  91. 
1920. 

Basidiocarps  perennial,  sometimes  de- 
veloping as  scattered  j)atclies  on  a  cottony 
mycelial  mat  f)n  the  surface  of  the  sub- 
stratum, often  becoming  rinios(>  with  age; 
pore  surface  Light  Ochraceous  Ruff  to 
Cinnamon  Buff,  pores  5-7  jier  mm.  angu- 
lar;   hyphal    pegs    present;    dissei)iments 


at  first  appearing  tottientose  under  a  30X 
lens,  llni  Is.  Ixm oming  thin  and  splitting 
with  age;  lube  layer  ivory  to  straw 
colored,  up  to  2  mm  thick;  context  con- 
colorous  with  tubes,  soft,  fibrous,  up  to  0.5 
nun  tln(  k.  taste  mild;  hy{)hal  system  di- 
mitic; subic  ular-  gcMierative  hyphae  (Fig. 
2  5a)  thin  wallecl,  with  abundant  clamp 
c ounce  tions.  often  branched,  2-5  yxn  diam; 
subicular  skeletal  hyphae  (Fig.  23b) 
thick  walled,  apparently  nonseptate,  2-3 
/'m  diam;  tramal  hyj)hae  predominantly 
thin  walled,  with  clamp  connections,  2-3 
;:x\\  diam;  cystidia  (Fig.  23c)  scarce  to 
conmion,  not  incrusted,  thin  walled,  hy- 
phoid,  up  to  2.5  /an  diam  and  projecting 
up  to  25  N.m  beyond  the  hymenium;  ba- 
sidia (Fig.  25d)  clavate,  four-sterigmate, 
10-13  X  4-6  ,"m;  basidiospores  (Fig.  23e) 
hyaline,  smooth,  negative  in  Melzer's  re- 
agent,  allantoid,   4-5.5    x    1-2   ,um. 

Poria  riifiosa  causes  a  white  heartrot 
of  living  trees  and  is  also  found  on  dead 
standing  and  fallen  junipers.  In  the  ad- 
vanced stages  the  rot  is  characterized  by 
small  em])ty  pockets.  It  has  a  wide  dis- 
tribution in  western  North  America  (Gil- 
bertson,  1961). 

Voucher  sjiecimens:  RLG  6961,  on  al- 
ligator juniper.  Sponsellor  Lake  Rd.,  Nava- 
jo Co..  AZ;  RLG  6963,  on  one-seed 
juniper,  between  Show  Low  and  Snow- 
flake,  Navajo  Co.,  AZ;  RLG  7560,  on 
cme-seed  juniper,  Stoneman  Lake  Rd.,  Co- 
conino Co..  AZ;  K.  D.  Butler,  on  alligator 
juniper,  Mingus  Mt.,  Yavapai  Co.,  AZ; 
.ILL  9106,  on  one-seed  juniper.  Silver 
Creek,  near  Portal,  Cochise  Co.,  AZ;  RLG 
10605,  on  LTtah  juniper,  Seegmuller  Mt. 
area  near  Wolf  Hole,  Mohave  Co.,  AZ 
(ARIZ). 

PoRi.'^  siNuos.^  (Fr.)  Cke.,  Grevillea  14: 
115.  1886. 

Pnlrporus  sinuosus   Fr..   Syst.   Myc.   1:    381.    1821. 

Basidiocarps  annual,  often  widely  ef- 
fused, tough,  corky,  easily  separable, 
taste  resinously  bitter;  margin  fertile  or 
narrowly  st(>rile.  Light  Buff,  soft,  fim- 
briate, to  1  nmi  wide;  pore  surface  Cream 
Color  or  <lr\ing  to  Cinnamon  Buff,  the 
tubes  to  \  mm  long,  the  pores  circular  to 
angular  oi-  siimous,  2-4  per  mm,  with 
thick,  entire  dissepiments  that  become  thin 
and  deeply  lacerate;  subiculum  whitish, 
corky,  azonale.  to  2  mm  thick;  hyphal  sys- 
tem dimitic;  subicular  generative  hyphae 


302 


GREAT  BASIN  NATITRALIST 


Vol.  35,  No.  3 


Fig.  22.  Poria  fcrox  (RLG  10817),  a,  subicu- 
lar  generative  hyphae;  b.  subicular  skeletal 
hyphae;  c,  fusoid  cystidioles;  d,  basidin;  e.  basidio- 
spores. 

Fig.  23.  Poria  rimosa  (RLG  6961  and  6963). 
a,  subicular  generative  hyphae;  b,  subicular 
skeletal  hj'phae;  c,  hyphoid  cystidia;  d,  basidia: 
e,  basidiospores. 

Fig.  24.  Poria  sinuoxa  (HHB  1133),  a.  subi- 
cular generative  hyphae;  b.  subicular  skeletal 
hyphae;  c.  fusoid  cystidioles;  d.  basidia;  e.  basidio- 
spores. 

Fig.  25.  Poria  tarda  (RLG  10135),  a.  subicu- 
lar hvphae;   b.   basidia;   c.   basidiospores. 

(Fig.  24a)  thin  walled,  rarely  branched, 
with  clamp  connections.  2-4  /.i.ni  diarn: 
subicular  skeletal  hyphae  (Fig.  24b) 
thick  walled,  aseptate,  rarely  branched, 
2-5  jjxn.  diam;  fusoid  cystidioles  (Fig.  24c) 
12-15  X  3-4  /-.m;  basidia  (Fig.  24d)  clavate, 
four-sterigniate,  with  a  basal  clam])  con- 
nection, 11-13  X  4.5-5  /.-.m;  basidiospores 
(Fig.  24e)  cylitidric.  some  slightly  curved, 
hyaline,  smooth.  n(\t(ativ(>  in  Melzer's  re- 
agent, 4-6  X  1.5  2  i>n\. 

Poria  si  nuosa  causes  i\  brown  cubic  al  rol 
of  conifer  logs  and  slash.  It  is  clisiriljni,.,! 
throughout  western  conilcrons   iof(vsts. 

Voucher  speciiiieii:  II.  M.  Burdsall  11  ^1 
on  Juuipcrus  s})..  Valoiu  ia  (^o..  Cibola  Nat. 
Forest  Rd.  No.  HSO  near  ( Irani s.  NM 
(ARIZj. 


Porta  ta]U)a  (Berk.)  Cke.,  Grevillea  14: 
109.  1S8(). 

Polyporus    tardus    Berk.,    London    J.    Bot.    4:    56. 
1845. 

Basidiocarps  annual,  usually  adnate, 
pore  surface  rose  pink  to  cream,  usually 
drying  Pinkish  Buff  to  Light  Buff;  tubes 
originating  as  isolated  cupules  and  then 
iniiting;  pores  3-5  per  mm;  sterile  margin 
usually  rather  wide,  thinning  out;  context 
white  to  cream,  soft,  thin,  Iwphal  system 
monomitic;  subicular  hyphae  (Fig.  25a) 
hyaline,  thin  walled,  simple-septate,  oc- 
casionally ampullate  at  the  septa,  fre- 
quently branched  at  right  angles,  some 
with  crystalline  incrustation,  2.5-6  jxva 
diam;  tramal  hyphae  similar;  cystidia 
none;  basidia  (Fig.  25b)  clavate,  four- 
sterigmate,  15-20  x  4-5  ixva;  basidiospores 
(Fig.  25c)  oblong  to  cylindric-ellipsoid, 
hyaline,  smooth,  negative  in  Melzer's 
reagent,  4-5  x  2-2.5  /im. 

Poria  tarda  is  a  common  fungus  in 
southern  Arizona  and  has  been  found  on 
dead  wood  of  many  trees  and  large  shrubs 
from  the  Sonoran  Desert  up  to  the  pond- 
erosa  pine  forest.  It  causes  a  white  rot. 

Voucher  specimens:  RLG  10135,  on  al- 
ligator juniper,  Sunnyside,  Huachuca 
Mts..  Cochise  Co.,  AZ;  RLG  10197,  on  al- 
ligator iimiper,  Canelo  Rd.,  Santa  Cruz 
Co.,  AZ;  FRC  71-118,  on  alligator  juniper, 
Suimyside,  Huachuca  Mts.,  AZ   (ARIZ). 

Pyrofomes  demidoffii  (Lev.)  Kotl.  et 
Pouz.,  Repert.  nov.  Spec.  Regn.  veg.  69: 
140.  1961. 

Polyporus  demidoffii  Lev.  in  Demidoff,  Voy.  Russ. 
Merid.  2:   92.   1842. 

Basidiocarps  ])erennial,  sessile,  solitary, 
tmgulate,  often  becoming  columnar,  to  15 
cm  wide,  7  cm  thick,  and  10  cm  high;  up- 
per surface  brownish  and  tomentose  in 
young  specimens,  becoming  blackened 
and  rimose  with  age,  concentrically  sul- 
cate;  margin  rounded.  Warm  Buff  to 
Ochraceous  Buff,  finely  tomentose  to 
blackened  and  rimose  in  older  specimens; 
pore  surface^  Light  Ochraceous  Buff  to 
Ocbracc'ons  I'uif.  smooth,  the  pores 
i-oinidc'(l.  J  )  |)C'i-  mm;  dissepiments  thick, 
entire;  (oiUc.xl  Orange  Ciimamon  to  Cin- 
namon Ridons.  woody,  azonate;  tube 
layers  Ochraceous  Buff  to  Antimony  Yel- 
low at  first,  e\entually  becoming  filled 
with  mycelium  and  concolorous  with  con- 


Sept.  1975 


(;ilbi:kts()in.  li;\'1) 


i)i;(;  x^   I'l'  ;\(; 


303 


'ig.  26.  Pyrofotncs  demuhffii  {«L(i  lUbOO). 
a.  liypliae  from  mycelial  felts;  b,  contextual 
skeletal  hyphae;  c,  much-branched  skeletal  hyphae 
from  trama;  d.  contextual  generative  hvphap;  e. 
fiisoul    fvstidioles:    f.   basidia;    g.    basidiospores 

tt^xt,  iiidisliiK  tl\-  stiiitifii'd.  each  hiA  or  to 
7  mm  tliick;  hy|)hal  system  dimitic;  con 
textual  generative  hy])hae  (Fig.  26d)  dil 
ficult  to  discern,  thin  walled,  hy^aline  in 
KOH  and  Melzer',s  reagent,  with  clamp 
connections,  2.5-4  /an  diam;  contextual 
skeletal  hyphae  ( Fig.  26b)  moderately 
thick  walled,  rarely  septate,  with  rare 
branching,  pale  brownish  in  KOH  and 
dextrinoid  in  mass  in  Melzer's  reagent, 
3-5.5  /an  diam;  tramal  tissue  similar  but 
with  some  much-branched  skeletal  h\phae 
(Fig.  26c),  2-5  /'m  diam;  tramal  tissue 
distinctly  dextrinoid  in  Melzer's  reagent; 
hyphae  of  mycelial  felts  (Fig.  26a)  in 
wood  frequently  branched,  some  thick 
walled,  aseptate.  1-^  /an  diam.  others  thin 
walled.  simple-sej)la1e  or  with  (xca^ional 
clamp  connections.  1.5-15  /an  diam;  in- 
conspicuous fusoid  cystidiolirs  (Fig.  26e)  in 
hymenium.  thin  walled,  not  incrusted, 
2()-30  X  3-5 /an;  basidia  (Fig.  26f)  broad  I  v 
clavate  from  a  narrow  base,  four-sterig- 
mate.  with  a  basal  (lamp  ( otmection.  22- 
35  X  8-10  /an;  basidiospores  i  Fi-.  26g  i 
pale  brownisli.  slightly  dextrinoid  in  \lel 
zer's    reagent.    ()\oid    lo    IxoiuJlx     ellipsoid 


or  tiiore  elongated,  angular,  thick  walled, 
most  truncate  at  apex  with  an  inconspicu- 
ous germ   ])ore,  6-12  x  5-7  /xm. 

Pyrojoiurs  dcinidoffii  is  probably  the 
most  important  heartrot  fungus  in  western 
junipers.  It  (  anses  a  white  rot  with  abun- 
dant my((>lial  lelts  in  the  decayed  wood. 
It  has  been  referred  to  as  Fomes  juniperi- 
nus  (yon  Schrenk)  Sacc.  et  Syd.  in  most 
American   literature. 

Voucher  specimens:  KLG  6960,  on  one- 
seed  juniper,  between  Show  Low  and 
Snowflake,  Navajo  Co.,  AZ;  RLG  7384 
and  7815,  and  ERC  71-28  and  71-326,  on 
one-seed  juniper,  Sycamore  Canyon,  Ata- 
scosa Mts.,  Santa  Cruz  Co.,  AZ;  RLG 
7559,  Stoneman  Lake  Rd.,  7  mi  E  of  High- 
way 79,  Coconino  Co.,  AZ;  RLG  7562,  on 
one-seed  juniper,  Dry  Creek,  6  mi  W  of 
Sedona,  Coconino  Co.,  AZ;  RLG  7890,  on 
Utah  juniper.  South  Rim,  Grand  Canyon 
Nat.  Park  Coconino  Co..  AZ;  RLG  9875, 
on  LTtah  juniper,  Mt.  Trumbull,  Mohave 
Co.,  AZ;  JLL  9070,  on  one-seed  juniper. 
Silver  Creek,  Chiricahua  Mts.,  Cochise 
Co.,  AZ  ( ARLZ ) . 


56.  1900. 

Rasidi.Harps 
to    gregarious; 
diam;    npper   >ii 
iiamou    Ikifl    to 
fibrillar  scales; 
to  Ruffy   Rrowi 


Rres.,  Fung.  Trid.  XL  p. 


(('ntrall\  stipitate,  single 
jjileus  ( irmlar,  0.5-3  cm 
ilacc  pale  brownish  (Cin- 
(>lay  Color)  with  radial 
sti})e  Light  Ruff  at  apex 
1  at  the  base,  glabrous  to 
minutely  pubescent  or  scaly%  to  3  mm 
diam  and  2  cm  long;  gills  pinkish  cream 
to  pale  buff  when  dried,  distant,  free  to 
adnate,  eflges  siimous  and  distinctly  granu- 
lose  under  a  30X  lens;  contextual  h}'phae 
variable,  some  (Fig.  27a  and  b)  simple- 
septate,  with  occasional  branching,  thin  to 
slighth'  thick  walled.  2.5-6  /an  diam,  others 
(Fig.  27c  I  very  thick  walled  to  almost 
solid,  aseptate.  rarely  branched,  5-9  /im 
diam;  |)letu-oc\  stidia  (Fig.  27d)  fusoid, 
barely  projecting,  35-60  x  6-8  /an;  cheilo- 
( ystidia  similar;  basidia  (Fig.  27e)  clav- 
ate, four-sterigmate,  simple-septate  at 
base,  38-60  x  9-13  /-m;  basidiospores  (Fig. 


2; 

li)    br( 

)adl^ 

r    c\ 

dindric,    si 

lightly 

curved. 

ii; 

valine. 

sm( 

.olh'. 

negative 

in  Melzer's  re- 

agent.  11 

2-16 

X  5. 

5-7  /an. 

Pr/rn/s 

fu/, 

'  vV/?/. 

s  causes  a 

browat 

cubical 

r-( 

)t  and  i 

onitt 

ion  finigii< 

;  on  dead  stand- 

ii 

igand  ! 

la  Me 

n  ini 

:ii[i(^rs  and 

also  on 

juniper 

fe 

■nee  po 

sts    1 

n  soi 

itliern  Arizona. 

304 


GREAT  BASIN  NATURALIST 


Vol.  35,  No.  3 


Fig.  27.  Panus  fulridus  (ERC  71-1^2  aiul 
71-158),  a,  thin-walled  contextual  livphao;  b. 
slightly  thick-walled  contextual  liyphae;  c,  thick 
walled  to  solid  contextual  liyphae;  d.  fusoid 
pl(;urocystidia;    e.    basidia:    f.    basidiosjiores. 

Voucher  specimens:  ERC  71-1  ^2,  on  al- 
ligator juniper,  Scotia  Canyon,  Huachiua 
Mts.,   Cochise   Co..   AZ;    RTX;    100  Ik   on 


junij)er  Nmkc 
Pataooiiia  \I|> 
10258,  on  jiii 
Canyon,  Ala^ 
AZVARIZ). 


Lr 


po^l.  VVashinoion  (Jamp, 
.  Santa  Cm/.  Co..  AZ;  RLG 
i\)cv  fence  post.  Sycamore 
nsa    Mt'^..   Santa   Cruz   Co., 


:r.\ti 


!RK    C 


I  ted 


Index    of    plant    diseases    in 
s.     TT.S.    Dept.   Agr.   Handb. 


Anony.aious.  1960. 
the  United  Sta 
No.  163.  VJl  p. 

Eriksson,  .!..  \xi)  L.  Rvvakdf.n.  1973.  The 
Corticiaceae  of  North  Europe.  Vol.  3.  Fungi- 
flora.     Oslo,   Norway,    pp.   287-546. 

Gir,BERTS0N,  R.  L.  1961.  Notes  on  western  poly- 
pores.    Pap.   Mich.   Acad.   Sci.   46:209217. 

Gir.BERTsoN.  R.  J,..  K.  ,T.  M.artiiv.  and  .1.  p. 
LiNDSEY.  1974.  Annotated  check  list  and 
host  index  for  Arizona  wood-rotting  fungi. 
Univ.  Ariz.  Agr.  Exp.  Sta.  Tech.  Bull.  209. 
48  p. 

Hedgcock.  G.  G..  and  W.  H.  I.ong.  1912.  Pre- 
liniinaiA"  notes  on  three  rots  of  juniper. 
Mycologin  4:109-1  13. 

IjEMKe,  p.  a.  1964.  The  genus  Aleurndiscux 
(sensLi  lafo)  in  Norlh  America.  Can.  .1.  Bot. 
42:72^-768. 

LiNDSEY,      ,T.      P.. 

'Wood-inhal) 
gnai'o   in   A 
I.iTir.K.    ]•:.    I... 
TT.S.    n.M)t. 


^ND 
tig 


RroGWAY.  R.  19 
nDineiu  laluie. 
by  the  author. 

Sii.vw,  C.  G.  197=; 
Pacific  NoT'tiiv 
Univ.   Agr.   Exj 


R.      L.      (ill.RERTSON.       1975. 

lomobasidionivcetes    on    sa- 
■izona.     Mycotaxon  2:83-103. 
,Tr.      1930.     Soufliweslern     frees. 
Agr.   Handb.   No.  9.     109  p. 


Color  standards  and  color 
Washington.    D.C.     Published 

Host  fungus  index  for  the 
>est  -I.  Hosts.  Wash.  State 
).    Sta.   Bull.    763.    121    p. 


BODY  SIZE,  ORCJAiN  SIZE,  AND  SEX  RAJ'IOS  IN  ADULT  AND 
YEARLINC;    BELDINC;    OROUND    SQUIRRELS 


Marl 


L.   M( 


WnlH-n    .1.     I'a 


AiJSTKACT.-^  A  fivcMMi-  sIikK  uT  I',i  ■](  I  i  1 1^'  ^^Miiinil  s(iiiii-rcls  was  roi 
Sierra  Nevada.  Bodv  \N.M-hl  and  l.ndv  len^illi  vane.!  seasunalK  dcpe. 
deposition  cycle,  age.  aiul  ,se.\.  Aduil  males  tended  to  he  l;ea\  ht  -jihI  h 
ticularly  in  the  last  half  of  the  active  season.  A  similar  pattern  was 
squirrels  were  often  distinguishable  from  adults  on  tlie  basis  ol  ImiU 
greater  in  adults  throughout  the  season,  and  mean  body  lengths  weie  giealc 
first  half  of  the  season.  Adults  also  had  larger  internal  organs  than  yearlings  , 
season.  In  liver  and  heart  this  difference  was  sustained.  Se.x  ratios  in  arluh'- 
1:1  hut  thei-e  was  c(insid(Mal)le  sp.itial  and  temporal  as\nnnetr\'  in  distiihulK 
tended    to    live    in   aieas    pci  iphnai    to   lush    nicailow  s  cm  i  upied  1)\    [em.ih's  .iiid  \( 


lu(  ted  at  liigh  altitude  in  the 
hug  upon  the  fat  depletion- 
igci-  liian  adidt  females.  i)ar- 
K'siMit  in  \earlings.  Yearling 
i/(\  MiMii  lM)(f\-  weights  wei-e 
gii'alcr    IN    aihilts   through   the 


llic  l)eginiung  of  the 
iid  in  yearlings  were 
(if   the   se.xes.    Males 


The  Belding  ground  sc|iiirrel  [Spermo- 
ph'dus  heldingi  bcldingi)  is  a  hiberiiator 
that  Hves  in  the  central  Sierra  Nevada 
Mountains  mainly  from  the  eastern  di- 
vide to  the  edge  of  the  Great  Basin  at 
altitudes  between  1,825  m  and  3,650  m 
(Storer  and  Usinger,  1970).  During  a 
five-year,  mark-release  study  of  5.  b. 
bcldingi  our  records  of  retrapj)ed  animals 
enabled  us  to  compile  data  on  individuals 
of  known  age  and  sex  for  prolonged 
periods.  In  the  course  of  this  study  it 
became  clear  that  three  functional 
groups,  based  upon  age.  existed  within  the 
pojDulation:  juveniles,  yearlings,  and 
adults.  Characteristics  of  juveniles  have 
been  previously  reported  (Morton,  Max- 
well, and  Wade,  1974).  Herein  we  rejiort 
on  seasonal  changes  in  body  size,  organs, 
and  on  sex  ratios  in  both  yearling  and 
adult  S.  b.  bcldingi. 

Methods 

The  study  \vas  conducted  from  1969 
through  1975  in  meadows  and  their  bor- 
dering areas  in  Lee  Vining  Canyon,  Mono 
County,  California.  Most  of  our  infor- 
mation stems  from  work  done  at  Big  Bend 
(elevation  ca  2.100  m)  and  es])ecially  at 
Tioga  Pass  (elevation  (a  3,000  m).  The 
active  seasons  are  similar  in  duration  for 
populations  at  both  areas  but  may  begin 
>^ix  weeks  or  more  a})art  due  to  chmatic 
differences  associatc^d  with  altitude  (Mor- 
ton, 1975).  All  data  reported  on  body 
weights,  bodv  lengths,  and  sex  ratios  of 
squirrels  of  known  age  are  from  Tioga 
Pass  animals.  Data  on  organ  weights 
were  combined  for  the  two  populations  at 


10-day  intervals  throughout  the  active 
season  in  order  lo  bolster  sample  size. 

Squirrels  were  ca])tured  alive  in  Toma- 
hawk wire-mesh  traps  baited  with  peanut 
butter.  Those  to  be  released  were  toe- 
clipjDod,  and  those  retained  for  specimens 
were  ethcrizecL  In  some  cases  specimens 
were  collected  with  a  .22  caliber  rifle. 
Body  weights  were  measured  to  the  near- 
est b.l  g  on  a  pan  balance.  Body  lengths 
were  taken  with  calipers  to  the  nearest 
0.1  ( ni.  Wet  weights  of  freshly  excised 
and  debrided  organs  were  measured  to  the 
nearest  0.01  g  on  an  anal^^ical  pan 
balance. 

Certain  small  meadows  or  sections  of 
large  meadows  were  used  only  for  mark- 
release  studies.  Sej)arate  data  logs  were 
maintained  for  each  toe-clipped  animal. 
In  our  terminology  juveniles  are  the 
young  of  the  year,  yearlings  were  born 
in  the  preceding  year,  and  adults  are  all 
animals  older  than  Aoarlings. 

Resi^lts 

The  first  S.  b.  bcldingi  to  emerge  each 
season  were  adult  males.  Within  a  few 
days,  however,  some  adult  females  and 
yearlings  could  be  found.  The  pace  of 
emergence  varied  somewhat  from  year  to 
year,  depending  upon  snow  cover.  Adults 
tended  to  enter  hibernation  earlier  than 
yearlings.  On  the  average  each  individual 
was  active  above  ground  for  about  three 
months  fMorton.  1975). 

BoDi  wEicHT. —  There  were  large  sea- 
sonal differences  in  bod^'  weight  due  pri- 
marily to  fat  depletion  or  deposition  and 
to  sex  and  age  differences   (Fig.  1). 


^Biology    Dcpartmcn 
-School    of    Mcdirine 


Odidcntnl     (.'ollogc,     I.os    iVngcIcs, 
iiivei'sity    ci[    Cnlifoinia.    San    Diego 


505 


306 


GREAT  BASIN  NATURALIST 


Vol.  35,  No.  3 


Fig.  1.  Seasonal  change  in  mean  body  weight 
of  Spermophilus  beldingi  beldingi  at  Tioga  Pass. 
Data  were  accumulated  over  five  seasons.  1969-73. 
Numerals  indicate  sample  size;  vertical  bars  de- 
note ±  2  S.E. 


Upon  emerging  in  mid-May  adult  fe- 
males at  Tioga  Pass  were  lighter  than 
adult  males,  but  during  pregnancy  they 
became  heavier  than  males.  In  late  July 
and  for  the  remainder  of  the  active  season 
mean  weights  of  adult  males  were  signifi- 
cantly greater  than  those  of  adult  females 
(P<0.05).  The  sexes  of  yearlings  were 
not  different  in  weight  until  late  July. 
Thereafter,  as  in  adidts,  males  were 
heavier.  Seasonal  trends  in  body  weight 
were  much  the  same  for  each  sex,  par- 
ticularly in  the  second  half  of  the  season. 
During  the  first  half  of  the  season  year- 
lings were  still  growing  rapidly.  As  a 
group,  yearlings  never  achieved  adult 
weight.  Adults  were  significantly  heavier 
(P<0.05)  than  their  yearling  counter- 
parts of  the  same  sex  at  every  class  in- 
terval throughout  the  season.  The  large 
weight  gain  observed  in  all  animals  dur- 
ing the  last  half  of  the  season  was  due  to 
fat  deposition  (Morton,  1975). 

Body  length. —  Growth  in  yearlings, 
as  indicated  by  body  length,  occurred 
throughout  the  season   (Fig.  2),  but  from 


23 


22 


21 


20- 


19 


18 


A   Yearling   Males 
O   Yearling  Females 


10        20 

30 

10        20 

30 

10        20 

30 

10        20 

30 

10        20 

May 

June 

July 

Aug 

Sept 

Fig.  2.  Seasonal  change  in  mean  ho{l\  length  of  Spermophilus  beldingi  brldingi  at  Tioga  Pass. 
Data  were  accumulated  over  five  seasons.  1969-73.  Numerals  indicate  sample  size;  vertical  bars 
denote  ±  2  S.E. 


Sept.  1975 


MOHTON.  PARMER:    SQUIRRELS 


307 


mid- July  on  yearling  females  as  a  group 
were  indistinguishable  from  adult  females 
in  body  length.  The  same  was  true  of 
males  except  that  the  smaller  males 
handled  in  August  and  September  invari- 
ably were  yearlings. 

Body  length  increased  in  adult  males 
as  the  season  progressed,  suggesting  that 
maximum  size  in  S.  b.  bcldingi  males 
ma\'  not  be  reached  until  be\'oiid  their 
second  year  of  life. 

Organ  weights. —  Liver  weights  in- 
creased rapidly  following  emergence  in 
all  animals  (Fig.  3),  but  the  increase  was 
more  rapid  in  females  than  in  males. 
Between  the  third  and  sixth  weeks  of 
activity  livers  of  females  were  larger  than 
those  of  males  (P<0.05).  Liver  hyper- 
trophy in  females  was  coincident  with 
lactation. 

By    the    twelfth    week    of   the    season 


yearlings  had  livers  of  adult  size.  Livers 
for  all  ages  and  sexes  were  smaller  at  the 
end  of  hibernation  than  at  the  beginning. 
Ap])arently  this  organ  atrophied  during 
hibernation. 

The  heart,  kidneys,  and  spleen,  were 
larger  in  adults  than  in  yearlings  during 
the  first  part  of  the  season  (Fig.  4).  This 
difference  was  particularly  noticeable  and 
])rolonged  in  heart  weight. 

Sex  Ratios. —  During  the  five  years  of 
this  study  a  total  of  341  yearhngs  (170 
males  and  171  females)  and  484  adults 
(238  males  and  246  females)  were 
handled.  The  sex  ratio  for  either  age 
group  did  not  differ  from  1 : 1  according  to 
a  chi-square  tost  (P>0.5()i. 

Discussion 

Body  size. —  Although  yearling  ground 
squirrels  often  represent  a  sid)stantial  ])or- 


A    Adult    Males 
•    Adult    Females 
A    Yearling  Males 
O    Yearling  Females 


60 

Days 

Fig.  3.  Seasonal  clianges  in  mean  liver  weight  of  Sprrnwphilus  bcldingi  beldingi  from  Big  Bend 
and  Tioga  Pass.  Numerals  indicate  sample  size.  Day  0  of  abscissa  refers  to  time  first  squirrels 
emerged  from  hibernation. 


308 


GREAT  BASIN  NATl^RALIST 


Vol.  35,  No.  3 


Days 

Fig.  4.  Seasonal  clianges  in  mean  weight  of 
spleen,  kidneys,  and  heart  in  Spcrmophilus  bcl- 
dingi  beldingi  from  Big  Bend  and  Tioga  Pass. 
Numerals  indicate  sample  size.  Symbols  as  in 
Figure  3.  Day  0  of  abscissa  refers  to  time  first 
squirrels  emerged  from  hibernation. 

tion  of  the  population  and  may  interact 
in  unique  ways  with  other  niembers 
(Michener  and  Michener.  1973),  there  is 
little  published  evidence  that  they  differ 
externally  from  older  animals.  Even  in 
such  large-bodied  species  as  S.  undulatus 
juveniles  reach  adult  size,  or  nearly  so. 
by  the  time  they  are  ready  for  hiberna- 
tion (Mayer  and  Roche.  1954).  A  priori 
this  might  be  predicted  since  ca]itive  \\\- 
A'eniles  of  hibernatory  Spcrmophilus.  par- 
ticularly those  from  high  latittide  or  high 
altitude,  tend  to  be  precocious  and  to  have 
e.xceptionallv  high  growth  rates  (Clark. 
1970;  Morton  and  Tung.  1971).  In  the 
case  of  S.  h.  /)('hli?is:i.  at  least,  by  the  time 
they  enter  hibernation  feral  juveniles 
have  foot  and  tail  lengths  indistinguish- 
able from  those  of  adults  (Morton  and 
Tung,  1971).  Nonetheless,  neither  maxi 
mum  body  weight  nor  itiaxinuun  boch 
length  is  achieved  in  .S'.  h.  hcldingi  until 
well  into  the  second  or  possibly  even  third 
year  of  life.  Furth(M-mor(\  we  have  found 
that  yearling  males  are  sexualh-  inunatuir 
and  do  not  reproduce  (Morton  and  (lal- 
lup.  unpubl.).  Yearling  females  do  vv- 
p/roduce.  Similar  age  (hfferences  in  re 
jiroductive  capacity  have  been  found  in 
5.  arrnatus  (Slade  and  Balph.  1971  i. 

The  differences  in  body  size  noted  in 
age  classes  of  S.  h.  beldingi  probably  are 


not  unique  among  ground  squirrels.  Such 
differences  are  likely  to  be  overlooked  un- 
less the  investigator  examines  large  num- 
bers of  animals  of  known  age  over  a  span 
of  several  consecutive  seasons. 

Sex  r.\tios. —  An  unbalanced  sex  ratio 
in  favor  of  females  has  been  reported  for 
many  ground  squirrel  populations.  This 
ratio  may  approach  or  exceed  3: 1  (Mc- 
Carlev.  1966;  Michener  and  Michener, 
1971;^Sheppard.  1972;  Turner,  1972).  In 
a  few  cases,  however,  the  ratio  foimd  did 
not  differ  significantly  from  1:1  (Clark. 
1970;  Murie,  1973;  present  study).  In  his 
study  of  S.  h.  orcgonus.  Turner  (1972) 
foimd  that  the  sex  ratio  was  1:1  in 
juveniles.  He  suggests  that  because  ju- 
venile males  tend  to  wander  and  explore 
more  than  females  they  incur  greater 
mortality,  resulting  in  an  unbalanced  sex 
ratio  in  older  animals.  In  S.  h.  hcldingi 
the  sex  ratio  in  juveniles  is  also  1:1;  males 
probably  wander  more  than  females  in 
that  they  have  larger  home  ranges  than 
females  and  are  more  likely  to  expand 
their  range  late  in  the  season  (Morton. 
Maxwell,  and  Wade,  1974).  Although 
this  behavior  would  seem  to  make  juve- 
nile males  more  susceptible  to  predation. 
we  have  no  evidence  that  it  does.  To  the 
( ontrarv.  males  and  females  occur  in 
equal  tnunbers  in  both  yearlings  and 
adidts.  The  sexes  are  not  distributed  ran- 
domly throtighout  the  habitat  occupied, 
however.  In  our  trap]iing  at  burrow  sys- 
tems located  in  lush  meadow  areas  re- 
served for  mark-release  studies,  adult  fe- 
males outnumbered  males  by  1.3:1  when 
all  data  were  summed.  The  ratio  tended 
to  fluctuate  seasonally,  however,  and  was 
sometimes  near  3:1,  particularly  at  mid- 
season.  We  often  captured  adult  males  at 
a  particular  burrow  system  oidy  a  few 
times  per  season,  during  the  first  days  or 
weeks  following  emergence  and  again  at 
th(^  \  erv  end  of  the  season.  This  suggests 
that  some  males  were  forced  from  the 
colony  \)\  conflicts  associated  \\dth  repro- 
(hiction  and  were  able  to  return  only  in 
time  to  (Mitcn-  their  trachtional  hibernacu- 
liun.  If  this  is  correct,  it  follows  that 
dis|)laced  males  should  be  found  in  areas 
jx'ripheral  to  main  colonies.  We  found 
this  to  be  the  (as(>.  When  males  were  de- 
sii-ed  for  spin  imens.  they  cotdd  usually 
be  found  b\-  collecting"  animals  scattered 
in    lodgepole    pine    stands    fringing    mea- 


Sept.  1975 


MORTON,  PARMER:    SQUIRRELS 


309 


flows,  and  in  rock)-  outcro])s  and  talus 
slopes  in  steep  areas  above  the  meadows 

A  tendency  for  males  to  live  in  areas 
peripheral  to  the  main  colonies  has  been 
observed  previously  in  S.  armatus  (Balph 
and  Stokes,  1963)  and  in  S.  richardsonii 
(Quanstrom,  1971).  Turner  (1972)  did 
not  believe,  however,  that  this  occurred 
in  S.  b.  oregonus.  The  spatial  distribution 
of  the  sexes  could  vary  enormously  de- 
pending upon  such  factors  as  habitat 
physiography  and  upon  intrinsic  charac- 
teristics of  the  population  itself. 

Undoubtedly  unbalanced  sex  ratios  ex- 
ist in  many  ground  squirrel  populations. 
Such  ratios  may  be  a  function  of  inter- 
populational  or  interspecific  differences  in 
breeding  systems  and  social  organization 
(Murie,  1973).  However,  an  investigator 
who  confines  his  work  to  locations  with 
maximum  animal  density  or  to  those  of 
easy  accessibility  could  obtain  an  inac- 
curate measurement  of  sex  ratio. 

Acknowledgments. —  We  wish  to 
thank  John  Gallup,  Roland  Leong,  Cath- 
erine Maxwell,  Allan  Tway,  and  Charles 
Wade  for  assistance  in  trapping  opera- 
tions. Southern  California  Edison  Co.  pro- 
vided housing  for  two  seasons.  Financial 
support  was  provided  by  Occidental  Col- 
lege and  by  National  Science  Foundation 
Grant  GB  29146X1. 

Literature  Cited 

Balph,  D.  R.,  and  A.  W.  Stokes.  1963.  On  the 
ethology  of  a  population  of  Uinta  ground 
squirrels.     Am.    Midi.    Nat.    69:106-126. 

Clark,  T.  W.  1970.  Richardson's  ground  squir- 
rels  {Spermophilus  richardsonii)  in  the  Lara- 


mie Basin,  Wyoming.  Great  Basin  Nat.  30: 
55-70. 

Mayer,  W.  T.,  and  E.  T.  Roche.  1954.  De- 
velopmental patterns  in  the  Barrow  ground 
squirrel,  Spermophilus  undulalus  barrowensis. 
Growtli   18:53-69. 

McCarley,  H.  1966.  Annual  cycle,  population 
dynamics  and  adaptive  behavior  of  Citellus 
tricedemlineatus.     J.    Mammal.   47:294-316. 

MiCHENER,     D.     R..     AND     G.     R.     MiCHENER.        1971. 

Sex  ratio  and  interyear  residence  in  a  popu- 
lation of  Spermophilus  richardsonii.  J. 
Mammal.  52:853. 

MiCHENER,     G.     R.,     AND     D.     R.     MiCHENER.        1973. 

Spatial  distribution  of  yearlings  in  a  Richard- 
son's ground  squirrel  population.  Ecology 
54:1138-1142. 

Morton,  M.  L.  1975.  Seasonal  cycles  of  body 
weights  and  lipids  in  Belding  ground  squir- 
rels.    Bull.    So.    Calif.   Acad.    Sci.   In   press. 

,  C.  S.  Maxwell,  and  C.  E.  Wade.     1974. 

Body  size,  body  composition,  and  behavior  of 
iuvenile  Belding  ground  squirrels.  Great 
Basin  Nat.  34:121-134. 

.    and    H.    L.    Tung.     1971.     Growth    and 

development  in  the  Belding  ground  squirrel 
{Spermophilus  beldingi  beldingi) .  J.  Mam- 
mal.  52:611-616. 

Murie,  J.  O.  1973.  Population  characteristics 
and  phenology  of  a  Franklin  ground  squirrel 
'{Spermophilus  franklinii)  colony  in  Alberta, 
Canada.    Am.   Midi.   Nat.   90:334-340. 

Quanstrom,  W.  R.  1971.  Behavior  of  Richard- 
son's ground  squirrel  Spermophilus  richard- 
sonii richardsonii.    Anim.   Behav.    19:646-652. 

Sheppard,  D.  R.  1972.  Reproduction  of  Rich- 
ardson's ground  squirrel  {Spermophilus  rich- 
ardsonii) in  southern  Saskatchewan.  Can.  J. 
Zool.,  50:1577-1581. 

Slade,  N.  a.,  and  D.  F.  Balph.  1974.  Popula- 
tion ecology  of  Uinta  ground  squirrels.  Ecol- 
ogy. 55:989-1003. 

Storer,  T.  I..  AND  Usinger,  R.  L.  1970.  Sierra 
Nevada  natural  history.  Univ.  Calif.  Press, 
Berkeley.    374  pp. 

Turner,  L.  W.  1972.  Autecology  of  the  Beld- 
ing ground  squirrel  in  Oregon.  Ph.D.  thesis, 
Univ.   of  Arizona,  Tucson.    149  pp. 


PHOTOPERIODIC  RESPONSES  OF  PHENOLOGICALLY 

ABERRANT  POPULATIONS  OF  PIERID  BUTTERFLIES 

(LEPIDOPTERA) 

Arthur  M.   Shapiro' 

Abstr.'^CT. —  T^vo  local  pierid  populations  in  western  Norlh  Ainerit.i  showing  regi(jnalh'  aix'rrant 
l)henologies  were  investigated  in  the  lahoratory.  Neither  a  parlialK  hivoltine  Piciis  iiapi  from  the 
Sierra  Nevada  foothills  in  El  Dorado  County.  California  i  sui  rnuiidi'd  hy  luiivoltine  populations),  nor 
a  vcrnal-univoltine  P.  occidenlalis  from  a  foothill  outlier  of  llie  (Colorado  Front  Range  (below  hi- 
voltine populations)  showed  unusual  resjwnses  to  controlled  developmental  regimes  in  the  labora- 
tory. Their  unusual  phenologies  are  hypothesized  to  be  the  produi  t  of  microclimate.  Failure  to  under- 
go genetic  adaptation  to  unusual  microclimates  is  discussed  witii  ])articulai-  reference  to  the  i)res- 
ence  or  absence  of  gene  flow  from  neaib\'  normal    populations. 


The  timing  of  life-history  phenomena 
in  an  insect  population  is  (leterminetl  by 
physiological  responses  to  en\ironniental 
stimnli.  These  proximate  controls  reflect 
a  genetic  basis  Ijelievecl  to  be  the  jiroduct 
of  natural  selection  for  seasonal  cycles 
appropriate  to  the  environment  of  the 
population.  In  the  western  LTnited  States 
topography  has  a  dramatic  impact  on  cli- 
mate, and  great  differences  may  occur 
over  short  ground  distances.  How  closely 
can  insect  populations  ada])t  to  their  im- 
mediate climates  on  a  microgeograj)hic 
scale?  Phenological  adaptation  is  merely 
one  case  of  the  more  general  jiroblem  of 
population  differentiation  (cf.  Ehrlich 
and  Raven,  1969;  Ehrlich  et  al.,  1975). 
In  most  organisms,  at  least  prior  to  the 
advent  of  electrophoretic  genetics,  popu- 
lation differentiation  was  assessed  on  the 
basis  of  visible  phenotypic  characters. 
Such  characters,  like  the  enzyme  systems 
studied  by  electrophoresis,  are  often  not 
translatable  into  specific  selection  pres- 
sures. In  markedly  seasonal  climates  the 
nature  of  selective  pressures  acting  on 
phenology  may  be  very  apparent.  Where 
local  deviations  from  the  broad  geograph- 
ic pattern  of  voltinism  are  observed  in  a 
species,  the  potential  exists  for  the  demon- 
stration of  microgeograj)hic  (or  ecotypic) 
differentiation.  This  is  the  fourth  paper 
in  a  series  exploring  the  evolution  of  sea- 
sonality in  the  butterfly  genus  Pieris  in 
western  North  America. 

In  various  multi\oltine  Pieridae  both 
phenotype  and  diapause  are  under  photo- 
periodic control.  The  two  sets  of  develop- 
mental options  (diapause /direct  develop- 
ment; vernal /esti\  a  1  j)henotype)  may  be 
physiologically  coupled   {Pieris  napi  Lin- 

'Departmcnt  of  Zoology.  Univcisily  n[  Cnlifoniin.  D.ivi';.  Ca 


naeus  complex)  or  not  (P.  protodice  Bois- 
duval  &  LeConte,  P.  occidciitdUs  Reakirt). 
Recent  st tidies  have  shown  that  univol- 
tinism  in  both  grouj)s  is  derivative  from 
multivoltinism,  accompanying  invasion 
of  a  short-summer  climate  (P.  occidcn- 
ta/is,  Sha})iro,  1975a)  or  persistence  in  a 
progressively  drier  one  (P.  napi,  Shapiro, 
1975b).  Such  patterns  are  defined  over 
broad  geographic  areas.  California 
P.  napi,  for  example,  is  differentiated  into 
a  commonly  biv'oltine,  heavily  pigmented 
subspecies  in  the  coastal  summer-fog  belt 
and  a  univoltine,  more  lightly  marked 
subspecies  in  the  interior,  where  summers 
are  clear  and  hot.  The  transition  between 
the  subspecies  apjjears  to  be  in  the  form 
of  a  stee])  cline  through  the  central 
Coast  Ranges   (Shapiro,  in  preparation). 

Recently  Lees  and  Archer  (1974)  have 
reported  the  existence  of  phenological  dif- 
ferences among  napi  populations  on  a 
mvich  finer  scale.  They  have  found  ap- 
parently relict  univoltine  populations  in 
suitable  (bog-heath)  habitats  completely 
surrounded  by  multivoltine  ones  in  the 
British  Isles.  Their  preliminary  interpre- 
tation of  this  situation  is  that  it  provides 
(n  ideiice  for  midtiple  invasions  of  Britain 
h\  napi  stocks  having  different  pheno- 
logical characteristics  and  source  regions. 
In  th(>  course  of  recent  work  on  pierid 
|)hen()logy  and  e\olution.  the  existence 
of  regionally  aberrant  populations  has 
b(>en  i)r()ught  to  my  attention  in  both  the 
napi  and  protodicc-occidcntalis  groups.  In 
both  ( ases  the  populations  appear  to  be 
uniciiie,  rather  than  forming  a  repeating 
pattern  as  in  British  P.  napi.  They  would 
therefore  seem  to  be  good  candidates  for 


310 


Sept.  1975 


311 


local   genetic    tliflcreiitiatioii    under   al\[j 
ical  microclimates 


he  Sierra  Foothills 


ricns  iiapi  ii 

Pieris  napi  from  interior  (-alitornia  are. 
as  noted  above,  luiivoltine  and  moiio- 
phenic  in  nature.  Under  laboratory  (au- 
ditions they  can  lie  reared  without  dia- 
pause; then  they  produce  the  eslixai 
phenoty|)(»  ''castoria''  Airtually  unknown 
in  the  wild  in  the  interior  (Sha])iro. 
1975b).  In  June  1974  Mr.  William  Pat- 
terson of  Sacramento,  California,  took 
se\-eral  wild  'Vy/.s/o/vV/""  of  hoth  sexes  in 
the  canyon  of  the  American  Ri\  er  below 
Auburn  in  the  Sierra  Nevada  foothills 
(El  Dorado  County,  650  feet).  The  oc- 
currence of  a  second  brood  there  was  con 
firmed  in  1975.  P.  napi  is  connnon  in  the 
canyon,  producing  its  usual  \ernal  pheiid 
type  in  March,  llie  second  brood,  whi(  li 
is  much  scarcer,  unlike  the  first  is  ex- 
tremely localized  ^vithin  llu^  camon  -al 
present  being  known  from  onh'  two 
densely  shaded  ravines  where  the  intro- 
duced cruciferous  weed  watercress   (Nas- 


lurliuni   offn  iiialc   W.   Br. 
turliu!ih(i(jii(ih(nni      Schii 
gr'ows     in     [xTincUient     stn 


Rorippa  na.s- 
:.  &  Thell.) 
ims  (Fig.  1). 
hutterflies  are 
bred       Sierran 


Most  .,r  ih,>  uiid  ,1,1, 

idonlicil        to        lal)orat( 
"cdstnrid'     I  h'ig.  1  i . 

On  J<)  Man  h  l<)7)  niii(>  in.de  and  three 
lemale  lirst-hrood,  vernal  phenotyjie  napi 
were  colIectcMl  in  one  of  these  ravines. 
These  included  two  coj)ulating  j)airs  in 
which  the  females  were  soft-winged,  in- 
dicating that  they  had  developed  in  the 
ravine^  itself.  The  eggs  from  these  females 
were  used  in  photoperiod  experiments 
( Table  1  ) .  ( Rearing  methods  are  de- 
scribed m  Shaj)iro.  'l975a  and  1975b.) 
Hie  results  are  entirely  typical  for  Sier- 
ran stock  and  do  not  suggest  that  x-Xmer- 
i(an  Ri^er  material  has  a  greater  pro- 
pel isit\'  to  flevelop  directly  than  do  stocks 
tVoju  purely  uniAoltiiie  localities,  at  least 
under  our  laboratory  regimes.  However, 
ihis  is  not  particularly  surprising.  The 
second  brooci  of  napi  in  the  American 
Ri^■er  gorge  is  nukh  rarer  than  the  first, 
indicating  that  is  is  onl}'  partial;  its  num- 
bers   also    fluctuate    from    year    to    year. 


B 


B 


(^5SS£S^IJ4;>rSjgx%,c-;.v|f^Vr^ 


Fig.    1.     Locations  of  ravines    ("B")    where   !)ivoItiiie  Pieris  tuipi 
univoltine  napi  are  generally  distriluited  at  low   (Icnsity.     l'S(;S    7. 


Ill  the  American  River  gorge; 
life    "Aulnun"'    quadrangle. 


312 


GREAT  BASIN  NATURALIST 


Vol.  35,  No.  3 


Table  1.  Incidence  of  diapause  ( D)  and  non- 
diapause  (ND)  pupae  in  bivoltine  (American 
River,  650')  and  univoltine  (Placerville,  1800') 
Pieris  nopi  from  El  Dorado  Co.,  Calironia,  reared 
on   watercress  at  27  C  under  two  photoperiods. 

Photophase:     Continuous         15  hr 
Stock:  Pupae:        D  ND         D     ND 


American   River 
Placerville 


15     29         16 
10     23  18 


(In  197:5  three  trips  by  Patterson  and 
Shapiro  in  season  turned  up  only  tNvo 
males  and  one  female.  A  later  search  of 


the  host  plant  at  the  optimum  time  failed 
to  turn  up  ;ni\  napi  immatures,  although 
ten  Pieris  rapac  lar^'ae  were  found.) 
These  circumstances  suggest  that  the 
production  of  a  second  brood  here  is  ac- 
cidental, resulting  from  the  peculiarly 
cool  and  moist  conditions  within  the  ra- 
\'ines.  There  is  no  evidence  that  the  bi- 
Aoltine  sites  are  in  any  sense  isolated 
from  adjacent  univoltine  ones,  nor  is  it 
clear  that  there  is  successful  re])roduction 
by  the  second  brood  in  all  years  nor  even 
that  there  is  genetic  continuity  from  year 
to  year  in  the  ravines;  perhaps  a  few  pu- 


t 


^P'- 


Fig.  2.  Wild  second-brood  Pieris  napi  from  the  American  River  gorge,  collected  by  W.  Patterson 
in  June  1974  fmales  at  top;  dorsal  (left)  and  ventral  (right)  surfaces).  The  heavily  marked  female 
is    atypical    for   an    inland    population. 


Fig.    3.     Phenotypes    of    representative    lab-reared  nondiapause  Pirris  napi  from  the  Ameiican  Riv( 
stock;  27  C.  continuous  light;  dorsal   (left)    and  ventral    (right). 


Sept.  1975 


SHAPIRO:   BUTTERFLIES 


313 


pae  will  develop  directly  there  whenever 
any  female  napi  happens  to  colonize  them. 
Experienced  California  collectors  (R.  L. 
Langston,  B.  Walsh)  agree  that  even 
near  the  coast  some  localities  produce 
second-brood  napi  every  year  and  others 
only  rarely  or  sporadically.  Exj)eriments 
have  shown  both  developmental  and 
phenotypic  differences  between  coastal 
and  inland  stocks  but  not  among  the 
coastal  stocks  themselves. 

Watercress  is  known  to  be  host  of  P. 
napi  in  various  Sierran  sites  up  to  about 
•5,000  feet  (Shapiro,  1975c).  The  only 
other  record  of  a  Sierran  ^'castoria" 
known  to  me  is  a  fresh  male  taken  flying 


Table  2.  Incidence  of  diapause  (D)  and  non- 
diapause  (ND)  pupae  in  two  split  broods  of  a 
Barbarea  verna-ieeAin^  univoltine  Pieris  napi 
(Gates  Canyon,  Inner  Coast  Ranges,  Solano  Co., 
750')  reared  at  27  C  on  continuous  light.  None 
of  the  differences  was  significant. 


Brood       Host 


Pupae: 


Brassica    kabet-^  7 

Nasturtium  officinale^'  5 

Brassica  kaber''^  6 

Lepidiuni  latifolium^'-''  5 


ND 

develop- 
mental 
time 
in  days 

21 

25.2 

11 

25.3 

14 

25.6 

16 

24.6 

Notes:  [a]  Tops,  (b)  Elongating  rosettes.  Ui  In  subsequent 
experiments  mature  tops  did  not  supirart  development. 
Butterflies   in   this   brood  were   stunted. 


among  first-brood  vernal  napi  at  Lang 
Crossing,  Nevada  County,  4,500  feet,  9 
.June  1975.  At  this  locality  napi  feeds  on 
both  watercress  and  native  vernal  cru- 
cifers.  There  are  several  possible  explan- 
ations of  this  odd  individual,  but  to  test 
the  hypothesis  that  watercress  feeding 
itself  inhibits  dia})ause,  split-brood  experi- 
ments were  conducted  in  1975  using  an 
Inner  Coast  Range  stock  (Gates  Canyon) 
with  no  previous  exposure  to  the  plant. 
No  evidence  of  a  dietary  influence  on  the 
incidence  of  diapause  was  found  in  this 
univoltine  strain   (Table  2). 

Pieris  occidcntalis  in  Colorado 

Haystack  Mountain  (5,589  feet)  is  an 
isolated  hill  eight  miles  northeast  of  Boul- 
der, Boulder  County,  Colorado  (Fig.  4), 
where  Dr.  Ray  E.  Stanford  of  Denver 
has  for  several  years  taken  small,  dark 
vernal  ''calycc'  phenotypes  of  Pieris  oc- 
cidentalis  indistinguishable  from  the 
single  brood  above  treeline  in  midsmnmer 
(Fig.  5).  He  has  no  summer  records  of 
P.  occidentalis  from  Haystack  Mountain 
but  finds  its  lowland  sibling  P.  protodice 
there  in  summer  instead.  Because  P.  pro- 
todice winters  only  very  locally  but  colo- 
nizes widely  in  summer,  this  is  not  sur- 
prising; it  does  however,  raise  the  pos- 
sibility that  P.  occidentalis  has  undergone 
a    phenological    shift    to    univoltmism    in 


7     r^-    r> 


t 


Fig.    4.     Location   of   Haystack    Mountain,    Boulder    County,    Colorado.    USGS    7.5-minute    "Boulder' 
and   "Niwot"   quadrangles. 


314 


GREAT  BASIN  NATURALIST 


Vol.  35,  No.  3 


Fig.    5.     Wild    Pieris    occidentalism    vernal    phenotype  C'calycc'').  from  Haystack  Mountain,  collected 
by  R.  E.  Stanford.  Males  at  left;  dors;d   (left)  and  ventral     (right)     surfaces. 


response  to  competition  from  P.  protodice 
(although  no  such  phenomenon  is  known 
at  other  localities  where  the  two  are  sym- 
patric).  Haystack  Mountain  is  probably 
the  lowest  elevational  record  for  P.  oc- 
cidentalis  in  Colorado.  In  the  Rockies 
proper  it  is  bi\oltine  at  middle  elevations 
(perhaps  locally  trivoltine)  and  univol- 
tine  in  the  Alpine  zone  (cf.  Brown,  Eff, 
and  Rotger,  1957)  and  has  two  seasonal 
phenotypes    (Shapiro,    1975d). 

A  laboratory  stock  was  established 
from  ova  laid  by  five  females  collected  by 
Stanford  on  6  April  1975.  Under  labora- 
tory conditions  their  developmental  and 
phenotypic  responses  (Table  3  and  Fig. 
6)  were  identical  to  both  Sierran  multi- 
voltine  and  Colorado  Alpine  stocks  (Sha- 
piro 1974,  1975d).  Once  again  we  have 
no  experimental  evidence  for  the  evo- 
lution of  a  phenological  ecotype  and 
are  therefore  forced  to  look  for  micro- 
climatic explanations.  Since  Haystack 
Mountain  is  effectively  in  the  Great 
Plains  climatic  regime,  which  is  hotter 
and  drier  than  the  usual  regime  of  P. 
occidentalis,  it  may  not  be  surprising 
that  conditions  there  would  be  associated 
wdth  summer  dormancy.  This  (juestion 
can  be  settled  only  b}'  laboratory  duplica- 
tion of  Haystack  Mountain  conditions  or 


by  testing  the  developmental  responses 
of  multivoltine  P.  occidentalis  stock  from 
elsewhere  reared  at  Haystack  Mountain. 
We  hope  to  carry  out  such  experiments 
within  the  next  couple  of  seasons. 

Phenological  differences  are  known  be- 
tween plains  and  lower  montane  popu- 
lations of  a  inunber  of  Colorado  butter- 
flies (.I.A.  Scott,  R.E.  Stanford,  pers. 
comm.),  but  they  may  go  in  a  direction 
o])posite  to  those  observed  in  Pieris  oc- 
cidentalis. Two  species  (Colias  olexandra 
Edwards,  Pieridae;  Plebeius  icarioides 
complex,  Lycaenidae,  both  Legume  feed- 
ers) are  bivoltine  on  the  plains  and  uni- 
voltine  in  the  mountains.  The  basis  for 
these  differences   is   uninvestigated. 

With  no  evidence  for  genetic  differen- 
tiation of  Haystack  Mountain  occidentalis. 
the    attractive    hypothesis    of    competitive 

Tahi.e  1  Incidence  of  diapause  iD)  and  non- 
diapause  (ND)  pupae  in  veiiial-univoltine  (Hay 
stack  Mountain,  Colordao,  o.oSO')  and  bivoltine 
(Donnor  Pass,  Colorado,  7,000')  Pieris  occident- 
alis reared  on  Brassica  kaher  at  27  C. 


Stock: 


Photophase:        Continuous  15   hr 

Pupae:        D  ND         D     ND 


Haystack    Mountain.    1975 
Donner  Pass.  1973 


0     22 
0     16 


Sept.  1975 


SHAPIRO:   BUTTERFLIES 


315 


^ '.  -^_f ^-  -1 


Fig.  6.  Phenotypes  of  representatives  lal)-reared  nondiapause  Picris  occidcntalis  from  the  Haystack 
Mountain  stock;  27  C.  continuous  light.  Estival  phenotvi)es  characteristic  of  nnitlivoltine.  Males  at 
left;    dorsal    (left)    and   venti-al    (right)    surfaces. 


seasonal  displacement  with  protodicc 
must  be  set  aside.  The  host  plants  of  both 
species  on  Haystack  Mountain  are  un- 
identified. Both  prefer  species  of  pepper- 
grass,  Lepidium,  throughout  their  ranges. 
On  the  plains  most  crucifers  are  vernal 
species,   as   in   lowland   California. 

DiSCEISSION 

Many  instances  are  on  record  of  eco- 
typic  differentiation  on  a  microgeogjraphic 
scale,  particularly  in  plants,  which  have 
more  versatihty  in  developing  isolating 
mechanisms  than  do  animals  (Jain  and 
Bradshaw,  1966). 

As  noted  above,  the  lack  of  a  genetic 
basis  for  biovoltinism  in  Sierran  Picris 
napi  is  not  very  surprising,  granted  the 
extremely  restricted  habitat  and  the  ex- 
tensive distribution  of  univoltine  butter- 
flies, with  ample  opportunity  for  gene 
flow.  The  failure  of  the  Haystack  Moun- 
tain P.  occidcntalis  to  differentiate  is 
more  intriguing.  It  is,  of  course,  possible 
that  it  has  differentiated  and  that  the  lab 
rearing  regimes  were  too  crude  or  inap- 
propriately selected  to  show  it.  It  is  cer- 
tain that  experiments   to   date,   involving 


simple  manipulation  of  constant  rearing 
temperatures  and  unchanging  day- 
lengths,  have  gi^en  an  oversimplified 
picture  of  the  developmental  versatility 
of  pierids  in  the  field.  If  microclimate 
determines  aberrant  voltinism  in  these 
stocks,  it  is  very  likely  that  humidity, 
for  exam])le,  may  interact  with  photo- 
period  and  temperature  in  controlling  de- 
\'elopment  in  natural  populations.  The 
same  genetic  information  may  allow 
Picris  occidcntalis  to  respond  ap])ropriate- 
ly  to  regimes  as  diverse  as  those  at  Hay- 
stack Mountain  (5,589  feet)  and  Love- 
land  Pass  (12,400  feet). 

Given  such  jilasticity,  we  may  wonder 
whether  the  Baldwin  effect  (Simpson, 
1953)  might  not  come  into  play  in  popu- 
lations in  extreme  environments.  Briefly, 
the  Baldwin  effect  postulates  the  buildup 
by  selection  of  a  genetically  obligate  basis 
for  the  ada})tations  produced  via  develo])- 
menlai  plasticity.  In  an  atypical  hut  pre- 
dictable climate  like  Haystack  Mountain, 
might  not  the  developmental  flexibility 
chara(  t(>ristic  of  montane  poj)ulations  be 
lost?  (^Alaskan  Picris  occidcntalis  nclsoni 
seem  to  be  evohing  in  this  direction; 
Shai)ir(>,   1975a.)    One  important  counter- 


316 


GREAT  BASIN  NATURALIST 


Vol.  35,  No  3. 


vailing  force  would  be  gene  flow,  which 
is  almost  certainly  operating  on  high- 
elevation  univoltine  occidcnUdis  in  Colo- 
rado (Shapiro,  1975d).  Haystack  Moun- 
tain is  about  30  air  miles  from  timberline 
and  much  closer  than  that  to  the  montane 
zone,  but  how  isolated  it  actually  is  is 
quite  unknown.  Nor  is  there  any  infor- 
mation bearing  on  how  long  occidentalis 
has  been  there — whether  it  is  a  Pleisto- 
cene relict  or  a  recent  colonization.  There 
are  much  more  isolated,  certainly  relict 
occidentalis  populations  in  other  localities 
east  of  the  Front  Range — the  Black  Hills 
of  South  Dakota  and  perhaps  the  Pine 
Ridge  of  northwestern  Nebraska — which 
deserve    study    in    this    regard. 

Acknowledgments 

Of  the  collectors  who  provided  vital  in- 
formation and  who  have  been  credited 
in  the  text,  special  thanks  are  due  Mr. 
William  Patterson  and  Dr.  Ray  Stanford. 
without  whose  help  these  experiments 
would  have  been  impossible.  Mr.  Mark 
Kauzer  assisted  in  field  w^ork  and  Mrs. 
Adrienne  R.  Shapiro  in  rearing.  This  re- 
search is  part  of  a  larger  study  of  colo- 
nizing ability  and  the  evolution  of  season- 
ality in  Pieris  funded  by  the  Committee 
on  Research,  UCD,  under  grant  D-804. 


LiTER.-^TURE  Cited 

Brown.  F.  M.,  D.  Eff,  .\nd  B.  Rotger.  1957. 
Colorado  butterflies.  Denver  Museum  of 
Natural   History.   Denver.     368   pp. 

Ehrlich.  p.  R..  and  p.  H.  Rwen.  1969.  Dif- 
ferentiation of  populations.  Science  165: 
1228-1233. 

EiiRLicH.  P.  R..  R.  R.  White.  M.  C.  Singer. 
S.  W.  McKechnie.  and  L.  E.  Gilbert. 
1975.  Checkerspot  butterflies:  a  historical 
perspective.   Science   188:221-228. 

Jain.  S.  K..  and  A.  D.  Bradshaw.  1966.  Evo- 
lutionary divergence  among  adjacent  pop- 
ulations. I.  The  evidence  and  its  theoretical 
analysis.    Heredity   21:407-441. 

Lees,  E..  and  D.  M.  Archer.  1974.  Ecology 
of  Pieris  napi  (L.)  (Lepidoptera.  Pieridae) 
m  Britain.   Ent.   Gazette  25:231-237. 

Shapiro.  A.  M.  1974.  Photoperiodic  control  of 
seasonal  polyphenism  in  Pieris  occidentalis 
Reakirt  (Lepidoptera:  Pieridae).  Wasmann 
J.    Biol.    31:291-299. 

.  1975a.  Photoperiodic  control  of  de- 
velopment and  phenotype  in  a  subarctic 
population  of  Pieris  occidenlalis  (Lepidoptera: 
Pieridae).  Canad.  Ent. 

.      1975b.     Developmental    and    phenotypic 

responses  to  photoperiod  in  uni-  and  bivoltine 
Pieris  napi  (Lepidoptera:  Pieridae)  in  Cali- 
fornia. Trans.  Rov.  Ent.  Soc.  London  127: 
65-71. 

.      1975c.     The     role     of     watercress,     Nas- 

lurtiuni  officinale,  as  a  host  of  native  and 
introduced  pierid  butterflies  in  California. 
J.   Res.  Lepid. 

.      1975d.     Ecotvpic    variation    in    montane 

butterflies.   Wasmann   J.    Biol.    32:267-280. 

Simpson.  G.  G.  1953.  The  Baldwin  Effect. 
Evolution    7:110-117. 


ADDITIONAL  RECORDS  OF  Ri:i?TILl^:S  I'ROM  .lAIJSCO.  MEXICO 

Philip   A.    Mcdira'.    Rii,l,,ir   (i.    Ain.ll-.    ,ni.l    l.nurs    I',.    I)iv,,n^ 


Abstract. —  Notewortliy  ri-cords  ol  tr|ililc^  iVoi 
first  state  recoi'ds  of  Colennyx  clrga/is  /u-nioi  tills,  llic 
notes  on  the  distribution  ;ind  roprodiK  tinti  of  F.umrcc 


l.diMii.  Affxito.  are  presented.  The 
KM  111  d  for  Gcophis  tarasra<\  and 
if/duhl/us   are   discussed. 


A  small  collection  of  amphibians  and 
reptiles  from  Jalisco,  Mexico.  \  icldc^l 
several  noteworthy  specimens. 

Coleonyx  elegans  nemoralis  I\laiil)(M'. 
One  male  (88  nnn  snout  to  Aciit  lo]i<>th. 
SVj  from  28.1  km  SW  Autlan,  .lali'sco. 
(Texas  Cooperative  Wildlife  Collection, 
i^exas  A  &  M  University,  TCWC  48035  ). 
collected  on  19  May  1974.  elevation  518 
m.  There  are  three  known  sjiecimens 
from  Jalisco.  The  first,  a  male  (Los  An- 
geles Coimty  Musemn,  LACM  37568 )  83 
mm  SV,  from  74.2  km  SW  Antlan.  Jalis- 
co, was  collected  by  J.  R.  Dixon  and  R. 
Heyer  on  23  July  19fv.  This  specimen 
has  10  preanal  pores;  7-7  supralabials;  h-7 
infralabials;  9  scales  nostril  to  nostril;  0 
gulars  contacting  mental;  4/4  gulars  con- 
tacting first  infralabials;  21  rows  of  tuber- 
cles across  at  midbody,  and  16  fourth  toe 
lamellae.  Our  specimen  (TCWC  48035 ) 
has  11  preanal  })ores;  8-8  supralabials;  7-8 
infralabials;  9  scales  nostril  to  nostril;  5 
gulars  contacting  mental;  1/5  gulars  con- 
tacting first  infralabials;  20  rows  of  tuber 
cles  across  at  midbody,  and  1  7  fourth  toe 
lamellae.  The  third  specimen  ( Brigham 
Young  University,  BYLT  41299).  a  male 
85  mm  SV,  from  14.5  km  N  Barra  de 
Navidad,  Jalisco,  was  collected  by  J.  Ott- 
ley  on  30  October  1974.  This  spetimen 
has  10  preanal  pores;  6-7  supralabiaL;  7 -S 
infralabials;  9  scales  nostril  to  nostril!;  (> 
gulars  contacting  mental;  2/2  gulars  con- 
tacting first  infralabials;  21  rows  of  tu- 
bercles across  at  midbody,  and  16  fourth 
toe  lamellae.  All  specimens  are  within  the 
range  of  variation  described  by  Klauber 
(1945).  These  specimens  extend  X\\v 
known  range  of  this  species  inland  some 
107  km  N  from  the  coastal  area  of  Cf)- 
lima,  the  heretofore  northenmiost  portion 
of  its  known  range  (Klauber.  1945;  Kluge. 
1975). 

Eunieces  brevirostris  iudiibitus  Taylor. 
A  series  of  20  specimens,  including  9  jm c- 

1U.C.I..A.,  P.  0.  Box  495,  Mercury,  Xcvnda  SOUJ  !. 
^Faculty  of  Natural  Sciences  &  Mnthematics,  Slofkl-u  Sl.il.' 
^Dcparimcnt  of  Wildlife  Sciences,  Texas  A  Sc  M  T"ui\ci^ii  v 


niles  (2l-)0  mm  SV.  x  --  26.7  mm)  and 
n  adidts  (52-75  mm  SV,  x  -  61.2  mm), 
collected  25  km  '>>¥.  Autlan,  Jalisco 
(J'CWC  ISO  50- 18055  I.  on  17  May  1974. 
J'his  siic  is  appro\iniat(4\  50  km  W  of  the 
ne.ucsl  i-eporled  hxalitv  and  about  100 
km  SSE  of  the  northernmost  locality  for 
this  subspecies,  both  in  Jalisco,  thus  par- 
tially filling  the  hiatus  in  the  distribution 
described  1)\  Dixon  (1969).  Our  speci- 
mens (>xhil)it  <i  signifi(  antlv  lower  mimber 
of  fourth  loe  lamellae  (  11-11.  x  -  11.8 
-  0.25)  and  Mipei-ciliaries  ( 5-()  to  7-7^ 
X  6.27  '  0.08  I.  but  olherwise  they  fall 
well  \\ithin  the  range  of  variation  for 
indul'ilus  given  In  Dixon  (1969).  This 
spec  ies  is  ()\  ()\  ivipatous  according  to  Tan- 
ner (1058).  \-\h()  reported  a  female  that 
(ontaiiu'd  two  fulK  devclojied  embryos. 
\A'e  secured  on(>  large  female  (75  mm  SV. 
weight  ().()7  g)  that  bore  six  live  young 
between  llie  time  of  capture  and  the  next 
morning.  Iheir  r.uige  in  length  was  24- 
28  mm  S\'  (X  2<).  >  mm)  and  in  weight 
from  0.  )7-0.  I()  g  (X  0.11  gi.  All  Kumeces 
\^('re  Found  in  pine-oak  woodland  imder 
rocks  within  20  m  of  a  |)ermanent  stream, 
elevation  1.1-3  3  m.  It  is  of  interest  to  note 
(hat  in  this  area  there  had  been  no  ap])re- 
ciable  rainfall  since  October  1973,  and 
most  decidous  vegetation  was  devoid  of 
lea\es.  Despite  |he  apparent  lack  of  mois- 
lure.  /''u/ncccs  h.  'mdiihitus  was  capable  of 
reprochu  ing.  perliaps  o\\ing  to  its  ovo^'i- 
\iparous  nature. 

(h'ophis  ididsciw  flartweg.  One  female 
from  l-\  km  S1-:  Autlan.  Jalisco  (TCWC 
l'79l-8),  collected  on  17  May  1974,  eleva- 
tion 1,1'33  m.  J1hs  apparently  is  but  the 
Fifth  known  sj)ecimen  oF  this  species  and 
the  stMond  from  th(>  slate  of  Jalisco. 
Downs  (1967)  rej)ortecl  on  three  speci- 
mens From  tli(>  ty])e  locality  at  Uruapan, 
\lichoa(an;  Dixon  (19()8i  reported  one 
spcH  imen  From  Ne\,ido  de  Colima  some 
FS    km    'o    [he    !■'..    Our    s[)ecimcMi    has    15 

nll..,^,..  P.iiiiM,,.,.  Xcw  Jci>ey  nS2-IU^ 


317 


318 


gri:at  basin  nai 


Vol.  35,  No.  3 


scale  rows;  146  ventrals;  41  caiidnls;  0  +  1 
temporals;  6  supralabials;  6  infralabials; 
1  loreal;  1  postocular;  no  preocular,  and 
10/10  maxillary  teeth.  The  first  pair  of 
chin  shields  is  twice  the  length  of  the 
second  }:)air;  sujiraocular  distinct,  larger 
than  loreal;  internasals  (H^ided  and  dis- 
tinct. The  dorsum  and  tail  are  dark  gray 
with  blackish  crossbands  (40  on  the  body, 
13  on  tail)  which  do  not  extend  across 
the  venter;  anterior  crossbands  2-3  scale 
rows  wide  and  bordered  on  the  anterior 
and  posterior  by  white-edged  scales,  pos- 
terior crossbands  1  scale  row  wide  and 
highly  irregular,  frequently  broken  later- 
ally and  dorsally;  head  distinct  from  neck; 
color  of  rostral  and  ])renasals  similar  to 
adjacent  scales;  eye  twice  into  snout 
length;  ^'enter  greenish  white  in  life, 
heavily  spotted  with  black.  The  total 
length  is  236  mm,  and  the  tail  is  45  nnn. 
This  specimen  has  fewer  Aentrals  than 
reported  by  Downs  (1967)  and  Dixon 
(1968).  Habitat  was  as  described  for 
Eumeces  brevirostris  indul>itus  above. 

The  authors   are   indebted   to   Dr.    An- 
tonio  Landazuri   Ortiz,   Director   General 


de  Fauna  Silvestre  of  Mexico,  for  pro- 
Aiding  the  necessary  collecting  ])ermits. 
We  also  tlijuik  Drs.  Wilmer  W.  Tanner 
,iM(l  Idlni  W.  Wright  for  loan  of  speci- 
mens in  llieir  care,  and  are  grateful  to 
Plnli[)  I.  M(Mlica  for  his  untiring  assis- 
tance   in    the    field. 

l.ri  i;i!  MLKI'.    ClTKO 

Dixo'V,  I.  I'.  I"I()S.  Notes  on  tlu-  snake  genus 
Gcf)f)/iis.  Iiom  Nevado  de  Colima  Jalisco, 
Me\i,,K    Sniiiliwest.   Nat..  13(,+):452-454. 

I '((]').  I'axononiic  review  of  the  Mex- 
ican skinks  nl  the  Eumeces  brevirostris  group. 
I.os  Auiivlv^  Coimtv  Mus.  Contrih.  Sci.  No. 
Ifi8: !-')(). 

Downs,  I''.  I..  l')f)7.  Intrageiieii(  I'ehitionships 
;iiiioiig-  (ohihiid  snakes  of  the  s<"iius  Geophis 
Wauh'i-.   Misc.    I'uhh   Mus.  /ool.    I'liiv.   Mich. 

Ki.AUHi.H.  I..  M.      I'*!-").     'I'lie  fie,  kos  of  tlie  genus 

Colc'i/nv  uith  (les(  liptions  of  new  suljspecies. 

'I^-.iiis.     San     Dh'Ko    So,.     Nat.     Hist.     lOi'll): 

I  >')-2Mi. 
Kir, a;.    ;\.    (..      l')7').      l^ln  l,ifj;eiieti,     lelationsiiips 

,inil    ,'\i)hihiiiiai  \     triTifls    in    tlie    eublepliarine 

ll/ai-,1  f;,Mins  Cnlrnnrx.  Copeia   1975   (l):24-35. 
■|'\Ni\i;H.    \\'.    \\'.      IM'xS.      Two    new    skinl^s    from 

Dinanf;,).    Me.xi,,).    Great    Basin    Nat.     18(2): 

")74)2. 


INVASION  OK  BIG  SAC^EBRUSH  [ARIEMISIA  rRlDENlATA) 

BY  WHITE  FIR  {^ABIES  CONCOLOR)  ON  THE  SOUTHEASTERN 

SLOPES  OF  THE  WARNER  MOUNTAINS,  CALIFORNIA 


rii 


IS  R.  Vale 


Abstract.-  -  \Vliito  I'ir  [  Ahics  concolor)  appears  to  he  i 
sagebrush  (Artemisia  tridcniata)  on  the  southeastern  slopes 
northeastern  California.  The  time  of  initial  tree  establishment 
increment  borings.  Possible  causes  of  the  invasion  involving  s 
tory.  and  grazing  use  of  the  area  during  years  of  the  (>stahlisii 
('omestic  livestock,  particularly  sheep,  appears  respousil)l<'  f 
tation    and    allowing    tree    invasion. 


i\a(hng  \('g('tation  (Joniiiiat(>(l  b\-  big 
if  the  ^^'al■ner  Mountains  of  e.xtrenie 
within  the  shrubs  was  detennined  by 
ic(ili(  (limalic  conditions,  fire  his- 
iiciil  were  cxidoied.  Heav\-  grazing  by 
V     .iltcrnig     the     sagi-brush-grass     vege- 


The  Wariicf  Moiiiitaiiis  of  cxtrGme 
northeastern  California  rise  to  3,000  ni 
above  sea  level  atid  snj)port  a  forest 
dominated  by  white  fir  {Ahics  concolor) 
and  ])onderosa  pine  {Pinus  ponderosa) 
(Fig.  1).  The  adjacent  valleys  lie  at 
1.220  ni  and,  where  not  irrigated  for  pas- 
ture or  hay  crojjs,  are  co^'ered  b}'  thick 
stands  of  big  sagebrush  {Artemisia  tri- 
dentata)  with  an  understory  of  herbac- 
eous plants.  In  the  southeastern  portion  of 
the  range,  the  transition  zone  between 
coniferous  forest  and  shrubs  is  charac- 
terized b}'  populations  of  small  fir  trees 
within  the  brush,  suggesting  a  recent 
downslope  movement  of  the  trees  (Fig. 
2).  The  cause  of  this  invasion  of  sage- 
brush by  white  fir  in  the  Warner  Moun- 
tains is  the  focus  of  this  paper. 

Relation  to  Other  Studies 

Many  observers  have  noted  invasions 
of  sagebrush  by  tree  sj)ecies  in  various 
parts  of  the  Intermountain  West.  Al- 
though pinyon  pines  (Pinus  rnonophylla 
and  P.  edulis)  and  junipers  {Juriiperus 
spp.)  are  the  most  common  invaders  of 
sagebrush  (Arnold  et  al.,  1964;  Black- 
burn and  Tueller,  1070;  Burkhardt  and 
Tisdale,  1969;  Cottam  and  Stewart,  1940; 
Wright  and  Fisser,  1968),  lodgepole  pine 
(Pinus  contorta)  also  has  been  found  ex- 
panding into  stands  of  Artemisia  (Patten, 
1969).  Moreover,  young  trees  of  ponder- 
osa pine  in  eastern  Oregon  and  Jeffrey 
pine  {Pinus  jeffrcyi)  in  eastern  Califor- 
nia may  be  readily  observed  within  sage- 
brush areas  along  forest  edges.  Although 
all  the  above  vegetation  changes  imply 
a  recent  establishment  of  trees  in  environ- 
ments formerly  unsuited  to  them,  studies 

'Assistant  Professor,  Department  of  Geogmphy,  University-  (i 


lia\'e  not  yet  documented  the  in^  asion  of 
the  relati^el}'  xeric  big  sagebrush  by  so 
mesic   a   species   as   white   fir. 

In  these  previous  studies,  the  initiation 
of  tree  establishment  is  often  found  to 
correlate  with  periods  of  intense  livestock 
grazing.  Cattle  may  deplete  the  herbac- 
eous plants  in  the  vegetation,  thereby  re- 
ducing the  full  utilization  of  the  habitat's 
resources  (e.g.,  soil  moisture);  this  "open- 
ing" of  a   formerly  "closed"   plant  cover 


public  domain  lands 
administered  by  the 
Bureau  of  Land  Manageme 


STUDY  SITE 


35 


Fig.    1.     The   Waiiier   Mountains    area    of   e.v- 
tieme    noitheastern    California. 


319 


320 


GREAT  BASIN  NATURALIST 


Vol.  35,  No.  3 


Fig.  2.     Eastern   slope   of   the   Warner   Mountains  with   the  study  area   in  the  middle  background. 


may  permit  the  establishment  of  species 
previously  excluded  (Robertson  and 
Pearse,  1943).  In  a  lodgepole  phie  inva- 
sion of  grassy  meadows,  Vankat  (1970) 
determined  that  most  invasive  trees  were 
established  immediately  after,  rather  than 
during,  a  time  of  grazing  by  sheep. 
Trampling  by  the  animals  apparently 
killed  seedling  trees,  but  the  heavy 
browsing  and  grazing  did  make  the  mea- 
dows susceptible  to  successful  invasion 
after  the   sheej)   were   removed. 

Another  cause  sometimes  imoked  to 
exj)lain  tree  advances  into  sagebrush  is 
climatic  fluctuation.  Patten  (1969)  and 
Arnold  et  al.  (1964)  both  suggest  that, 
during  abnormally  wet  j)eriods.  trees 
may  become  established  in  shrub  environ- 
ments typically  too  dry  for  them.  John- 
sen  (1962)  offers  a  \ariation  on  this 
theme  by  speculating  that  long  drouglits 
may  initiate  the  invasion  by  rechicing 
the  cover  of  brush  and  herbaceous  ])lants; 
when  moist  conditions  return,  trees  arc 
able  to  sprout  and  survive  in  the  "opened'" 
vegetation  stand.  This  sec[uence  of  events 
is  analogous  to  the  "opening"  of  a 
"closed"  plant  community  by  livestock 
grazing. 


Fire  suppression  is  often  invoked  to  ex- 
plain the  iiiA'asion  of  woody  growth, 
usually  trees,  into  grasslands  in  the  south- 
ern Intermountain  West  (e.g.,  Foster, 
1917;  Pearson,  1931;  .Tohnsen,  1962). 
Blackburn  and  Tueller  (1970),  moreover, 
suggest  that  a  decrease  in  fires,  together 
with  overgrazing,  accounts  for  the  inva- 
sion of  pinyon  pine  and  juniper  into 
brush  in  eastern  Nevada.  Abundant  evi- 
dence exists  suggesting  that  fires  retard 
the  spread  of  woody  plants  in  the  south- 
ern Intermountain  West,  although  inter- 
pretation of  tree  invasion  into  sagebrush 
as  a  resj)onse  to  fire  su])pression  is  com- 
plicated hv  the  fact  that  both  the  trees 
and  the  slwub',  suffer  from  freipieiit  fires. 

MiniioDs 

A  site  V)  km  soulli  of  the  town  of 
(>edar\ille  was  selec  IcmI  (oi-  sampling  the 
age  striK  liii-(>  of  ihe  \oiiiil;  lre(>s.  The  site 
appeared  to  he  represeiital  i\  e  of  the  ex- 
posure, slope.  cUkI  \  (^i^etatioii  character- 
ized b\  invading  white  fir  in  the  south- 
eastern Warner  Moinita.ins.  Five  plots, 
each  50  m  l)\  60  m,  were  located  at  in- 
tervals of  0.5    km  along  the  lower  limit 


Sept.  1975 


^^^LK•.   white  fik  I!NV\sTC)^^ 


321 


of  young  trees:  an  atlditional  plot  was  lo- 
cated in  a  stand  of  young  fir  trees  at  a 
higher  elevation  where  a  southeastern  (>\- 
posure  caused  a  high(>r  forest-shruh  tr.ni- 
sition.  The  selection  of  plots  along  the 
lower  Hmit  of  these  Ivvv^  wa^  designed 
to  estahlish  the  [jerioil  during  \vhicli 
white  fir  invaded  that  portion  of  the  area 
usually  considered  least  hospitahle  to  it. 
AA'ithin  each  plot,  all  trees  greater  than 
20  cm  dbh  were  cored  with  an  increment 
borer,  while  the  time  elapsed  since  tree 
establishment  was  estimated  to  be  (ught 
years  plus  those  indicated  by  the  tree 
rings.  Trees  with  diameters  smaller  than 
20  cm  were  recorded  by  estimated  height. 

Results  and  Discussion 

Although  invasion  began  between  1915 
and  1919,  most  white  fir  became  estab- 
lished between  1925  antl  1944  (Table 
1).  After  1944  a  decrease  in  successful 
tree  establishment  is  suggested  by  a  gap 
in  the  age  structure.  More  recent  repro- 
duction seems  improved,  judging  from  the 
relatively  large  number  of  trees  between 
0.5  and  2  meters  in  height.  The  absence 
of  dead  trees  of  any  size  precludes  the 
possibility  that  older  cohorts  suffered 
mortality,  an  e^-ent  which  would  compli- 


cate 
iu\-< 


nation  of  the  initial   tree 


Climate. —  The  years  correlated  with 
I  lie  beginning  of  tree  invasion  constitute 
the  driest  ])eriod  in  the  historical  record, 
i.e..  since  the  18()0s  (Fig.  3).  Consequent- 
ly, the  first  white  fir  sprouted  and  sur- 
vived, not  when  climatic  conditions 
would  ha\('   l)e(>n  most  favorable  for  this 

Tablk  1.  Age  structure  of  trees  in  sample 
plots  and  numbers  of  seedlings  by  height. 

Trees  whose  diameters  exceed  20  cm 
Year  of  establishment  Number  of  trees 


Prior  to  1915 

0 

1915-1919 

1 

1920-1924 

1 

1925-1929 

6 

1930-1934 

8 

1935-1939 

10 

1940-1944 

6 

1945-1949 

2 

1950-1954 

1 

1955-1959 

4 

Trees   wliose    diameters 

are    less  than   20    cm 

Height  HI   m 

Number  of  trees 

2-3 

8 

Less  than  2 

39 

1870-71 


Fig.  .3.  Running  five-year  means  of 
tation  years  (I  July- 30  June),  for  Cedai 
source:    U.S.   Weather  Bureau. 


Precipitation  Year 

F    the    percentage   of    average    precipitation,   based   on   precipi- 
le   and    Fort  Bidsvell    in   Surprise  Valley.  California.   Data 


522 


(,KEAT  BASIN  NATURALIST 


Vol.  33,  No.  3 


niesic  species,  but  when  drought  should 
have  hiuderod  ils  ostablishnieut.  The 
notion  that  dry  conditions  might  have 
favored  tree  invasion  l)\  reducing  the 
sagebrush-grass  cover  is  untenai)le,  be- 
cause a  drought  so  severe  as  to  decrease 
the  xerii  sagebrush  toidd  not  possibh' 
siniuhaneouslv  inc  reas(>  the  niesic  white 
fir. 

The  coincidence  of  drought  and  initial 
establishment  of  the  white  fir  in  this 
study,  then,  suggests  that  the  trees  in- 
vaded in  spite  of  the  weather  rather  than 
because  of  it.  Moreover,  it  is  apj^arent 
that  nonclimatic  factors  prevented  tree 
invasion  during  the  more  moist  periods 
existing  in  the  study  area  prior  to  1910. 

Fire  supprkssion. —  Although  the 
Modoc  Forest  Reser^  e  was  established  in 
1904,  wildfires  in  the  Warner  Mountains 
apparently  continued  to  be  connnon  un- 
til after  the  creation  of  the  Civilian  Con- 
servation Corps  in  1933;  the  year  1924, 
for  example,  is  reported  to  have  been  a 
particularly  serious  fire  year  (Cook,  n. 
d.).  More  specifically,  information  from 
the  U.S.  Forest  Service  indicates  that  the 
area  immediately  adjacent  to  the  study 
plots  was  burned  by  wild  fires  l)etween 
1921  and  1930,  aiid  that  fires  were  com- 
mon within  the  national  forest  during 
that  decade  (U.S.  Forest  Service.  197-1). 
Supporting  the  impression  that  fire  su]i- 
[)ression  was  not  well  developed  in  the 
region  by  the  time  of  tree  invasion,  the 
rangeland  outside  of  the  forest  resei-\'e, 
and  in  which  the  white  fir  became  estab- 
lished, was  not  given  official  protection 
until  passage  of  the  Taylor  Grazing  Act 
in  1934.  The  Bureau  of  Land  Manage- 
ment office  in  Susanville,  California,  out 
of  which  the  federal  rangelands  in  the 
region  are  administered,  reports  no  rec- 
ords of  fire  sui)pression  activities  on  the 
east  slope  of  the  Warner  Mountains 
prior  to  1930  (U.S.  Bureau  of  Land  Man- 
agement, 1974). 

The  State  of  (California  was  not.  and  is 
not,  res})onsible  for  fire  control  in  the 
study  area,  although  it  presently  protects 
private  rangelands  west  of  the  Warner 
Mountains  (California  Division  of  For- 
estry, 1975).  Moreover,  the  state  did  not 
have  any  system  for  fire  supj)ression 
during  the  initial  years  of  tree  establish- 
ment, and  its  fire  protection  policies  can- 
not be  considered  to  have  been  effective 


until     after     1943     (Clar,     1969;     Davis, 
1963). 

It  seems  \ali(l  to  conclude  that  suc- 
cessful fire  suppression  on  the  east  slope 
of  the  southern  Warner  Mountains  was 
not  effective  until  after  the  period  of 
initial  tree  establishment.  Although  sub- 
sequent fire  control  has  probably  aided 
the  maintenance  of  the  trees,  it  cannot 
be  invoked  to  exjilain  the  initiation  of  tree 
advanc(^  dowiislope. 

Grazing. —  The  intensity  of  grazing 
by  domestic  livestock  in  the  southeastern 
Warner  Mountains  has  varied  greatly 
since  the  initial  settlement  of  Surprise 
Valley,  inniiediately  east  of  the  Warner 
Range,  in  1(S(>1.  The  number  of  cattle  on 
raiichcs  in  Modoc  County  as  a  whole  was 
little  changed  between 'lS90  and  1945, 
but  it  doid)letl  in  the  following  twenty 
years;  resident  sheep,  by  contrast,  in- 
creased rapidly  between  1890  and  1930, 
but  by  1940  the\'  had  declined  j)recip- 
itously    (Fig.    4). 

Much  of  Modoc  Count}'  is  heavily  for- 
ested, thus  restricting  grazing  by  domestic 
livestock  to  areas  of  brush  or  grass.  Such 
habitats  in  the  Warner  Mountains  and  ad- 
jacent A'alleys  have  supported  large  num- 
bers of  animals,  particularly  sheep,  over 
the  last  century.  Contributing  to  the 
heavy  grazing  of  these  rangelands  was 
the  seasonal  migration  of  sheep  from  the 
inountains  in  summer  to  the  semiarid 
lowdands  of  northern  Nevada  in  winter, 
a  pattern  well  established  by  the  1870s 
(Olmsted,  1937).  Moreover,  sheep  drives 
from  Idaho  and  Oregon  to  shipping 
points  in  western  Nevada  jiassed  through 
the  Warner  Mountains   (Olmsted,   1937). 


Fig.  !•.  Niiinl)('i->  1)1  uitll.t"  and  slu'cj)  on 
i-iuicliofi  in  Mcxloc  (bounty.  Data  soiirco:  IT.S. 
Bureau   of  Census. 


Sept.  1975 


IITF.    FIU    IN\-ASI()X 


323 


By  1900  the  ranges  of  Modoc  County  are 
said  to  have  been  greatl\-  overgrazed 
(Brow^i,    1951;   Pease.   196-3). 

Establishment  of  f(>deral  forest  res(>r\('s 
(later  to  become  national  forests)  in  the 
early  19()0s  apj^arently  (hd  littl(>  to  re- 
(hice  innnediately  the  grazing  j)ressures 
in  the  higher  elevations  of  Modoc 
County,  including  the  Warner  Moun- 
tains. TYansient  sheej)  continued  to  be 
driven  across  national  forest  land  from 
Oregon  initil  the  Forest  Service  banned 
such  use  in  1914  (Tierney,  1946).  Pease 
(1965)  suggests  that  the  elimination  of 
grazing  by  transients  promjited  the  estab- 
lishment of  new  sheep  ranches,  with  resi- 
dent flocks,  in  Surjirise  Valley;  this  con- 
tributed to  the  ra])id  increase  of  resident 
sheep  in  Modoc  County  l^etween  1910 
and  1920.  Also  accentuating  the  heavy 
grazing  pressures  at  this  tiine,  the  Forest 
Service  intentionalh'  allowed  overstock- 
ing on  national  forest  lands  in  north- 
eastern California  during  World  W^ar  I  to 
help  meet  war  demands  for  food  and  wool; 
even  after  the  war,  heavy  stocking  con- 
tinued because  it  was  felt  a]n-u])t  reduc- 
tions in  livestock  numbers  might  ha^e 
created  economic  hardship  for  area 
ranchers    (Tierney.    1946). 

Even  while  the  national  forest  lands 
were  under  nominal  regulation,  the  pub- 
lic domain  continued  to  be  completely 
free  and  open  range.  These  latter  federal 
lands  in  northwestern  Nevada  served,  in 
part,  as  wintering  grounds  for  sheep 
that  were  moved  from  California  during 
the  autumn  season.  Olmsted  (1957) 
claims  that  150,000  sheep  were  grazed  in 
Surprise  Valley  in  1920,  and.  when  com- 
j)ared  to  resident  sheep  rejiorted  on 
ranches  in  all  of  Modoc  Coimty  in  that 
year  (109,000),  it  is  apparent  that  much 
use  of  Surprise  Valley  ranges,  including 
much  ])ublic  domain  acreage,  was 
by  transient  flocks.  The  peak  in  grazing 
pressure  by  sheep  in  the  1920s  was  ap- 
|)arently  even  greater  than  that  suggc^sted 
by  the  numbers  of  resident  sheej). 

The  end  of  uncontrolled  sheep  grazing 
on  the  public  domain  came  with  the  pas- 
sage of  the  Taylor  Grazing  Act  in  1934 
(Olmsted,  1957;  Pease,  1965).  With 
more  stringent  regulations  against  tran- 
sient flocks,  the  public  lands  coidd  jio 
longer  be  used  by  migratory  sheep  herd- 
ers. 

The   grazing   history   suggests    that   the 


white  fir  iii\asion  (oinc  ided  with  the 
])eak.  and  tli(>  period  immediately  fol- 
lowing the  peak,  (jf  >lieep  grazing  in  the 
region.  Sheep  may  have  reduced  the  cox- 
erage  of  grass  and  shrubs,  thereb}'  in- 
creasijig  the  a^■aiiabilit}'  of  soil  moisture 
and  allowing  the  establishment  of  seed- 
ling trees.  Sheep  browse  shrubs,  hut  not 
conifers,  on  ^^  inter  range,  thus  encour- 
aging   the    tree    invasion. 

Conclusions 

The  evidence  suggests  that  grazing  b}" 
domestic  livestock  altered  the  sagebrush- 
grass  vegetation  on  the  east  slope  of  the 
Warner  1\ fountains  and  allowed  the  estab- 
lishment of  white  fir  seedlings.  Yet,  while 
grazing  seems  responsible  for  the  initia- 
tion of  tree  in^'asion,  the  accelerated  rate 
of  tree  establishment  during  the  period 
1935-1944  may  have  been  the  result  of 
increased  precipitation  on  the  openings 
in  the  vegetation  cover  ])roduced  by 
grazing.  Such  an  explanation  would  ac- 
count for  the  decrease  in  tree  establish- 
ment after  1944  because,  by  that  time, 
the  plant  cover  would  have  sufficiently 
recovered  from  the  effects  of  the  earlier 
heavy  grazing  to  minimize  the  avail- 
ability of  suitable  seedbeds  and  soil  mois- 
ture. The  cause  of  the  present  abundance 
of  seedling  trees  is  more  obscure,  but  may 
be  related  to  a  resurgence  of  grazing  pres- 
sure due  to  increasing  numbers  of  cattle 
in  recent  years    (Fig.  4). 

In  the  northern  Intermountain  West 
generally,  grazing  by  domestic  livestock 
may  be  adequate  to  explain  the  vsdde- 
spread  invasion  of  trees  into  sagebrush- 
grass  vegetation  dialing  the  late  nine- 
teenth and  early  twentieth  centuries.  The 
example  from  California,  discussed  in 
this  paper,  suggests  that  fire  suppression 
has  been  too  recent,  except  perhaps  lo- 
cally, to  correlate  with  these  tree  invas- 
ions. (Control  of  fire  may  account  for 
tre(^  establishment  in  other  vegetation 
types,  however,  during  this  time.)  Re- 
gional (dimatic  fluctuations,  by  them- 
selves, also  seem  inadequate  to  account 
for  the  (>x])ansion  of  trees  into  sagebrush- 
grass  vegetation  during  the  latter  half  of 
the  1800s,  a  time  characterized  by  "fluc- 
tuating Init  below  axerage  moisture"  in 
western  North  America  (Fritts,  1965). 
This  portrayal  of  climate  does  not  suggest 
conditions   sufficiently   wet   to   favor  tree 


324 


(iKEAT  BASIN  NATUKALIST 


Vol.  .35,  No.  3 


growth  in  formerly  xeric  brush.  More- 
over, tree  invasion  in  the  Warner  Moun- 
tains began,  not  during  a  wet  period,  but 
during  an  extended  time  of  below  average 
precipitation.  Grazing  by  domestic  live- 
stock, then,  remains  the  most  likely  gen- 
eral cause,  apjilicable  on  a  regional  basis, 
to  account  for  the  widespread  ijivasion  of 
sagebrush  vegetation  by  tree  species. 
Periods  of  extended  drought  may,  in  cer- 
tain ])laces,  accentuate  the  plant  Vover  de- 
terioration caused  by  grazing,  thus  en- 
couraging tree  estab'lishment  once  more 
moist  conditions  returii. 


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The  pinyon-iuniper  typo  of  Arizona:  ef- 
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Blackburn,  W..  and  P.  Tufxler.  1970.  Pin- 
yon  and  juniper  invasion  in  Ijlack  sage- 
brush communities  in  east-central  Nevada 
Ecol.  51:841-848. 

Brown,  W.  1951.  California  Northeast:  The 
bloody  ground.  Oaklaiifl.  California.  Bio- 
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Burkhardt.  J.,  and  E.  Tisdale.  1969.  Nature 
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CiAR.  C.  1969.  Evolution  of  California's  Wild- 
land  Fire  ProtcM  tion  System.  Sacramento. 
St^ate  of  Califonna.  The  Resources  Agency. 
?5  p. 

Cook.  F,  n.  d.  History  of  Modoc  Counlv.  Vol- 
cano. California.  California  Tiavelei-  Inc 
104  p.. 

CoTTAM,  W..  AND  G.  Stewart.  1910.  Plant  suc- 
cession  as  result  of  grazing  of  meadow  desic- 


cation   by    erosion    since    settlement    in    1862 
J.   For.   38:613-626. 

Davis.  L.  1965.  The  economics  of  wildlife  pro- 
k'ction  with  emphasis  on  fuel  break  svstems 
Sacramento,  State  of  California.  The  Re- 
sources  Agencv.     166   p. 

Iuxster.  ,J.  1917.  The  spread  of  timbered  areas 
HI   central   Te.xas.     ,J.   For.    15:442-145. 

luuTTs.  H.  1965.  Tree-ring  evidence  for  cli- 
matic ciumgcs  in  western  North  America 
Mon,   Weather   Rev.   93:421-443. 

.loiiNSEN.  T.  1962.  One-seed  juniper  invasion  of 
northern  An/.ona  grasslands.  Ecol.  Monog. 
32:187-207.  ^ 

Oimsted,  p.  1957.  The  Nevada-California-Ore- 
gon border  triangle:  a  study  in  sectional  his- 
tory. M.A.  tliesis,  Univ.  Nevada,  Reno. 
284  p. 

Patten,  13.  1969.  Succession  from  sagebrush 
to  mixed  conifer  forest  in  the  northern 
Rocky  Mountains.  Am.  Midi.  Nat  82- 
229-240. 

Pe.\rson.  G.  1931.  Forest  types  in  the  south- 
west as  determined  bv  climate  and  soil.  U  S 
Dept.   Ag.   Tech.    Bull.   247. 

Pe.\se.  R.  1965.  Modoc  County:  a  geographic 
time  continuum  on  the  California  volcanic 
tableland.  Cniv.  California  Publ.  Geog  Vol 
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Robertson,  J.,  and  C.  Pearse.  1945.  Artificial 
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present.  Alturas,  California,  Modoc  County 
Superintendent     of     Schools.      139     p. 

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Letter  to  author. 

U.S.    Forest    Service.      |i)7k     Letter    to    author. 

Vankat,  ,1.  1970.  Vegetation  cliange  in  Se- 
quoia National  Park.  California.  Ph.D.  dis- 
sertation.   Univ.    California.   Davis.      197   p. 

Wright.  .[..  and  H.  Fisser.  1968.  Juniperus 
osteospeiiua  in  northwestern  Wyoming:  their 
(bstribution  and  ecology.  Sci.  Monograph 
No.  7.  Laramie.  Wyoming.  LTniv.  Wvoming 
Ag.   Exper.   Stat.     31    p. 


MORPHOLOGY  OF  EPHEMERAL  AND  J^ERSISTENT  LEAVES 

OF  TFIREE  SUBSPECIES  OF  BIG  SAGEBRUSH 

GROWN  IN  A  UNIFORM  ENVIRONMENT 


W.  T.  McDonough.  R.  ().  Hainl 


(I    I',,    15.  Canipbe: 


Abstract.-  Measui-ciiipnts  uere  made  of  inorj)h()li)gi(  al  (  liai  a(  tciistic  s  of  ephenieral  and  j)er- 
sistent  leaves  of  three  subspecies  of  big  sagebrush  (Arlc/iiisia  Iri/lr/ila/a  Nutt.)  grown  in  a  uni- 
form environment.  No  morphological  feature  clearh'  separaterl  the  subspecies.  Genetic-environ- 
mental interactions  on  leaf  morphology  apparently  reduce  its  \alue  as  a  sole  criterion  for  (hstin- 
guishing  the  subspecies. 


Taxonomic  subdivisions  of  a  species 
may  differ  in  morphological  and  physio- 
logical characteristics,  and  these  (hffer- 
ences  may  be  genetically  (jr  einirf)n- 
mentally  controlled  to  varying  extents. 
There  is  considerable  interest  in  subspe- 
cies of  big  sagebrush  (Artemisia  tridcnt- 
ata  Nutt.)  because  of  differences  in  palat- 
ability,  habitats  occuj^ied,  and  aggressive- 
ness-characteristics of  interest  to  range 
managers  fMcArthur  et  al.,  1974).  Iden- 
tification is  based  chiefly  on  the  growth 
form  and  morphology  of  leases  and  on 
the  inflorescence  of  plants  collected  from 
their  normal  habitats  (Winward,  1970). 
Biochemical  differences  have  also  been 
noted  in  reproducti^•e  and  vegetative 
parts  (Winward  and  Tisdale,  1969; 
Hanks  and  Jorgensen,  1973;  Stevens  and 
McArthur,  1974).  Biochemical  patterns 
are  less  subject  to  enviromnental  mod- 
ification (Winward,  A.  H.,  1975.  Per- 
sonal communication.  Oregon  State 
Univ..  Corvallis)  but  are  inconvenient 
for  use  in  field  indentification. 

To  determine  the  extent  to  which  leaf 
morphology  of  big  sagebrush  subs])ecies 
is  genetically  rather  than  environmen- 
tally controlled,  we  grew  plants  of  three 
subspecies  under  the  same  greenhouse 
conditions  to  identify  distinguishing  leaf 
characteristics   that   persist  in   a   uniform 


environment. 


Methods 


Potted  year-old  plants  of  each  of  three 
subspecies,  basin  big  sagebrush  (tridcnt- 
ata  Nutt.),  mountain  big  sagebrush  (vasc- 
yana  Rydb.),  and  Wyoming  big  sage- 
brush {wyomingensis  Beetle),  were  estab- 
lished from  seeds  collected  the  fall  of  1973 
near  the  Sheep  Ex})erimental  Range,  Du- 

'Plant  physiologist,  range  scientist,  and  range  tcclmici 
and  Range  Experiment  Station,  Ogden,  Utali  84401.  stati 
tained     in    cooperation    with    Utah     State     University,     Logan. 


bois,  Idaho.  The  (25  cm)  pots  were  ran- 
domly arranged  on  a  greenhouse  bench 
and  given  routine  care.  From  the  50 
]jlants  of  each  subspecies,  6  plants  and 
10  mature  leaves  (ephemeral  and  per- 
sistent; from  each  plant  were  randomly 
selected  for  determinations  of  length, 
width,  length /width,  number  and  depth 
of  lobes.  Lobes  were  distinguished  from 
occasional  minor  dentations  by  the  pres- 
ence of  a  secondary  Acin.  Leaves  were 
fixed  to  cards,  photographed,  and  the 
35-mm  transparencies  projected  for  mea- 
surements. 

At  the  time  of  sampling,  growing 
plants  averaged  27  cm  in  height  and  had 
from  6  to  1 7  branches.  Significance  of  dif- 
ferences at  the  5  percent  level  was  eval- 
uated by  variance  analysis  and  multiple 
range  tests. 

Results 

Representative  leaves  are  shown  in 
Figure  1;  mean  dimensioius,  in  Table  1. 
Only  limited  differentiation  was  observed 
for  both  types  of  lea\(vs  among  the  sub- 
species. 

In  the  e|)hemeral  leaves,  variation  in 
lobe  tium])er  was  not  significant.  Width 
and  length/width  differed  only  in  subsp. 
vascyana.  Subsp.  wyomingensis  differed 
from  subsp.  trident  at  a  in  lobe  length  and 
from  subsp.  vaseyana  in  lobe  depth. 

In  the  jiersistent  leaves,  length  dis- 
tinguishes subs]).  wyomingensis  from  the 
other  subspecies  and  length/width  dis- 
tinguishes wyomingensis  from  tridentata. 
Number  of  lobes  was  uniformly  three 
in  all  subs]iecies,  and  lobing  generally  was 
so  shallow  that  attempted  measurements 
were  uiu'eliable. 

ui,     respective!  V.     USiXV     I'orest     Service.     Interniounlain     P'orcl 
med    in    I.ogan.     l.'tah.     at    l-"orestry    Sciences    Lal)or,'ilory.     mam- 


325 


326 


GREAT  BASIN  NATURALIST 


Vol.  35,  No.  3 


T.VBLE     1  . 

brush   grown 


Mean    dimensions'     ± standard    devi 
1   a    unifonn    environment. 


of     til 


iul)species    of    l)ig    sage- 


Subspecies 

Length 
(mm) 

Width 

(mm) 

Length/Widtli 

Number 

of 

lobes 

Lobe  deptli 
(mm) 

Ephemeral  Leaves 

Tridcnta/a 

Vaseyana 

Wyomingeusis 

50.9=1     ±7.1 
47.9=1''   ±5.4 
43.3''     ±4.7 

19.3=1 
15.6" 
19.6=1 

±6.8 
±5.9 
±6.1 

5.0=1     ±1,2 
3.6"     ±  1 .8 
2.5=^     ±1.0 

Persistent  Leaves 

4.0=1 
3.8=1 
4.0=1 

±1.1 
±0.9 
±1.0 

9.8=1"   ±^j 
8.4"     ±3.4 
11.4=1     ±3.6 

Tridentata 

Vaseyana 

Wyomingeusis 

14.3=1     ±2.5 
14.9=1     ±3.0 
11.9>^     ±2.3 

3.1=1 
3.4=1 
3.0=1 

±0.7 
±0.7 
±0.8 

4.9=1     ±1.2 
4.5=1"   ±0.6 
4.1"     ±0.8 

3=1 
3=1 
3=1 

- 

'Means    for    anv    measurement 


same    letter    in    superscript 


Fig.  1.  Ephemeral  (above)  atid  persistent 
(below)  leaves  of  sagebiiish  grown  in  a  uniform 
environment — subsp.  tridentata  (T).  vaseyana 
(V).    and    wyomingeusis    (W). 

Wiiiward  (197()j  exaniiiiofl  only  per- 
sistent leaves  of  field-grown  plants.  Only 
differences  in  length/width  were  con- 
sidered usefnl  in  separating  the  subspe- 
cies. Values  of  5.6,  4.0,  and  3.1  were  as- 
signed, respectively,  to  subsp.  tridentata, 
vaseyana,  and  wyomingensis.  I.ea^-es  of 
plants  grown  under  uniform  conditions 
are  in  the  same  sequence  but  have  some- 
what divergent  values  that  do  not  differ- 
entiate  subsp.    vaseyana    (Table    1). 


Conclusions 

Even  with  careful  measurements,  no 
characteristic  of  ephemeral  or  persistent 
leaA'es  from  plants  growai  under  uniform 
conditions  was  found  to  clearly  separate 
the  three  subspecies.  Only  subsp.  vase- 
yana is  distinguishable  by  two  character- 
istics of  ephemeral  leaves  and  subsp. 
wyomingensis  by  one  characteristic  of 
persistent  leaves.  Distinguishing  leaf  mor- 
phology may  be  so  dependent  upon  en- 
vironmental inodification  that  it  is  not 
uniformly  useful  in  identification.  Al- 
ternatively, distinct  morphological  dif- 
ferences may  be  obscured  by  genetic  in- 
trogression  among  subspecies  and  hybrids 
(Plummer,  A.  P.,  1975.  Personal  com- 
munication. U.S.  Dep.  Agric.  For.  Serv., 
Intermt.  For.  and  Range  Exp.  Stn.,  Og- 
flen,  Utah)  or  because  insufficient  num- 
bers of  plants  are  sampled  from  the  same 
or  other  areas. 


r.ITER  VrURF.    CiTF.n 

Hainks.  I).  I...  \M)  K.  R.  .JoRr.ENSEN.  1973. 
Chroniatograpliic  identification  of  big  sage- 
brush  seed.   ,1.   Range  Manage.  26:304. 

McArtiiur.  E.  D..  B.  C.  Giunta.  and  A.  P. 
Pr.uMiMER.  1974.  Shrubs  for  restoration  of 
depleted  ranges  and  disiurbed  areas.  Ltali 
Sci.    35:28-33. 

Stevens,  R..  and  E.  D.  McAruiur.  1974.  A 
siini)le  field  teduiiciue  for  identification  of 
some  sagebrusii  taxa.  J.  Range  Manage. 
27: '>25- 526. 

\\'iN\\ARD.  A.  H.  l'>70.  Taxonomy-  and  ecology 
of  big  sagebruslr  Ph.D.  Thesis.  Univ.  Idaho. 
Moscow.     80   ]). 

\\IN\VARD.      A.      H..       \.M)      K.      W.      TlsDAI.E.        1969. 

.A    simplified    .hrninal    iiiriho.l    for    sagebrusii 
id(Mitifi(.iti.)ii.    I   iiiv.   Id.ih..  Stn.  Note   11.  2  p. 


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TABLE  OF  CONTENTS 

Urosaurus  and  its  phylogenetic  relationship  to  Uta  as  determined  by  oste- 
ology and  myology  (Reptilia:  Iguanidae).  Charles  Fanghella,  David  F. 
Avery,   and   Wilmer  W.   Tanner   245 

Distribution  and  adundance  of  the  black-billed  magpie  (Pica  pica)  in  North 

America.   Carl  E.   Bock  and  Larry  W.  Lepthien  269 

Nectar  composition  of  hawkmoth-visited  species  of  Oenothera  (Onagraceae). 

Robert  E.   Stockhouse,   II   273 

A  revision  of  the  nearctic  species  of  Clinohelea  Kieffer  (Diptera:   Ceratopo- 

gonidae).  William  L.  Grogan,  Jr.  and  Willis  W.  Wirth  275 

Basidiomycetes  that  decay  junipers  in  Arizona.  R.  L.  Gilbertson  and  J.  P. 

Lindsay   288 

Body  size,  organ  size,  and  sex  ratios  in  adult  and  yearling  Belding  ground 

squirrels.  Martin  L.  Morton  and  Robert  J.  Parmer 305 

Photoperiodic    responses    of    phenologically    aberrant    populations    of    pierid 

butterflies    (Lepidoptera).    Arthur    M.    Shapiro    310 

Additional    records    of    reptiles    from    Jalisco.    Me.xico.    Philip    A.    Medica. 

Rudolf   G.    Arndt.    and   James   R.    Dixon    317 

Invasion  of  big  sagebrush  (Artemesia  Iridentata)  by  white  fir  {Abies  con- 
color)  on  tlie  southeastern  slopes  of  the  Warner  Mountains,  California. 
Thomas  R.  Vale  ---     319 

Morphology  of  ephemeral  and  persistent  leaves  of  three  subspecies  of  big 
sagebrush  grown  in  a  uniform  environment.  W.  T.  McDonough.  R.  O. 
Harniss,   and  R.   B.  Campbell  325 


IE  GREAT  BASIN  NATURALIS 

me  35  No.4         December  31, 1975        Brigham  Young  Universi 


■OOL 


** 
•^1   - 


^iSF^' 


^^Sl^ 


^^-y^W 


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,  Zoology;  Stanley  L. 

Welsh,  Botany;  Clayton  M.  White,  Zoology. 
Ex  Officio  Editorial  Board  Members.  A.  Lester  Allen,  dean.  College  of  Biological  and 

Agricultural  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  continuously  published  from  one  to  four  times  a  year  since  then  by  Brigham 
Young  University,  Provo,  Utah.  In  general,  only  original,  previously  unpublished 
manuscripts  pertaining  to  the  biological  natural  history  of  the  Great  Basin  and  western 
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The  Great  Basin  Naturalist 

Published  at  Provo,  Utah,  by 
Brigham  Young  University 


Volume  35 


December  31,  1975 


No.  4 


ENDANGERED,   THREATENED,   EXTINCT,   ENDEMIC, 
AND   RARE  OR  RESTRICTED   UTAH   VASCULAR   PLANTS 

Stanley  L.  Welshi,  N.  Duane  Atwood-,  and  James  L.  ReveaP 

Abstract. —  The  status  of  382  vascular  plant  taxa  with  distribution  in  Utah  is  presented.  Some  66 
species  are  possibly  endangered,  198  threatened,  7  extinct,  and  20  extirpated  within  the  state;  4  spe- 
cies have  questionable  taxonomic  status.  Included  in  the  list  are  nearly  225  species  of  endemic 
plants,  many  of  which  are  among  the  possibly  endangered,  threatened,  and  extinct  or  extirpated 
plants.  Bibliographic  citations,  type  locality,  status,  and  distribution  by  counties  is  included  for  each 
species  or  infraspecific  taxon.  Two  new  species  are  described:  Psoralen  pariensis  and  Eriogonum 
natum.  One  new  name.  Astragalus  barnebyi,  is  proposed;  and  one  new  variety,  Eriogonum  umbel- 
latum  var.  deserticum,  is  proposed.  The  following  new  combinations  are  made:  Cycladenia  hum- 
ilis  var.  jonesii;  Aralia  racemosa  ssp.  bicrenata;  Heterotheca  jonesii;  Hymenoxys  depressa;  Xantho- 
cephalum  sarothrae  var.  pomariense;  Thelypodium  integrifolium  var.  complanatum;  Thely podium 
sagittatum  var.  ovalifolium;  Arenaria  kingii  var.  plateauensis;  Psorothamnus  thompsonae;  Najas  caes- 
pitosus;  Oenothera  gouldii;  Eriogonum  corymbosum  var.  revealianum;  Penstemon  humilis  var.  ob- 
tusifolius;  Penstemon  lentus  var.  albiflorus;   and  Viola  purpurea  var.  charlestonensis. 


The  vascular  plant  flora  of  Utah  is 
both  large  and  complex.  Its  components 
are  diverse,  representing  numerous  flor- 
istic  elements  from  many  parts  of  North 
America,  including  unique  and  provin- 
cial elements  restricted  to  the  state.  Spe- 
cies of  many  major  geographical  group- 
ings of  plants  occur  within  the  multiplic- 
ity of  habitats  available  within  Utah,  all 
to  a  greater  or  lesser  degree  of  their  entire 
range.  Some  of  these  species  are  at  the 
edge  of  their  total  range,  and  these  occur 
in  smaller  portions  of  the  state.  Other 
taxa  occur  only  in  one  or  few  peculiar, 
limited  edaphic  situations  or  habitats, 
while  others  are  more  widespread  and 
cover  a  broad  altitudinal  or  latitudinal 
expanse  of  Utah. 

Those  plants  that  occur  only  within 
the  state,  or  within  the  natural  basins  that 
overlap  the  artificial  political  boundaries 
of  the  state,  are  known  as  local  endemics. 
Their  range  can  be  widespread  within 
Utah,  but  more  often  they  are  restricted 
in    distribution    to    very    limited    areas. 


These  are  plants  that  are  of  much  inter- 
est to  scientists,  because  they  present  liv- 
ing proof  of  the  origin  and  evolution  of 
species,  origin  of  floras,  and  indications  of 
relationships  of  plant  species.  These 
plants  are  those  entities  which  have  orig- 
inated here  or  are  mere  remnants  of  spe- 
cies which  have  had  a  much  broader  area 
of  distribution  in  the  past. 

The  impress  of  man  and  his  activities 
onto  the  natural  habitats  of  Utah  has  re- 
duced the  area  available  to  most  native 
or  indigenous  plants.  Those  species  of 
broad  extent  and  wide  ecological  toler- 
ances have  withstood  these  activities  best, 
with  only  a  reduction  in  their  range  and 
number.  Less  frequently  their  range  has 
increased  in  size  as  less-tolerant  plants  of 
adjoining  areas  have  been  reduced.  How- 
ever, many  of  the  most  unique  species 
have  areas  of  distribution  that  are  very 
small,  with  only  a  few  known  individuals. 
In  some  examples  the  species  have  ap- 
parently ceased  to  exist  in  the  Utah  flora. 
In    most    cases    the    new    habitats    made 


^Department     of    Botany,     Brigham     Young     Universily,     Provo,     Utah     84602. 
-Bm-eau   of   Land    Management,    Cedar   City,    Utah    84720. 

^Departments   of   Botany,    University   of   Maryland,    College   Park,    Maryland   20742,    and   National    Museum    of   Natural    His- 
tory,    Smithsonian    Institution,    Washington,     D.C.     20560. 


327 


328 


GREAT   BASIN   NATURALIST 


Vol.  35,  No.  4 


available  by  the  acthdties  of  man  have 
been  occupied  by  introduced,  cultivated, 
and  adventive  plants  from  the  Old  World. 
These  are  the  crop  plants  and  weeds  of 
modern  agricultural  and  industrial  so- 
ciety. 

The  phalanxes  of  intolerant  native 
plants  have  been  retreating  under  the  im- 
pacts of  agriculture  and  grazing  for  more 
than  a  century.  Industrial  development 
during  most  of  that  period  was  limited 
in  extent,  if  not  in  effect.  In  Utah  most 
of  these  activities  were  restricted  mainly 
to  the  broad  valleys  and  river  basins, 
where  plant  communities  that  were  rela- 
tivel}^  fragile  are  now  almost  totally 
lacking  within  the  state.  However,  these 
sites  apparently  contained  few  of  the  nar- 
rowly endemic  plant  species.  With  the 
advent  of  the  second  half  of  the  twentieth 
century,  there  has  occurred  a  resurgence 
of  economic  activities,  mineral  explora- 
tion, and  a  greater  use  of  the  public  lands 
which  hitherto  had  been  considered  as 
useful  (if  considered  useful  at  all)  only 
for  grazing  and  watershed.  The  rapid 
spread  of  industrial  development  into  pre- 


viously undeveloped,  low-elevation,  arid 
lands  in  the  southern  portions  of  the  state 
is  all  the  more  impressive  when  one  con- 
siders that  most  of  the  narrowly  restric- 
ted plants  occur  in  those  areas  (Figs.  1 
and  2).  Plant  species  which  were  once 
remote  from  the  impacts  of  civilization — 
industrial,  agricultural,  or  recreational  ac- 
tivities— are  now  threatened  not  only  by 
the  effects  of  ranching,  construction,  and 
off-road  travel,  but  even  by  the  very 
ageiicies  of  government  which  are  estab- 
lished by  law  to  oversee  in  the  public 
trust  the  proper  use  and  protection  of  the 
public  lands.  At  the  present  time,  hardly 
a  part  of  Utah,  even  that  set  aside  as 
national  parks,  monuments,  or  wilderness 
areas,  is  safe  from  degradation  by  masses 
of  people  or  by  those  seeking  to  exploit  the 
very  natural  resources  and  features  these 
unique  areas  were  established  to  protect. 
Inroads  into  the  most  remote  and  most 
arid  portions  of  the  state  now  guarantee 
further  reduction  of  the  unique  flora  of 
Utah.  Naturally,  those  entities  that  will 
suffer  greatest  from  the  commercializa- 
tion of  the  state  will  be  those  which  have 
specific  and  naturally  restricted  areas  of 


Fig.  1.  Distributional  incidence  of  endangered, 
threatened,  extinct,  or  extirpated  Utah  plants  by 
county;  total  number  is  greater  than  the  total 
for  the  state  due  to  overlap. 


Fig.  2.  Distribution  of  endangered,  threatened, 
extinct,  and  rare  or  restricted  Utah  plants,  by 
phytogeographical  subdivision. 


Dec.  1975 


WELSH,   ET   AL.:    ENDANGERED    UTAH    PLANTS 


329 


distribution.  Only  the  most  enlightened 
management  and  protection  from  un- 
reasonable exploitation  will  ensure  the 
continuation  of  the  rich  indigenous  flora 
that  these  plants  represent. 

Commerical  exploitation  need  not  bring 
about  the  demise  of  species,  although  the 
basic  nature  of  the  flora  will  change.  It 
is  a  fact  of  biology  that  in  even  the  most 
enlightened  and  carefully  planned  devel- 
opment, there  will  be  a  reduction  in  the 
native  vegetation.  Reclamation  attempts 
will  be  made  with  Old  World  introduc- 
tions, or  with  selected  ecotypes  of  indig- 
enous plants,  and  not  with  those  that  oc- 
curred in  the  region  prior  to  its  modifi- 
cation. Both  of  these  types  of  plants  can 
hardly  replace  the  quality  of  those  cleared 
and  destroyed  by  construction  activities. 
Revegetation  of  a  disturbed  site  can  be 
more  or  less  productive  from  an  economic 
standpoint,  but  natural  plant  succession 
will  require  generations  before  any  real 
resemblance  of  the  natural  vegetation 
will  again  be  possible,  and  then,  depend- 
ing on  the  nature  of  the  disturbance,  the 
native  vegetation  occupying  the  site  can 
be  of  an  entirely  different  composition 
than  formerly.  Indigenous  taxa  with 
specific  habitat  requirements  will  not 
survive  if  those  habitats  are  altered,  and 
there  is  no  known  technology  which  can 
simulate  some  of  the  peculiar  habitats 
present  in  the  state — which  are  now  oc- 
cupied by  narrowly  restricted  plants — and 
no  technology  can  ever  replace  an  extinct 
species    (Reveal  1973b). 

Because  of  the  rate  at  which  Utah  is 
being  impressed  by  all  the  forces  of  a 
modern  society,  it  seems  important  that 
the  endangered,  threatened,  extinct,  en- 
demic, and  rare  or  restricted  vascular 
plants  of  Utah  be  inventoried,  and  that 
their  known  or  historic  areas  of  distribu- 
tion be  outlined.  That  task  is  the  basic 
goal  of  this  paper,  but  this  is  only  a 
pioneering  effort.  Much  work  remains  to 
be  done. 

While  the  present  paper  was  in  prep- 
aration, the  secretary  of  the  Smithsonian 
Institution,  S.  Dillon  Ripley,  transmitted 
to  the  Congress  of  the  United  States  a 
"report  on  endangered  and  threatened 
species  of  the  United  States."  This  report 
(cited  herein  as  "Ripley  1975")  lists  num- 
erous plants  from  Utah  as  either  threat- 
ened,   endangered,    possibly    extinct,    or 


probably  extinct.  Each  category  was  ab- 
breviated as  T,  E,  PoEx,  or  PrEx,  respec- 
tively. 

That  report  was  prepared  during  the 
calendar  year  of  1974  as  mandated  by 
the  Endangered  Species  Act  of  1973  (Pub- 
lic Law  93-205)  in  which  the  secretary  of 
the  Smithsonian  was  to  report  to  Congress 
within  one  year  on  all  of  the  "species  of 
plants  which  are  now  or  may  become  en- 
dangered or  threatened"  in  the  United 
States  (sec.  12).  The  Congress  provided 
no  funding  for  the  preparation  of  this  re- 
port, and  the  time  restrictions  prevented 
a  detailed  field  examination  of  all  spe- 
cies included  in  the  list.  Some  states, 
notably  California  and  Texas,  with  active 
offices  concerned  with  endangered  plant 
species,  were  able  to  provide  precise  data; 
other  states  were  able  to  supply  some  in- 
formation, and  still  others  had  little  or  no 
input  into  the  final  report,  except  that 
gathered  by  the  committee  established  by 
the  secretary  to  prepare  a  statement  for 
the  Congress.  The  data  for  the  state  of 
Utah  was  provided  by  a  number  of  indi- 
viduals (including  the  authors  of  this 
paper,  and  Reveal  served  on  the  Smith- 
sonian committee),  but  time  did  not  per- 
mit the  type  of  critical  examination  of 
each  taxon  which  has  been  largely  pos- 
sible for  the  present  paper.  The  Smithso- 
nian report  (Ripley  1975)  was  published 
in  the  Federal  Register  (Schreiner  1975) 
as  a  "notice  of  consideration,"  and  we 
understand  that  the  Department  of  In- 
terior will  submit  a  revised  listing  in  the 
near  future.  Disagreement  as  to  the  desig- 
nation of  degree  of  endangerment  be- 
tween this  paper  and  that  of  Ripley  (1975) 
represents  the  results  of  a  more  detailed 
and  concentrated  survey  of  the  Utah 
flora,  and  is  based  on  detailed  literature 
search  and  personal  information  of  the 
present  authors  and  that  of  their  col- 
leagues who  have  reviewed  the  manu- 
script. These  differences  have  largely  been 
resolved  and  will  appear  in  the  Smithso- 
nian's revised  list  to  be  submitted  early 
in  1976  to  the  Secretary  of  the  Interior. 
Much  of  the  information  has  been  ac- 
quired through  many  years  of  investi- 
gation of  the  Utah  flora,  both  in  the  field 
and  in  the  herbarium,  and  has  been  stim- 
ulated by  the  Ripley   (1975)   report. 

The  designation  of  the  status  of  an  in- 
dividual taxon  as  belonging  to  a  partic- 


330 


GREAT   BASIN   NATURALIST 


Vol.  35,  No.  4 


ular  category  is  subjective.  Still,  it  is 
based  on  the  best  information  available 
to  us  at  the  present  time  (a  requirement 
of  the  Endangered  Species  Act).  A  plant 
species  is  considered  as  "endangered" 
when  its  known  area  of  distribution  is 
very  small,  and  when  the  expected  de- 
velopment or  exploitation  of  the  area  oc- 
cupied has  already  occurred  or  is  immi- 
nent. A  "threatened"  plant  is  one  of  some- 
what larger  known  areal  extent,  but 
which  has  experienced  or  is  now  exper- 
iencing a  reduction  of  its  natural  distribu- 
tional area.  These  are  the  two  major  cat- 
egories required  by  law  to  be  designated. 
Plants  that  are  "rare  or  restricted"  are 
those  which  have  been  collected  only  oc- 
casionally or  which  are  known  from  very 
limited  regions  of  the  state;  they  may  or 
may  not  be  widely  distributed  elsewhere 
outside  the  political  confines  of  Utah.  In- 
troduced species  are  excluded  from  this 
category,  even  though  they  might  be  re- 
stricted or  even  rare. 

Plants  considered  to  be  "endemic"  are 
those  whose  entire  distributional  area  is 
within  Utah  or  within  one  of  the  natural 
drainage  basins  which  overlap  the  politi- 
cal boundaries  of  the  state.  Some  plants 
in  this  category  especially  are  poorly 
known  taxonomically  and  biologically, 
and  in  these  cases  we  have  so  indicated 
the  need  for  a  careful  systematic  evalu- 
ation of  the  taxon  to  determine  the  status 
of   the   plant   entity. 

All  statements  about  status  of  those 
plants  not  endemic  to  Utah  are  with  re- 
gard to  the  occurrence  of  those  plants 
within  Utah.  In  many,  if  not  most  of  these 
cases,  the  species  are  widespread  and  com- 
mon to  abundant  in  other  portions  of 
their  area  of  distribution.  If  they  are  limi- 
ted, rare,  or  possibly  extirpated  from 
Utah,  only  that  portion  of  their  range  is 
considered  in  making  the  designation. 
This  follows  the  guidelines  established  by 
California  and  Texas  in  which  the  en- 
dangered and  threatened  lists  are  based 
solely  upon  the  situation  of  the  plant  in 
question  within  the  confines  of  the  state 
boundaries   (Table  1). 

Plants  listed  in  one  of  the  categories 
designated  above  by  Ripley  (1975)  are 
included  here,  whether  or  not  they  are 
considered  as  something  other  than  threat- 
ened, endangered,  or  extinct  on  the  new 
list  that  will  be  published  in   1976.  Ob- 


vious errors,  which  will  not  be  repeated 
in  the  future,  such  as  Lewisia  maguirei 
and  Penstemon  decurvus,  to  mention  only 
two,  are  excluded. 

Voucher  specimens  for  some  of  the  re- 
ports are  cited  in  the  distribution  state- 
ments (e.g.,  Harrison  6370,  for  Cymop- 
terus  basalticus) .  Bibliographic  citations 
in  support  of  distributional  data  are  in- 
cluded for  many  species,  especially  for 
those  which  are  obscure  or  are  poorly  rep- 
resented in  herbaria.  We  are  not  making 
any  attempt  to  provide  precise  location 
data  (except  as  might  be  obtained  from 
published  type  localities)  in  fear  of  com- 
mercial exploitation  of  some  plants  (i.e., 
catci  and  orchids)  and  because  of  the  pos- 
sibility of  destruction  of  selected  popula- 
tions by  those  having  a  vested  interest  in 
ridding  parcels  of  land  of  any  species  of 
plant  that  might  fall  under  the  protection 
of  the  provisions  of  the  Endangered  Spe- 

Table  1.  Numerical  summary  of  the  endan- 
gered, threatened,  extinct,  extirpated  and  rare  or 
endemic  species  in  Utah. 


o 

T3 

t 

0) 

0) 

'S 

1 

60 

CO 

0) 

c 

1 

-o 

U 

u 

S 

JS 

c« 

County 

W 

H 

M 

w 

tf 

Beaver 

1 

7 

0 

1 

11 

Box    Elder 

1 

5 

0 

0 

1 

Cache 

2 

8 

0 

0 

3 

Carbon 

2 

6 

0 

0 

11 

Daggett 

3 

9 

0 

1 

4 

Davis 

0 

0 

0 

0 

4 

Duchesne 

3 

9 

1 

1 

14 

Emery 

4 

19 

1 

0 

28 

Garfield 

9 

39 

1 

1 

38 

Grand 

6 

17 

0 

1 

30 

Iron 

2 

15 

1 

0 

9 

Juab 

0 

7 

0 

3 

10 

Kane 

12 

31 

0 

4 

26 

Millard 

2 

9 

1 

0 

14 

Morgan 

0 

0 

0 

0 

0 

Piute 

0 

12 

0 

1 

8 

Rich 

1 

1 

0 

0 

1 

Salt  Lake 

0 

8 

0 

1 

8 

San  Juan 

7 

25 

0 

2 

27 

Sanpete 

1 

6 

1 

0 

9 

Sevier 

2 

13 

0 

0 

7 

Summit 

2 

3 

0 

3 

6 

Tooele 

0 

3 

0 

3 

7 

Uintah 

7 

17 

1 

1 

14 

Utah 

1 

7 

0 

1 

13 

Wasatch 

0 

1 

1 

0 

5 

Washington 

12 

41 

0 

3 

24 

Wayne 

5 

14 

0 

0 

29 

Weber 

0 

7 

0 

0 

4 

Dec.  1975 


WELSH,   ET   AL.:    ENDANGERED    UTAH    PLANTS 


331 


cies  Act  of  1973.  Nevertheless,  those  agen- 
cies charged  with  protection  of  endan- 
gered or  threatened  plants  must  deter- 
mine the  precise  localities  of  those  entities 
in  order  to  plan  for  their  survival. 

Maps  are  presented  (Figs.  1  and  2) 
which  demonstrate  the  unequal  distribu- 
tion of  these  unusual  plants.  In  Figure  1 
the  number  of  taxa  known  of  each  county 
is  indicated.  The  total  appears  to  be  very 
large,  but  this  is  due  to  many  of  the  spe- 
cies being  listed  in  several  counties  when 
area  of  distribution  occurs  outside  a  single 
county.  Phytogeographic  regions  of  Utah 
are  plotted  in  Figure  2,  and  the  number  of 
taxa  considered  to  be  unusual  is  cited  for 
each.  The  number  of  taxa  totals  are  less 
than  for  the  entire  state  because  some  dis- 
tributional data  are  so  indefinite  as  to  not 
be  plotable.  Also,  the  figures  represent  an 
attempt  to  plot  plants  within  their  main 
area  of  distribution;  double  representation 
has  been  avoided.  This  accounts  for  the  ap- 
parent discrepancies  between  the  two 
maps. 

Recommendations 

Land  use  planning  should  take  into  ac- 
count the  presence  of  the  unusual  plants 
present  in  the  state  of  Utah.  If  and  when 
the  Secretary  of  Interior  proclaims  any 
species  of  plants  found  in  Utah  as  en- 
dangered or  threatened,  they  will  fall  un- 
der the  protective  provisions  of  the  En- 
dangered Species  Act  which,  in  section 
seven,  calls  upon  all  federal  departments 
and  agencies  "to  insure  that  actions  auth- 
orized, funded,  or  carried  out  by  them  do 
not  jeopardize  the  continued  existence  of 
such  endangered  species  and  threatened 
species  or  result  in  the  destruction  or 
modification  of  habitat  of  such  species." 
Those  which  are  protected  by  law  must 
be  determined,  and  their  areas  of  distri- 
bution should  become  known  to  the  agen- 
cies responsible  for  their  protection. 
Questionable  taxonomic  units  should  be 
investigated  in  order  to  determine  their 
nature  and  area  of  distribution.  Plants 
thought  to  be  extinct  should  be  sought 
in  carefully  coordinated  field  studies.  The 
results  of  these  studies  should  be  reported 
to  the  Secretary-  of  the  Interior  so  that  in- 
formation about  the  various  species  may 
be  updated  from  time  to  time,  and  so  the 
status  of  each  taxon  can  be  reviewed. 
When  new  species  are  described  from 
Utah,   their   status    should   be   evaluated, 


and  if  they  should  prove  to  be  endangered 
or  threatened,  that  information  should  be 
presented  to  the  Secretary  so  that  such 
species  may  be  considered  for  the  federal 
endangered  species  list. 

In  a  recent  issue  of  the  Federal  Register, 
Green  wait  (1975)  proposed  a  "rule- 
making" for  various  amendments  to  the 
Endangered  Species  Act  of  1973.  These 
amendments  introduce  the  term  "plant" 
into  many  sections  of  the  act.  However, 
as  Lachenmeier  (1974)  has  pointed  out, 
there  are  several  legal  and  constitutional 
questions  about  the  act  that  need  to  be 
resolved  still,  and  from  a  botanical  point 
of  view,  there  still  exist  certain  discrim- 
inatory differences  between  "wildlife" 
and  "plants."  These  must  be  resolved  in 
the  future. 

In  and  for  the  state  of  Utah,  we  recom- 
mend that  a  review  board  be  appointed 
to  oversee  the  protection  of  Utah's  most 
unique  plants,  and  funding  should  be 
forthcoming  to  allow  a  real  understand- 
ing not  only  of  the  endangered  and  threat- 
ened species,  but  the  entire  state's  flora. 
The  state  should  adopt  the  federal  act  and 
include  within  its  own  listing  those  spe- 
cies of  the  state  which  are  endangered 
or  threatened  within  its  borders.  Policies 
should  be  adopted  that  will  determine 
basic  land  use  with  regard  to  these  unique 
plants,  and  "critical  habitats"  as  provided 
by  the  act  should  be  investigated  through- 
out the  state  (see  also  Green  wait  &  Geh- 
ringer  1975).  Above  all,  prompt  actions 
must  be  taken  by  the  state  to  preserve  and 
protect  the  state's  unique  flora,  and  the 
members  of  the  botanical  community 
must  be  ready  to  aid  and  assist  all  levels 
of  government  in  this  important  endeavor. 

Note 

In  the  following  list,  we  are  abbrevi- 
ating journals  according  to  the  abbrevi- 
ations listed  by  Lawrence  et  al.  (1968) 
and  herbaria  according  to  listing  given  by 
Holmgren  and  Keuken  (1974).  We  have 
attempted  to  give  the  type  information  as 
published  in  the  original  place  of  publica- 
tion; however,  we  are  not  attempting  to 
typify  any  name,  nor  is  the  herbarium 
cited  considered  a  statement  of  lectotyp- 
ification.  This  information  is  given  for  the 
convenience  of  the  reader.  For  the  most 
part,  the  abbreviations  of  the  authors  fol- 


332 


GREAT   BASIN    NATURALIST 


Vol.  35,  No.  4 


low   the   unpublished   suggestions   of   the 
Index  Kewensis  staff. 


Acknowledgments 

The  authors  wish  to  acknowledge  the 
critical  reviews  supplied  by  our  colleagues 
in  the  study  of  Utah's  plant  taxonomy. 
Especially  important  help  and  criticism 
was  supplied  b}'  Arthur  H.  Holmgren, 
Lois  Arnow,  and  Andrew  H.  Barnum. 


Apiaceae 

Angelica  wheeleri  S.  Wats.,  Amer.  Nat- 
uralist 7:  301.  1873. 

Type:   Utah,  Wheeler  s.n.    (gh). 

Status:  Endemic,  evidently  rare  and 
local,  possibly  threatened. 

Distribution:  Northern  and  central 
Utah   (Mathias  &  Constance  1945). 

Cymopterus  basalticus  M.  E.  Jones,  Contr. 
W.  Bot.   12:    6.   1908. 

Type:  Half-way  Station,  west  of  Wa 
Wa,  Millard  or  Beaver  Co.,  Utah,  7.000 
feet,  15  May  1906,  M.  E.  Jones  s.  n. 
(pom). 

Status:  Restricted  and  locally  abun- 
dant, neither  threatened  nor  endan- 
gered (Ripley,  T) . 

Distribution:  Western  Utah  (Mil- 
lard Co.,  Harrison  6370;  Matthews  14 
[bry]),  and  adjacent  Nevada  (Math- 
ias &  Constance  1945). 

Cymopterus  coulteri  (M.  E.  Jones)  Math- 
ias, Ann.  Missouri  Bot.  Gard.  17:  276. 
1930.  based  on  C.  corrugatus  var.  coul- 
teri M.  E.  Jones,  Contr.  W.  Bot.  12:19. 
1908. 

Type:  Juab,  Juab  Co.,  Utah,  4,000 
feet,  30  Apr  1880.  M.  E.  Jones  1691 
(us). 

Status:  Endemic,  rare  and  restric- 
ted, threatened   (Ripley,  T). 

Distribution:  Western  Utah  (San- 
pete Co.,  Mabey  5300  [bry]). 

Cymopterus  duchesnensis  M.  E.  Jones. 
Contr.  W.  Bot.  13:   12.  1910. 

Type:  Among  loose  rocks  on  south- 
ern slopes  of  mesas,  Myton,  Duchesne 
Co.,  Utah,  5,000  feet,  20  May  1908, 
M.  E..  Jones  s.n.  (pom). 

St.\ti:s:  Endemic,  rare  and  restric- 
ted, threatened    (Ripley,  E). 

Distribution:  Duchesne  and  Uintah 
counties,    Utah     {Welsh    180    [bry]): 


Holmgren  &  Reveal  1887    [ny,  utc]; 
Holmgren    &    Holmgren    5169     [bry, 

NY,    utc]). 

Cymopterus  jonesii  Coult.  &  Rose,  Rev. 
N.  Amer.  Umbell.  80.  1888. 

Type:  Frisco,  Beaver  Co.,  Utah, 
8,000  feet,  22  Jun  1880,  M.  E.  Jones 
1808   (US). 

Status:  Rare  and  restricted. 

Distribution:  Southwestern  Utah 
and  Nevada  (Mathias  &  Constance 
1945). 

Cymopterus  higginsii  Welsh,  Great  Basin 
Nat.  35:  377.  1976. 

Type:  Shadscale  dominated  bajada, 
on  gravelly  pedimental  fan  east  of 
None  Butte,  ca  17  miles  east  of  Glen 
Canyon  City,  Kane  Co.,  Utah,  31  May 
1975,  5.  L.^ Welsh  12740  (bry). 

Status:  Endemic,  local;  possibly 
threatened. 

Distribution:  Eastern  Kane  Co., 
Utah. 

Cymopterus  minimus  (Mathias)  Mathias, 
Brittonia  2:  245.  1936,  based  on  Aulo- 
spermum  minimum  Mathias,  Ann.  Mis- 
souri Bot.  Gard.  17:  353.  1930. 

Type:  On  the  upper  part  of  the 
"Breaks"  at  Cedar  Breaks,  Iron  Co., 
Utah,  ca  10,500  feet,  Mathias  723 
(mo). 

Status:  Endemic,  rare  and  restric- 
ted to  Cedar  Breaks,  possibly  endan- 
gered  (Ripley,  E). 

Distribution:  Cedar  Breaks,  Iron 
Co.,  Utah. 

Cymopterus  newberryi  (S.  Wats.)  M.  E. 
Jones,  Zoe  4:  47.  1893,  based  on  Peu- 
cedanum  newberryi  S.  Wats.,  Proc. 
Amer.  Acad.  Arts  11:    145.  1876. 

Status:  Widespread  and  at  least  lo- 
cally abundant,  neither  threatened  nor 
endangered   (Ripley,  T). 

Distribution:  Garfield,  Grand, 
Kane,  Millard,  San  Juan,  Uintah, 
Washington,  and  Wayne  counties,  Utah 
(bry,  utc),  and  northern  Arizona 
(Mathias  &  Constance  1945). 

Cymopterus  rosei  M.  E.  Jones,  Contr.  W. 
Bot.  12:   17.  1908. 

Type:  Richfield,  Sevier  Co.,  Utah, 
18  Jun  1898,  M.  E.  Jones  30  (us). 

Status:  Endemic,  rare  and  local,  pos- 
sibly  threatened    (Ripley,   T) . 


Dec.  1975 


WELSH,   ET   AL.:    ENDANGERED    UTAH    PLANTS 


333 


Distribution:  Iron,  Sanpete,  Sevier, 
and  Washington  counties,  Utah  {Ma- 
guirc  1913U  Holmgren  10932  [utc]). 

Ligusticum  porteri  Couh.  &  Rose  var. 
brevilobum  (Rydb.)  Mathias  &  Con- 
stance, Bull.  Torrey  Bot.  Club  68:  123. 
1941,  based  on  L.  brevilobum  Rvdb.. 
Fl.  Rocky  Mts.  613,   1064.   1917.  " 

Type:  Aquarius  Plateau,  Utah  :5  Aug 
1905,  Rrdberg  &  Calrton  7473  (nyj. 

Status:  Endemic,  rare  and  local; 
possibly  threatened. 

Distribution:  Aquarius  Plateau, 
Garfield  Co.,  Utah  (Mathias  &  Con- 
stance 1941). 

Lomatium  latilobum  (Rydb. )  Mathias, 
Ann.  Missouri  Bot.  Card.  25:  281.  1937, 
based  on  Cynomarathriim  latilobum 
Rydb.,  Bull.  ToiTey  Bot.  Club  40:  73. 
1913. 

Type:  Proposed  dam  site,  near  Wil- 
son Mesa,  Grand  Co.,  Utah,  Rydberg 
&  Garrett  8371   (ny). 

Status:  Endemic,  rare  and  local, 
possibly  threatened. 

Distribution:  Grand  Co.,  Utah 
(bry,-  utc). 

Lomatium  megarrhizum  (A.  Nels.) 
Mathias,  Ann.  Missouri  Bot.  Gard.  25: 
282.  1937,  based  on  Peucedanum  megar- 
rhiza  A.  Nels.,  Bull.  Torrev  Bot.  Club 
26:  130.  1899. 

Status:  Rare  and  local,  possibly 
threatened. 

Distribution:  Emery  Co.,  Utah 
{Higgins  1268  [bry])  and  southwest- 
ern Wyoming  (Mathias  &  Constance 
1945;  Higgins  1972b). 

Lomatium  minimum  Mathias,  Ann.  Mis- 
souri Bot.  Gard.  25:  273.  1937. 

Type:  Near  the  hotel,  on  dr}'  slopes 
bordering  Bryce  Canyon,  Garfield  Co.. 
Utah,  8,600  feet,  Mathias  670  (mo). 

Status:  Endemic,  local,  possibly 
threatened    (Ripley,  T) . 

Distribution:  Garfield,  Iron,  and 
Kane  counties,  Utah  (bry,  utc). 

Lomatium  scabrum  (Coult.  &  Rose)  Ma- 
thias, Ann.  Missouri  Bot.  Gard.  25:  275. 
1937,  based  on  Cynomarathrum  scab- 
rum  Coult.  &  Rose,  Contr.  U.S.  Natl. 
Herb.  7:  247.  1900. 

Type:  Frisco,  Beaver  Co.,  Utah,  2,400 
meters,  Jones  1864  (us). 


Status:  Endemic  (but  possibly  in 
Mohave  Co.,  Arizona),  locally  common, 
not  threatened  nor  endangered. 

Distribution:  Beaver,  Iron  (ut), 
Millard,  and  Washington  counties.  Utah 
(bry)  . 

Musineon  lineare  (Rydb.j  Mathias,  Ann. 
Missouri  Bot.  Gard.  17:  265.  1930, 
based  on  Daucophyllum  lineare  Rydb., 
Bull.  Torrey  Bot.  Club  40:   69.  1913. 

Type:  Near  Logan,  Cache  Co.,  Utah, 
9  Aug  1895,  Rydberg  s.n.   (ny). 

Status:  Endemic,  possibly  threat- 
ened. 

Distribution:  Cache  Co.,  Utah 
(Holmgren  3595  [utc]). 

Apo(;yn.\ceae 

Cycladenia  humilis  Benth.  var.  jonesii 
(Eastw.)  W^elsh  &  Atwood,  stat.  et 
comb,  nov.,  based  on  C.  jonesii  Eastw., 
Leafl.  W.  Bot.  3:  159.  1942. 

Type:  San  Rafael  Swell,  Emery  Co., 
Utah.   19  May  1914,  M.  E.  Jones  s.n. 

(C.'^S). 

Status:  Endemic,  rare  and  endan- 
gered  (Ripley,  E). 

Distribution:  Emery  and  Grand 
counties,  Utah. 

This  entity  is  known  in  Utah  from 
three  basic  collections:  the  type  as  cited 
above,  a  collection  by  Cottam  (ut)  also 
from  the  San  Rafael  Swell,  and  a  collec- 
tion from  Castle  Valley.  Grand  Co.,  Utah 
(Welsh  1970b). 

Araliaceae 

Aralia  racemosa  L.  ssp.  bicrenata  (Woo- 
ton  &  Standley)  Welsh  &  Atwood, 
Stat.  &  comb,  no^ .,  based  on  A.  bicre- 
nata Wooton  &  Standley,  Contr.  U.S. 
Natl.  Herb.   16:    157.   1913. 

Status:  Rare;  restricted  to  Zion 
Narrows,   Washington   Co.,   Utah. 

Distribution:  Washington  Co..  Utah 
{Welsh  12366  [bry]  ),  and  from  south- 
eastern Canada  south  to  Georgia,  west- 
ward to  Arizona  and  northern  Mexico 
(Kearney  &  Peebles  1951);  the  sub- 
species is  from  Utah  and  Arizona. 

ASCLEPIADACEAE 

Asclepias  cutleri  Woodson,  Ann.  Missouri 
Bot.  Gard.  26:  263.  1939. 
Status:  Rare,  threatened. 


334 


GREAT   BASIN    NATURALIST 


Vol.  35,  No.  4 


Distribution:  Grand  {Cottam  5799 
[bry]  and  San  Juan  {Holmgren  3144 
[uTc])  counties,  Utah,  and  from  north- 
eastern Arizona. 

Asclepias  engelmanniana  Woodson,  Ann. 
Missouri  Bot.  Card.  28:  207.  1941. 

Status:  Rare. 

Distribution:  Grand  (Rydberg  & 
Garrett  8504  [ut])  and  San  Juan 
(Welsh  et  al.  2930  [bry])  counties, 
Utah;  from  Nebraska  to  Texas  and 
Arizona. 

Asclepias  labriformis  M.  E.  Jones,  Proc. 
Calif.  Acad.  Sci.  II,  5:   708.  1895. 

Type:  Capital  Wash,  near  the  Henry 
Mts.,  Wayne  Co.,  Utah,  5,000  feet,  in 
sandy  gulch,  19  Jul  1894,  M.  E.  Jones 
5650  (pom). 

Status:  Endemic  to  the  state  but 
common,  neither  threatened  nor  en- 
dangered. 

Distribution:  Emery,  Garfield,  San 
Juan,  Uintah,  and  Wayne  counties, 
Utah  (bry). 

Asclepias  ruthiae  Maguire  &  Woodson, 
Ann.  Missouri  Bot.  Gard.  28:  245.  1941. 

Type:  Frequent,  sandy  soil,  vicinity 
of  water  tanks,  Calf  Spring  Canyon, 
5,000  feet,  San  Rafael  Swell,  18  miles 
southeast  of  Castle  Dale.  Emery  Co., 
Utah,  10  May  1940,  Maguire  &  Ma- 
guire 18310  (uTc). 

Status:  Endemic,  common  in  dis- 
junct populations  on  the  San  Rafael 
Swell;  possibly  threatened  (Ripley,  T). 

Distribution:  Emery  (Cottam  5500, 
Harrison  8067,  Higgins  &  Reveal 
1285a  [bry],  Grand  [see  Woodson 
1954),  and  Wayne  (Harrison  11223 
and  11891  [bry])  counties,  Utah. 

Asteraceae 

Chamaechaenactis  scaposa  (Eastw.) 
Rydb.,  Bull.  Torrey  Bot.  Club  33:  156. 
1906,  based  on  Chaenactis  scaposa 
Eastw.,  Zoe  2:  231.  1891. 

Status:  Restricted,  rare  except  lo- 
cally, not  threatened  nor  endangered. 

Distribution:  Carbon,  Duchesne, 
Emery,  San  Juan,  Uintah,  and  Wayne 
counties,  Utah,  and  western  Colorado 
(bry.  utc). 

Cirsium  rydbergii  Petrak,  Beih.  Bot. 
Centralbl.  35  (2):  315.  1917. 


Type:  Along  the  San  Juan  River 
near  Bluff,  San  Juan  Co.,  Utah,  1,200- 
1,500  meters,  1-2  Jul  1911,  Rydberg 
10001  (g?). 

Status:  Habitat  specific,  in  hanging 
gardens,  restricted  and  possibly  threat- 
ened. 

Distribution:  Garfield,  Grand, 
Kane,  San  Juan,  and  Wayne  counties, 
Utah  (bry),  and  Black  Mesa,  Apache 
Co.,  Arizona  (Howell  &  McClintock 
1960). 

Enceliopsis  nutans  (Eastw.)  A.  Nels.,  Bot. 
Gaz.  47:  433.  1909,  based  on  Encelia 
nutans  Eastw.,  Zoe  2:  230.  1891. 

Status:  Habitat  specific,  in  heavy 
clay  soils,  restricted  and  rare. 

Distribution:  Carbon,  Duchesne, 
Emery,  Grand,  Uintah,  and  Wayne 
counties,  Utah  (bry,  utc),  and  western 
Colorado  (Harrington  1954). 

Erigeron  abajoensis  Cronq.,  Brittonia  6: 
168.  1947. 

Type:  Abajo  Mts.  (eastern  range), 
3,000-5,300  meters,  San  Juan  Co.,  Utah, 
17  Aug  1911,  Rydberg  &  Garrett  9755 
(ny). 

Status:  Endemic,  rare  and  threat- 
ened (Ripley,  T). 

Distribution:  Garfield,  Kane,  and 
San  Juan  counties,  Utah  (bry). 

Erigeron  arenarioides  (D.  C.  Eaton) 
Rydb.,  Fl.  Rocky  Mts.  1067.  1917,  based 
on  Aster  arenarioides  D.  C.  Eaton  ex  A. 
Gray,  Proc.  Amer.  Acad.  Arts  8:  647. 
1873. 

Type:  Rocky  gulch  above  Cottonwood 
Canyon,  Wasatch  Mountains,  Salt  Lake 
Co.,' Utah,  8,000-9,000  feet,  Aug  1869, 
Waston  547  (us). 

Status:  Endemic  and  rare,  restric- 
ted to  rock  crevices,  threatened. 

Distribution:  Salt  Lake,  Tooele, 
Utah,  and  Weber  counties,  Utah  (bry, 
UTC;  Cronquist  1947). 

Erigeron  cronquistii  Maguire,  Brittonia 
5:   201.   1944. 

Type:  Cliffs,  north  side  of  Logan 
Canyon  l^  mile  below  forks.  Bear  River 
Range,  Cache  Co.,  Utah,  5,800  feet,  20 
May  1939,  Maguire  16681   (ny). 

Status:  Endemic,  rare  and  threat- 
ened  (Ripley,  T). 

Distribi'tion:  Cache  Co..  Utah 
(Cronquist  1947). 


Dec.  1975 


WELSH,   ET  AL.:    ENDANCJERED    UTAH    PLANTS 


335 


Erigeron  garrettii  A.  Nels.,  Manual  Bot. 
Centr.  Rocky  Mts.  526.  1909. 

Type:  Big  Cottonwood  Canyon,  Salt 
Lake  Co.,  Utah.  28  Jun  1905,  Garrett 
1310  (rm). 

Status:  Endemic,  rare  and  possibly 
threatened  (Ripley,  T). 

Distribution:  High  altitudes  in  Box 
Elder  (ut).  Salt  Lake  and  Utah  coun- 
ties, Utah  (bry;  Cronquist  1947). 

Erigeron  flagellar  is  A.  Gray  var.  triloba- 
tus  Maguire  ex  Cronq.,  Brittonia  6:  258. 
1947. 

Type:  Canyon  to  Cedar  Breaks,  12 
miles  east  of  Cedar  City,  Iron  Co.,  Utah, 
5  Aug  1934,  Maguire  14947   (ny). 

Status:  Endemic  and  rare,  endanger- 
ed (Ripley,  E). 

Distribution:  Iron  Co.,  Utah; 
known  only  from  the  type  locality 
(Cronquist  1947). 

Erigeron  kachinensis  Welsh  &  Moore, 
Proc.  Utah  Acad.  Sci.  45:  231.  1968. 

Type:  Hanging  gardens  and  seeps, 
near  Kachina  Natural  Bridge,  Natural 
Bridges  N.M.,  San  Juan  Co.,  Utah,  13 
Aug  1963,  Welsh  &  Moore  2398  (bry). 

Status:  Endemic,  rare  and  endan- 
gered   (Ripley,  E). 

Distribution:  San  Juan  Co.,  Utah; 
known  only  from  the  type  locality 
(Welsh  &  Moore  1968). 

Erigeron  maguirei  Cronq.,  Brittonia  6: 
165.  1947. 

Type:  Dry  rocky  sandy  canyon  bot- 
tom. Calf  Spring  Wash,  1.5  mile  up 
San  Rafael  Swell,  16  Jun  1940,  5,500 
feet,  Maguire  18459  (ny). 

Status:  Endemic,  rare  and  possibly 
extinct  (Ripley,  E). 

Distribution:  Emery  Co.,  Utah; 
known  only  from  the  type  locality. 

Erigeron  mancus  Rydb.,  Fl.  Rocky  Mts. 
902,  1067.  1917.  ' 

Type:  LaSal  Mts.,  Grand  or  San 
Juan  Co.,  Utah,  7  Jul  1911,  Rydberg  & 
Garrett  8671   (ny). 

Status:  Endemic,  rare  and  possibly 
threatened  (Ripley,  T) . 

Distribution:  LaSal  Mts.,  Grand  and 
San  Juan  counties,  Utah  (Cronquist 
1947.) 

Erigeron  religiosus  Cronq.,  Brittonia  6: 
258.    1947. 


Type:  Clear  Creek,  Zion  N.P.,  Wash- 
ington Co.,  Utah,  8  Sep  1938,  Eastwood 
8z  Howell  6339  (cas). 

Status:  Endemic,  rare  and  endan- 
gered (Ripley,  E). 

Distribution:  Kane  [Harrison 
11085  [bry]  and  Washington  counties, 
Utah   (Cronquist  1947). 

Erigeron  sionis  Cronq.,  Brittonia  6:  258. 
1947. 

Type:  Zion  N.P.,  Washington  Co., 
Utah,  1-3  Aug  1925,  Pilsbry  s.n.   (ph). 

Status:  Endemic,  very  rare  and  en- 
dangered (Ripley,  E). 

Distribution:  Washington  Co., 
Utah;  known  only  from  type  locality. 

Flaveria  campestris  J.  R.  Johnston,  Proc. 
Amer.  Acad.  Arts  29:  287.  1903. 

Status:  Rare,  restricted,  but  not  ap- 
parently threatened  or  endangered. 

Distribution:  Grand  Co.,  Utah  (Ma- 
guire 1937;  Harrison  et  al.  1964);  Colo- 
rado, New  Mexico  and  northern  Mex- 
ico. 

Gaillardia  flava  Rydb.,  N.  Amer.  Fl.  34: 
139.  1915. 

Type:  Lower  Crossing  of  the  Price 
River  (see  Jones  1965),  Emery  Co., 
Utah,  2  Jul  1898,  M.  E.  Jones  6412 
(us). 

Status:  Endemic,  rare  and  endan- 
gered. 

Distribution:  Emery  Co.,  Utah 
(Cottam  &  Hutchings  2176   [bry]). 

Gaillardia  spathulata  A.  Gray,  Proc. 
Amer.  Acad.  Arts   12:    59.   1876. 

Type:  Rabbit  Valley,  Wayne  Co., 
Utah,  7,000  feet,  1875,  Ward  s.n.  (gh). 

Status:  Endemic,  common  through- 
out its  range,  neither  threatened  nor  en- 
dangered. 

Distribution:  Carbon,  Emery,  Gar- 
field, Grand,  and  Wayne  counties,  Utah 
(bry,  utc). 

Haplopappus  scopulorum  (M.  E.  Jones) 
Blake  in  Tidestrom,  Contr.  U.S.  Natl. 
Herb.  25:  546.  1925,  based  on  Bigelovia 
menziesii  var.  scopulorum  M.  E.  Jones, 
Proc.  Calif.  Acad.  Sci.  II,  5:  692.  1895. 

Type:  Near  Cedar  City,  Iron  Co., 
Utah,  6,000  feet,  10  xMay'l894,  M.  E. 
Jones  5204v  is  the  first  of  two  collec- 
tions cited  (pom). 

Status:  Uncommon,  habitat  specific, 


336 


GREAT   BASIN   NATURALIST 


Vol.  35,  No.  4 


neither     endangered     nor     threatened 
(Ripley,  T). 

Distribution:  Iron,  Kane  (?),  and 
San  Juan  (Welsh  &  Moore  2447;  Welsh 
8813;  Atwood  4100  [bry])  counties, 
Utah,  and  Coconino  Co.,  Arizona  (Mc- 
Dougall  1973). 

Helianthus  anomalus  Blake,  J.  Wash. 
Acad.  Sci.  21:  333.  1931. 

Type:  Desert  between  Hanksville  and 
Henry  Mts.,  Wayne  Co.,  Utah,  5  Jul 
\9^0^,  Stanton  4806    (us). 

Status:  Restricted  to  dunes  and  sandy 
washes,  where  locally  abundant,  neither 
threatened  nor  endangered. 

Distribution:  Emery,  Garfield, 
Grand,  Juab,  Kane,  Millard,  San  Juan, 
Tooele,  Uintah,  Washington,  and 
Wayne  counties,  Utah,  and  northern 
Arizona  (Blauer  1966). 

Helianthus  canus  (Britton)  Wooton  & 
Standley,  Contr.  U.S.  Natl.  Herb.  6: 
190.  1913,  based  on  Helianthus  petiolar- 
is  var.  canus  Britton,  Mem.  Torrey  Bot. 
Club  5:  334.  1894. 

Status:  Rare,  possibly  threatened. 

Distribution:  San  Juan  Co.,  Utah 
(Cottam  2544'  [bry]  ),  and  from  western 
Texas  to  southern  California  and  north- 
ern Mexico. 

Helianthus  deserticolus  Heiser,  Proc.  In- 
diana Acad.  Sci.  70:  209.  1961. 

Type:  3.3  miles  west  of  Hurricane, 
Washington  Co.,  Utah,  29  Jun  1957, 
Stoutamire  2574    (ind). 

Status:  Rare,  restricted  and  possibly 
threatened. 

Distribution:  Washington  Co.,  Utah, 
Mohave  Co.,  Arizona,  and  Clark  Co., 
Nevada  (Heiser  1961). 

Heterotheca  jonesii  (Blake)  Welsh  &  At- 
wood, comb.  no\'.,  based  on  Chrysopsis 
jonesii  Blake  in  Tidestrom,  Contr.  U.S. 
Natl.  Herb.  25:  536.  1925,  a  substitute 
name  for  C.  caespitosa  M.  E.  Jones, 
Proc.  Calif.  Acad.  Sci.  II,  5:  694.  1895, 
not  Nutt. 

Type:  Springdale,  Washington  Co., 
Utah,  4,000  feet,  16  May  1894,  M.  E. 
Jones  5249u  (pom). 

Status:  Endemic,  rare,  local,  and  en- 
dangered. 

Distribution:  Washington  and  (jar- 
field  {Harrison  12345  [bry])  counties, 
Utah. 


Heterotheca  grandi  flora  Nutt.,  Trans. 
Amer.  Philos.  Soc.  II,  7:  315.  1840. 

Status:   Rare,  local  and  restricted. 

Distribution:  Washington  Co.,  Utah 
{Welsh  et  al.  9530  [bry]);  also  in  Cali- 
fornia and  Arizona. 

Heterotheca  subaxillaris  Britton  &  Rusby, 
Trans.  New  York  Acad.  Sci.  7:  10.  1887. 

Status:   Rare,  possibly  endangered. 

Distribution:  Grand  Co.,  Utah 
{Welsh  &  Moore  2744  [bry]);  wide- 
spread elsewhere  in  the  United  States. 

Hulsea  heterochroma  A.  Gray,  Proc. 
Amer.  Acad.  Arts  7:   359.  1868. 

Status:  Local,  rare  and  possibly 
threatened. 

Distribution:  Washington  Co.,  Utah 
[Higgins  &  Atwood  1410  [bry];  Hig- 
gins  1972b);  California  and  southern 
Nevada. 

Hymenopappus  filifolius  Hook.  var.  al- 
pestris  (Maguire)  Shinners,  Rhodora 
61:  155.  1959,  based  on  H.  nudipes  var. 
alpestris  Maguire,  Amer.  Midi.  Nat- 
uralist 37:  144.  1947  (includes  var. 
nudipes  (Maguire)  Turner). 

Type:  Cedar  Breaks  rim.  Iron  Co., 
Utah,  23  Jun  1940,  Maguire  19023 
(ny). 

Status:  Restricted  but  locally  com- 
mon,   not   threatened    nor   endangered. 

Distribution:  Duchesne,  Garfield, 
Iron,  Kane,  Piute,  Sevier,  Summit,  and 
Utah  counties,  Utah,  and  Lincoln  Co., 
Wyoming  (Turner  1956). 

Hymenopappus  filifolius  Hook.  var.  to- 
mentosus  (Rydb.)  Turner,  Rhodora  58: 
237.  1956,  based  on  H.  tomentosus 
Rydb.,  Bull.  Torrey  Bot.  Club  27:  633. 
1900. 

Type:  St.  George,  Washington  Co., 
Utah,  1877,  Palmer  270   (ny). 

Status:  Endemic,  rare  and  restricted, 
threatened  (Ripley,  T). 

Distribution:  Kane  and  Washington 
counties,  Utah  (Turner  1956). 

Hymcnoxys  depressa  (Torr.  &  Gray  ex 
Gray)  Welsh  &  Reveal,  comb,  nov., 
based  on  Actinclla  depressa  Torr.  & 
Gray  ex  Gray,  Mem.  Amer.  Acad.  Arts 
II,  4:   100.  1849. 

Status:  Rare  and  .  local,  possibly 
threatened. 

Distribution:     Emery     (bry)     and 


Dec.  1975 


WELSH,   ET   AL.:    ENDANGERED    UTAH    PLANTS 


337 


Garfield  (ut)  counties,  Utah,  and  from 
Colorado. 

Lepidosparluni  latisquamum  S.  Wats., 
Proc.  Amer.  Acad.  Arts  25:  133.  1890. 

Status:  Restricted  and  rare,  possibly 
threatened. 

Distribution:  Millard  Co.,  Utah 
(bry),  and  adjacent  Nevada  west  to 
eastern    California. 

Lygodesmia  grandiflora  (Nutt.)  Torr.  & 
Gray  var.  striata  Maguire,  Amer.  Midi. 
Naturalist  37:  145.  1947. 

Type:  Frequent  on  adobe  clay,  1  mile 
south  of  Price,  Carbon  Co.,  Utah,  5  Jun 
1940,  Maguire  18417   (ny). 

Status:  Endemic,  rare,  restricted 
and  endangered. 

Distribution:  Carbon  Co.,  Utah; 
known  only  from  the  type  locality. 

Machaer  anther  a  glabriuscula  (Nutt.) 
Cronq.  &  Keck  var.  confertifolia  Cronq., 
Leafl.  W.  Bot.  10:   11.  1963. 

Type:  Barren  clay  slopes  in  pinyon- 
juniper  zone,  11  miles  northeast  of 
Henrieville,  Garfield  Co.,  Utah,  7,000 
feet,  31  May  1961,  Cronquist  9164 
(ny). 

Status:  Endemic,  rare  and  threat- 
ened. 

Distribution:  Garfield  and  Kane 
counties,  Utah  (bry,  utc)  . 

Machaeranthera  grindelioides  (Nutt.) 
Shinners  var.  depressa  (Maguire) 
Cronq.  &  Keck,  Brittonia  9:  237.  1957, 
based  on  Haplopappus  nuttallii  var. 
depressa  Maguire,  Amer.  Midi.  Natural- 
ist 37:   144.  1947. 

Type:  Warm  Point,  5  miles  southwest 
of  Desert  Range  Experiment  Station 
headquarters,  Millard  Co.,  Utah,  10 
Jun   1941,  Maguire  20859   (ny). 

Status:  Rare  and  restricted,  neither 
threatened  nor  endangered  (Ripley,  T) . 

Distribution:  Beaver  and  Millard 
counties,  Utah  (bry),  and  Eureka,  Lin- 
coln and  White  Pine  counties,  Nevada 
(Maguire  1947). 

Machaeranthera  kingii  (D.  C.  Eaton) 
Cronq.  &  Keck,  Brittonia  9:  238.  1957, 
based  on  Aster  kingii  D.  C.  Eaton  in 
King,  Rep.  Geol.  Explor.  40th  Parallel 
5:    141.  1871. 

Type:    Wasatch  Mts.,  above  Cotton- 


wood Canyon,  Salt  Lake  Co.,  Utah, 
1869,  9,000  feet,  Watson  507   (gh). 

Status:  Endemic,  rare  and  restricted, 
possibh'  threatened. 

Distribution:  Cache,  Salt  Lake  and 
Utah  counties,  Utah   (bry,  utc). 

Malacothrix  clcvelandii  A.  Gray.  Bot. 
Calif.   1:   433.  1876. 

Status:    Local  and  infrequent. 
Distribution:  Washington  Co.,  Utah 
[Ativood   4966    [bry,    wts]);    also    in 
Nevada  and  California. 

Partheniuni  ligulatum  (M.  E.  Jones) 
Barneby,  Leafl.  W.  Bot.  5:  20.  1947, 
based  on  P.  alpinum  var.  ligulatum  M. 
E.  Jones,  Contr.  W.  Bot.  13:    16.  1910. 

Type:  On  nearly  bare  clayey  and 
gravelly  knolls  on  ridges,  Theodore 
[now  Duchesne],  Duchesne  Co.,  Utah, 
6,000  feet,  M.  E.  Jones  s.n.    (pom). 

Status:  Endemic,  rare  and  restricted, 
threatened    (Ripley,   T). 

Distribution:  Duchesne  and  Emery 
counties,  Utah   (bry,  ny). 

Senecio  dimorphophyllus  Greene  var. 
intermedins  Barkley,  Trans.  Kansas 
Acad.   Sci.  65:    363."  1962. 

Type:  Edge  of  swampy  places.  Geys- 
er Pass,  LaSal  Mts.  Utah,  10,500  feet. 
Pay  son  &  Pay  son  4097   (mo). 

Status:  Endemic,  rare,  restricted 
and  threatened   (Ripley,  T) . 

Distribution:  San  Juan  Co.,  Utah; 
known  only  from  the  type  area. 

Sphaeromeria  capitata  Nutt.,  Trans. 
Amer.  Philos.  Soc.  II,  7:  402.   1841. 

Status:    Local  and  rare,  threatened. 

Distribution:  Grand  Co.,  Utah 
(utc);  Wyoming  and  southern  Mon- 
tana. 

Townsendia  aprica  Welsh  &  Reveal,  Brit- 
tonia 20:  375.  1968. 

Type:  Ca  6  miles  south  of  Fremont 
Junction  along  Utah  Highway  72,  on 
low,  rolling  exposed  gray  clay  slopes, 
among  scattered  igneous  boulders,  Se- 
vier Co.,  Utah,  ca  6,500  feet,  1  May 
1966,  Reveal  &  Welsh  721    (bry). 

Status:  Endemic,  rare  and  restricted, 
endangered  (Ripley,  E) . 

Distribution:  Sevier  Co.,  Utah  (bry; 
Welsh  &  Reveal  1968). 

Townsendia  mensana  M.  E.  Jones,  Contr. 
W.  Bot.  13:   15.  1910. 


338 


GREAT  BASIN   NATURALIST 


Vol.  35,  No.  4 


Type:  Benches  of  the  Uinta  Mts., 
near  Theodore  [now  Duchesne],  Du- 
chesne Co.,  Utah,  14  May  1908,  M.  E. 
Jones  s.n.   (pom). 

Status:  Endemic,  locally  restricted, 
possibl}'  threatened. 

Distribution:  Duchesne  and  Uintah 
counties,  Utah  (Reveal  1970b). 

Townsendia  minima  Eastw.,  Leafl.  W. 
Bot.  1:  206.  1936. 

Type:  Bryce  Canyon,  Garfield  Co., 
Utah.  19  Jun  1933,  Eastwood  &  Howell 
727   (cAs). 

Status:  Endemic,  edaphically  restric- 
ted, possibly  threatened. 

Distribution:  Garfield  and  Kane 
counties.  Utah  (Beaman  1957;  Reveal 
1970b). 

Viguiera  soliceps  Barneby.  Leafl.  W.  Bot. 
10:   316.  1966. 

Type:  Locally  plentiful  on  gumbo- 
clay  knolls  and  bluffs,  lower  Cotton- 
wood Canyon  near  its  confluence  with 
Paria  River,  about  41  miles  southeast 
of  Cannon ville,  Kane  Co.,  Utah,  4,500 
feet,  12  Jun  1966,  Barneby  14435  (ny). 

Status:  Endemic,  local  on  Tropic 
Shale  formation,  threatened  (Ripley. 
E). 

Distribution:  Cottonwood  Canyon 
east  to  Last  Chance  Canyon,  Kane  Co., 
Utah  (bry,  utc). 

Xanthocephalum  sarothrae  (Pursh)  Shin- 
ners  var.  pomariense  (Welsh)  Welsh, 
comb,  nov.,  based  on  Gutierrezia  saro- 
thrae (Pursh)  Britton  &  Rusby  var. 
pomariensis  Welsh.  Great  Basin  Nat. 
30:  19.  1970. 

Type:  Sand}'  flat,  base  of  sandstone 
cliffs,  Frontier  formation,  mouth  of 
Orchard  Creek  Draw,  Dinosaur  N.M.. 
Uintah  Co.,  Utah,  2  Oct  1969,  Welsh 
et  al.  9471   (bry). 

Status:  Endemic,  rare  and  threat- 
ened. 

Distribution:  Uintah  Co.,  Utah 
(Welsh  1970b). 

Berberidaceae 

Berberis  fendleri  A.  Gray,  Mem.  Amer. 
Acad.  Arts  II,  4:  5.  1849. 

Status:  Rare,  floristically  restricted 
to  seeps  and  hanging  gardens. 

Distribution:  San  .luan  (Maguirc 
5904,  Holmgren  13850    [utc];  Moore 


204a;  Welsh  &  Moore  3839  [bry];  Ma- 
guire  1937);  southern  Colorado  and 
New  Mexico. 

Betul.'^ceae 

Betula  X  utahensis  Britton,  Bull.  Torrey 
Bot.  Club  31:   165.  1904. 

Type:  City  Creek  Canyon,  Salt  Lake 
Co.,  Utah,  18  Apr  1900,  Stokes  s.n. 
(ny). 

Status:  Rare;  a  putative  hybrid  be- 
tween B.  occidentalis  Hook  and  B.  papy- 
rifera  Marsh. 

Distribution:  In  Utah  known  only 
from  the  t}'pe  locality;  the  hybrid  is 
widespread  north  of  Utah  (Dugle 
1966). 

Ostrya  knowltonii  Coville,  Gard.  &  Forest 
7:' 114.  1894. 

St.atus:  Rare,  disjunct  in  hanging 
gardens,  along  seeps  and  on  slickrock 
in  sandstone  canyons. 

Distribution:  Grand  [Cottam  2145 
and  556  [bry,  utc]),  Kane  {Welsh  & 
Toft  11871  [bry]),  San  Juan  (Welsh 
et  al.  2939.  2961  and  3721;  Moore 
336;  Welsh  11893;  Welsh  &  Moore 
11783;  Welsh  &  Atwood  11693  and 
11711;  Atwood  4103  [bry]);  northern 
Arizona,  southwestern  New  Mexico 
and  western  Texas  (Little  1953;  Correll 
&  Johnston  1970). 

Boraginaceae 

Cryptantha  barnebyi  I.  M.  Johnston,  J. 
Arnold  Arbor.  29:  240.  1948. 

Type:  Ca  30  miles  south  of  Ouray 
on  white  shale  knolls,  Uintah  Co.,  Utah, 
17  Jun  1947,  Ripley  &  Barneby  8748 
(gh). 

Status:  Endemic,  rare,  threatened 
(Ripley,  T). 

Distribution:  Uintah  Co.,  Utah 
(Higgins  1971). 

Cryptantha  brevi flora  (Osterh.)  Payson, 
Ann.  Missouri  Bot.  Gard.  14:  318.  1927, 
based  on  Oreocarya  breviflora  Osterh. 
ex  Payson,  Univ.  Wyoming  Publ.  Sci., 
Bot.  1:   169.  1926. 

Type:  6.5  miles  north  of  Jensen, 
Uintah  Co.,  Utah,  19  Jun  1925,  Oster- 
hout  6414  (rm). 

St.-vtus:  Endemic.  .  locally  common 
ill  the  Uinta  Basin,  neither  threatened 
nor   endangered    (Ripley,   E). 


Dec.  1975 


WELSH,   ET  AL.:    ENDANGERED    UTAH    PLANTS 


339 


Distribution:  Daggett,  Duchesne 
and  Uintah  counties,  Utah  (Higgins 
1971). 

Cryptantha  capitata  (Eastw.)  I.  M. 
Johnston,  J.  Arnold  Arbor.  21:  66.  1941, 
based  on  Oreocarya  capitata  Eastw., 
Leafl.  W.  Bot.  1:  9.  1937. 
Status:  Rare  and  obscure. 
Distribution:  Garfield,  Kane, 
Washington,  and  Wayne  counties, 
Utah  (Higgins  1971),  and  in  Coconino 
Co.,  Arizona. 

Cryptantha  compacta  Higgins,  Great  Ba- 
sin Nat.  28:   196.  1968. 

Type:  Ca  8  miles  west  of  Desert 
Range  Experiment  Station  headquar- 
ters, along  Utah  Highway  21,  Millard 
Co.,  Utah,  18  Jun  1968/ Higgins  1613 
(bry). 

Status:  Endemic,  rare  and  local, 
threatened  (Ripley,  T). 

Distribution:  Millard  Co.,  Utah 
(Higgins  1971). 

Cryptantha  elata  (Eastw.)  Payson,  Ann. 
Missouri  Bot.  Card.  14:  285.  1927, 
based  on  Oreocarya  elata  Eastw.,  Bull. 
Torrey  Bot.  Club  30:   241.   1903. 

Status:  Rare  and  possibly  threatened 
(Ripley,  T). 

Distribution:  Grand  Co.,  Utah,  and 
adjacent  Mesa  Co.,  Colorado  (Higgins 
1971). 

Cryptantha  grahamii  I.  M.  Johnston,  J. 
Arnold  Arbor.  20:   391.  1939. 

Type:  On  bench  west  of  Green  River, 
north  of  the  mouth  of  Sand  Wash, 
Uintah  Co.,  Utah,  28  May  1933, 
Graham  7924   (gh). 

Status:  Endemic,  locally  abundant, 
on  white  shale  outcrops  along  Willow 
Creek,  possibly  threatened  (Ripley, 
E). 

Distribution:  Uintah  Co.,  Utah 
(Higgins  1971). 

Cryptantha  johnstonii  Higgins,  Great  Ba- 
in Nat.  28:    195.  1968. 

Type:  15  miles  west  of  U.S.  High- 
way 50-6  along  the  road  from  Wood- 
side  to  Castle  Dale,  Emery  Co.,  Utah, 
25  May  1968,  Higgins  1510  (bry). 

Status:  Endemic,  locally  common, 
threatened  (Ripley,  E). 

Distribution:  Emery  Co.,  Utah 
(Higgins  1971). 


Cryptantha  jonesiana  (Payson)  Payson, 
Ann.  Missouri  Bot.  Gard.  14:  323. 
1927,  based  on  Oreocarya  jonesiana 
Payson,  Univ.  Wyoming  Publ.  Sci.,  Bot. 
1:   168.  1926. 

Type:  San  Rafael  Swell,  Emery  Co., 
Utah,  15  May  1914,  M.  E.  Jones  s.n. 
(pom). 

Status:  Endemic,  rare  except  in  clay 
barrens  in  the  San  Rafael  Swell,  threat- 
ened (Ripley,  E). 

Distribution:  San  Rafael  Swell, 
Emery  Co.,  Utah   (Higgins  1971). 

Cryptantha  longiflora  (A.  Nels. )  Payson, 
Ann.  Missouri  Bot.  (7ard.  14:  326. 
1927,  based  on  Oreocarya  loniflora  A. 
Nels.,  Erythea  7:   67.  1899. 

Status:  Rare  and  restricted  in  Grand 
Valley. 

Distribution:  Grand  Co.,  Utah, 
and  west  central  Colorado  (Higgins 
1971). 

Cryptantha  rnensana  (M.  E.  Jones)  Pay- 
son,  Ann.  Missouri  Bot.  Gard.  14:  333. 
1927,  based  on  Krynitzkia  rnensana  M. 
E.  Jones,  Contr.  W.  Bot.  13:  14.  1910. 

Type:  Emery,  Emery  Co.,  Utah,  16 
May  1894,  M.  E.  Jones  5445p  (pom). 

Status:  Rare  and  restricted  to  clay 
soils;  threatened. 

Distribution:  Carbon,  Emery,  and 
Grand  counties,  Utah   (Higgins  1971). 

Cryptantha  ochroleuca  Higgins,  Great  Ba- 
sin Nat.  28:  197.  1968. 

Type:  On  an  outcrop  100  meters 
south  of  Red  Canyon  Campground, 
along  Utah  Highwav  12,  21  Jul  1968, 
Higgins  1788  (bry)^ 

Status:  Endemic,  rare  and  endan- 
gered (Ripley,  E) . 

Distribution:  Garfield  Co.,  Utah 
(Higgins  1971)  . 

Cryptantha  osterhoutii  (Payson)  Payson, 
Ann.  Missouri  Bot.  Gard.  14:  329.  1927, 
based  on  Oreocarya  osterhoutii  Pavson, 
Univ.  Wyoming  Publ.  Sci.,  Bot.  1:"  167. 
1926. 

Status:  Rare  or  obscure. 
Distribution:  Disjunct  in  Duchesne, 
San  Juan,  and  Wayne  counties,  Utah, 
and   in    Mesa    Co.,    Colorado    (Higgins 
1971). 

Cryptantha  paradoxa  (A.  Nels.)  Payson, 
Ann.    Missouri    Bot.    Gard.    14:     330. 


340 


GREAT   BASIN    NATURALIST 


Vol.  35,  No.  4 


1927,  based  on  Oreocarya  paradoxa  A. 
Nels.,  Bot.  Gaz.  56:  69.  1913. 

Status:  Rare,  San  Rafael  Swell, 
threatened. 

Distribution:  Emery  Co.,  Utah, 
western  Colorado  and  northwestern 
New  Mexico  (Higgins  1971). 

Cryptantha  rollinsii  I.  M.  Johnston,  J. 
Arnold  Arbor.  20:   391.  1939. 

Type:  Shale  hillside  on  Thome's 
Ranch,  near  Willow  Creek,  ca  22  miles 
south  of  Ourav,  Uintah  Co.,  Utah,  16 
Jun  1937,  Rollins  1715  (gh). 

Status:  Endemic,  locally  common, 
neither  threatened  nor  endangered. 

Distribution:  Uinta  Basin,  in  both 
Duchesne  and  Uintah  counties,  and  on 
the  San  Rafael  Swell,  Emeny'  Co.,  Utah 
(Higgins  1971). 

Cryptantha  semiglabra  Barneby,  Leafl. 
W.  Bot.  3:   197.  1943. 

Status:  Rare  and  restricted,  threat- 
ened (Ripley,  T) . 

Distribution:  Vicinity  of  Fredonia, 
Coconino,  and  Mohave  counties,  Ari- 
zona, and  in  Washington  Co.,  Utah 
(Higgins    1971). 

Cryptantha  striata  (Osterh.)  Pay  son,  Ann. 
Missouri  Bot.  Card.  14:  264.  1927,  based 
on  Oreocarya  stricta  Osterh.,  Bull.  Tor- 
rey  Bot.  Club  50:  217.  1923. 

Status:  Rare,  but  neither  threatened 
nor  endangered  (Ripley,  T) . 

Distribution:  Daggett,  Summit,  and 
Uintah  counties,  Utah;  also  in  Moffat 
Co.,  Colorado,  and  Carbon  Co.,  Wyo- 
ming (Higgins  1971). 

Cryptantha  tenuis  (Eastw.)  Payson,  Ann. 
Missouri  Bot.  Gard.  14:  327.  1927, 
based  on  Oreocarya  tenuis  Eastw.,  Bull. 
Torrey  Bot.  Club  30:  244.  1903. 

Type:  Court  House  Wash,  near 
Moab,  Grand  Co.,  Utah,  25  May  1892, 
Eastwood  s.n.  (cas). 

St.\tus:  Endemic,  common  and  wide- 
spread. 

Distribution:  Emery,  Grand,  San 
Juan,  and  eastern  Wayne  counties, 
Utah   (Higgins  1971). 

Cryptantha  wether illii  (Eastw.)  Payson, 
Ann.  Missouri  Bot.  Gard.  14:  324. 
1927,  based  on  Oreocarya  wetJierillii 
Eastw.,  Bull.  Torrey  Bot.  Club  30:  242. 
1930. 


Type:  Court  House  Wash,  near 
Moab,  Grand  Co.,  Utah,  25  May  1892, 
Eastwood  s.n.  (cas). 

Status:  Endemic,  common  and  wide- 
spread. 

Distribution:  Carbon,  Emery,  Gar- 
field, Grand,  and  Wayne  counties,  Utah 
(Higgins  1971). 

Hackelia  patens  (Nutt.)  I.  M.  Johnston 
var.  harrisonii  J.  L.  Gentry,  Southw. 
Naturahst  19:    140.  1974. 

Type:  Pine  Valley  Mts.,  ca  1.5  miles 
up  Forsyth  Trail  from  Pine  Valley,  30 
May  1968,  7,300  feet,  Gentry  2002 
(ny). 

Status:  Endemic,  restricted  but  lo- 
cally common. 

Distribution:  Utah,  Wasatch,  and 
Washington  counties,  Utah  (Gentry 
1974). 

Mertensia  arizonica  Greene,  Pittonia  3: 
197.  1897. 

Type:  "Arizona,"  without  definite 
locality,  but  more  likely  from  southern 
Utah,  1869,  Palmer  s.n.   (us). 

Status:  Endemic  (?),  locally  com- 
mon; the  var.  arizonica  is  not  known 
from   Arizona    in    modern    times. 

Distribution:  Beaver,  Garfield,  Iron, 
Piute,  and  Washington  counties,  Utah 
(Higgins  1972a). 

Mertensia  viridis  A.  Nels.  var.  cana 
(Rydb.)  L.  O.  Williams,  Ann.  Missouri 
Bot.  Gard.  24:  115.  1937,  based  on  M. 
cana  Rydb.,  Bull.  Torre^'  Bot.  Club  36: 
698.  1909. 

Status:  Rare  and  restricted,  possibly 
threatened. 

Distribution:  Bald  Mtn.,  Summit 
Co.,  Utah  (Maguire  14699;  Weber 
5894  [uTc] )  and  north  central  Colorado 
(Williams    1937;    Higgins    1972a). 

Mertensia  viridis  A.  Nels.  ^-ar.  dilatata 
(A.  Nels.)  L.  O.  Williams,  Ann  Mis- 
souri Bot.  Gard.  24:  113.  1937,  based  on 
M.  coriacea  var.  dilatata  A.  Nels.,  Bull. 
Torrey  Bot.  Club  29:  403.  1902. 

Status:  Rare,  known  from  a  single 
collection  from  the  Uinta  Mts. 

Distribution:  Daggett  Co.,  Utah 
{Williams  599  |rm]);  southeastern 
Wyoming  and  adjacent  Colorado  (Wil- 
liams 1037;  Higgins  1972a). 


Dec.  197'. 


WELSH.   ET   AL.:    ENDANGERED    UTAH    PLANTS 


341 


Brassicaceae 

Arabis  deniissa  Greene  ^ar.  languida  Rol- 
lins, Rhodora  43:  388.  1941. 

Status:  Rare  and  possibly  threat- 
ened (Ripley,  T). 

Distribution:  Daggett  Co.,  Utah 
(Rollins  1941);  Albany  and  Sweet- 
water counties,  Wyoming. 

Arabis  dcmissa  Greene  \c\r.  russcola  Rol- 
lins, Rhodora  43:  387.  1941. 

Type:  Ca  18  miles  north  of  Vernal, 
Uintah  Co.,  Utah,  Jun  1937,  Rollins 
1757  (gh). 

Status:  Rare  and  threatened  (Rip- 
ley, T). 

Distribution:  Daggett  and  Uintah 
counties,  Utah,  and  Albany  Co.,  Wyo- 
ming (Rollins  1941). 

Caulanthus  divaricatus  Rollins.  Contr. 
Gray  Herb.  201:   8.  1971. 

Type:  Ca  75  miles  west  of  Blanding 
and  10  miles  east  of  Hite,  San  Juan 
Co.,  Utah,  16  May  1961,  Cronquist 
9033   (NY). 

Status:  Endemic,  rare  though  wide- 
spread, neither  threatened  nor  endan- 
gered. 

Distribution:  Carbon,  Emery,  Gar- 
field, Grand,  San  Juan,  and  Wayne 
counties,  Utah  (bry.  utc). 

Draba  maguirei  C.  L.  Hitchc,  Univ. 
Wash.  Publ.   Biol.    11:    71.   1941. 

Type:  Rocky  soil  on  east  slope  of  Mt. 
Naomi,  Bear  River  Range,  Cache  Co., 
Utah,  9,600  feet,  Maguire  et  al.  14161 
(wtu). 

Status:  Endemic,  restricted  and 
possibly  threatened. 

Distribution:  Cache  Co..  Utah 
(Hitchcock  1941). 

Draba  maguirei  C.  L.  Hitchc.  var.  burkei 
C.  L.  Hitchc,  Univ.  Wash.  Publ.  Biol. 
11:  72.  1941. 

Type:  Cottonwood  Canyon,  Wells- 
ville  Mts.,  Box  Elder  Co.,  Utah,  25  May 
1932,  Burke  2968  (utj. 

Status:  Endemic,  restricted  and 
threatened. 

Distribution:  Box  Elder  (Hitch- 
cock 1941)  and  Weber  {Clark  2332 
[bry,  wsco])   counties,  Utah. 

Draba  pectinipila  Rollins,  Rhodora  55: 
231.  1953. 


Status:  Endangered  or  possibly  ex- 
tirpated. 

Distribution:  Flaming  Gorge,  Dag- 
gett Co.,  Utah,  and  adjacent  Sweet- 
water Co.,  Wyoming  (Rollins  1953). 

Draba  sobolifera  Rydb.,  Bull.  Torrev  Bot. 
Club  30:  251.  1903. 

Type:  Marysvale,  Piute  Co.,  Utah, 
above  timber  line,  1894,  M.  E.  Jones 
5936  (us). 

Status:  Endemic,  rare  and  threat- 
ened   (Ripley,  T). 

Distribution:  Garfield  and  Piute 
counties,   Utah    (Hitchcock    1941). 

Draba  subalpina  Goodman  &  Hitchc, 
Ann.  Missouri  Bot.  Card.  19:  77.  1932. 

Type:  Cedar  Breaks,  Iron  Co.,  Utah, 
Goodman  &  Hitchcock  1622  (mo). 

Status:  Endemic,  infrequent  and 
possibly  threatened   (Ripley,  T). 

Distribution:  Garfield,  Iron  and 
Kane   counties,  Utah    (bry,    utc). 

Draba  zionensis  C.  L.  Hitchc,  Univ. 
Wash.  Publ.  Biol.  11:  49.  1941. 

Type:  Zion  Canyon,  Washington  Co., 
Utah,  7  May  1923,  M.  E.  Jones  s.n. 
(pom). 

Status:  Endemic,  rare  and  threat- 
ened (Ripley,  T). 

Distribution:  Juab  (Cottam  7201 
UT)  and  Washington  counties,  Utah 
(Hitchcock  1941). 

Glaucocarpum  suffrutescens  (Rollins) 
Rollins,  Madrono  4:  233.  1938,  based 
on  Thelypodium  suffrutescens  Rollins 
in  Graham,  Ann.  Carnegie  Mus.  26: 
244.  1937. 

Type:  West  of  Willow  Creek,  on 
Thome's  Ranch,  eastern  slope  of  Big 
Pack  Mtn.,  Uintah  Co.,  Utah,  23  May 
1935,  Graham  8950   (gh). 

Status:  Endemic,  rare  and  endan- 
gered   (Ripley,  E). 

Distribution:  Uintah  Co.,  Utah; 
known  only  from  the  tApe  locality 
(Rollins   1938). 

Lepidium  barnebyanum  Reveal,  Great  Ba- 
sin Nat.  27:  178.  1967,  based  on  L.  mon- 
tanum  Nutt.  ex  Torr.  &  Gray  ssp.  de- 
missum  C.  L.  Hitchc,  Madrofio  10:  157. 
1950. 

Type:  Indian  Creek  Canyon,  on 
white  shale  ridge  tops,  ca  4  miles  south- 
west of  Duchesne,  Duchesne  Co.,  Utah, 


342 


GREAT   BASIN   NATURALIST 


Vol.  35,  No.  4 


15  Jun   1947,  Ripley  &  Barneby  8699 

(WTU). 

Status:  Endemic,  rare  and  endan- 
gered (Ripley,  E). 

Distribution:  Duchesne  Co.,  Utah; 
knowTi  onl}'  from  the  type  locality 
(Reveal  1967). 

Lesquerella  garrettii  Payson,  Ann.  Mis- 
souri Bot.  Gard.  8:  213.  1921. 

Type:  Big  Cottonwood  Canyon,  Salt 
Lake  Co.,  Utah,  28  Jun  1908,  Garrett 
1544  (mo). 

Status:  Endemic,  rare  and  possibly 
threatened   (Ripley,  T). 

Distribution:  Salt  Lake,  Utah  and 
Wasatch  counties,  Utah  (Rollins  & 
Shaw  1973). 

Lesquerella  hemiphysaria  Maguire, 
Amer.  Midi.  Naturalist  27:   456.  1942. 

Type:  Frequent,  limestone  breaks, 
south  side  of  Middle  Fork  Park,  Wa- 
satch Plateau,  10  Aug  1940,  10,800 
feet,  Maguire  20053    (utc). 

Status:  Endemic,  locally  common 
but  restricted. 

Distribution:  Carbon,  Emery,  San- 
pete, and  Utah  counties,  Utah  (Rollins 
&  Shaw  1973). 

Lesquerella  rubicundula  Rollins,  Contr. 
Dudley  Herb.  3:   178.  1941. 

Type:  Red  Canyon,  Garfield  Co., 
Utah,  2,300  meters,  6  Jul  1912,  Eggle- 
ston  8198  (na,  the  type  now  trans- 
ferred to  us). 

Status:  Endemic,  rare  and  threat- 
ened (Ripley,  T) . 

Distribution:  Garfield,  Kane,  and 
Piute  counties,  Utah  (Reveal  1970a; 
Rollins  &  Shaw  1973). 

Lesquerella  tumulosa  (Barneby)  Reveal, 
Great  Basin  Nat.  30:  97.  1970,  based 
on  L.  hitchcockii  Munz  ssp.  tumulosa 
Barneby,  Leafl.  W.  Bot.  10:  313.  1966. 

Type:  Bare  white  shale  knolls,  6.5 
miles  southeast  of  Cannonville,  Kane 
Co.,  Utah,  12  Jun  1966,  Barneby 
14424  (NYj. 

Status:  Endemic,  rare  and  endan- 
gered. 

Distribution:  Kane  Co.,  Utah; 
known  only  from  the  type  locality 
(Reveal    1970a). 

Lesquerella  utahensis  Rydb.,  Bull.  Torre v 
Bot.  Club  30:  252.  1903. 


Type:  American  Fork  Canyon,  Utah 
Co.,  Utah,  31  Jul  1880,  M.  E.  Jones 
1354  (ny). 

Status:  Endemic,  rare  to  locally  com- 
mon. 

Distribution:  Davis,  Duchesne, 
Juab,  Salt  Lake,  Summit,  Tooele, 
Uintah,  Utah,  Wasatch  and  Weber 
counties,  Utah  (RoUins  &  Shaw  1973). 

Parry  a  rydbergii  Botsch.,  Bot.  Mater. 
Gerb.  Bot.  Inst.  Komarova  Akad.  Nauk 
SSSR  17:  178.  1955,  based  on  P.  platy- 
carpa  Rydb.,  Bull.  Torrey  Bot.  Club  39: 
326.  1912,  not  Hook.  f.  &  Thomas. 

Type:  Uinta  Mts.,  Aug  1869,  Wat- 
son 54  (ny). 

Status:  Endemic,  rare  and  threat- 
ened (Ripley,  T). 

Distribution:  Crest  of  the  Uinta 
Mts.,  Duchesne  {Murdoch  54  [bry]) 
and  Daggett-Uintah  counties  {Waite 
252  and  297   [bry]). 

Physaria  grahamii  Morton  in  Graham, 
Ann.   Carnegie  Mus.  26:    220.   1937. 

Type:  Chandler  Canyon,  Uintah  Co., 
Utah,  3  Aug  1935,  6,000  feet,  Graham 
9976  (us). 

Status:  Endemic,  rare  and  possibly 
extinct  (Ripley,  PoEx). 

Distribution:  Uintah  Co.,  Utah; 
known  only  from  the  type  collection 
as  recent  efforts  to  recollect  this  spe- 
cies have  failed  (Waite  1973). 

Thely podium  integrifolium  (Nutt.)  Endl. 
var.  complanatum  (Al-Shehbaz)  Welsh 
&  Reveal,  stat.  nov.,  based  on  T.  in- 
tegrifolium ssp.  complanatum  Al-Sheh- 
baz, Contr.  Gray  Herb.  204:  105.  1973. 
Status:  Local  and  infrequent. 
Distribution:  Box  Elder  Co.,  Utah 
(uTC;  Al-Shehbaz  1973);  found  to  the 
west  of  LTtah. 

Thelypodium  rollinsii  Al-Shehbaz.  Contr. 
Gray  Herb.  204:    97.   1973. 

Type:  On  alkaline  soil  in  the  vicinity 
of  the  Sevier  RiAer.  ca  12  miles  north 
of  Scipio  along  U.S.  Highway  91,  29 
Jul  1969,  Al-Shehbaz  &■  Al-Shehbaz 
6913   (gh). 

Status:  Endemic,  locally  common, 
neither  threatened  nor  endangered. 

Distribution:  Carbon,  Juab,  Mil- 
lard, Piute,  Sanpete,  and  Sevier  coun- 
ties, Utah   (Al-Shehbaz  1973). 


Dec.  1975 


WELSH.   ET  AL.:    ENDANGERED   UTAH   PLANTS 


343 


Thely podium  sagittatum  (Nutt.)  Endl. 
ex  Walp.  var.  ovalifoUum  (Rydb.) 
Welsh  &  Reveal,  stat.  &  comb,  nov., 
based  on  T.  ovaJifoUum  Rydb.,  Bull. 
Torrey  Hot.  Club  30:   253.   1903. 

Type:  Pangnitch  Lake,  Garfield  Co., 
Utah,  7  Sep  1894,  M.  E.  Jones  601 5e 
(us). 

Status:  Rare  and  restricted,  possibly 
threatened. 

Distribution:  Garfield  and  Iron 
counties,  Utah,  and  White  Pine  Co.. 
Nevada  (Al-Shehbaz  1973). 

Cactaceae 

Echinocereus  engelmannii  (Parry)  Le- 
maire  var.  purpurcus  L.  Benson,  Cact. 
Succ.  J.   (Los  Angeles)   41:    127.   1969. 

Type:  Mojavean  desert  north  of  St. 
George,  Washington  Co.,  Utah,  5  May 
1949,  2,900  feet,  Benson  15637   (pom). 

Status:  Endemic,  rare  and  endan- 
gered (Ripley,  E). 

Distribution:  Washington  Co., 
Utah;  known  only  from  the  vicinity  of 
the  type  locality  (Benson  1969b). 

Ferocactus  acanthodes  (Lemaire)  Britton 
&  Rose,  Publ.  Carnegie  Inst.  Wash.  248 
(3):  129.  1922,  based  on  Echinocactus 
acanthodes  Lemaire,  Cact.  Gen.  Nov. 
Sp.  106.  1839. 

Status:    Restricted  and  endangered. 
Distribution:        Washington       Co., 
Utah  (bry);  southern  Nevada,  Arizona 
and  California  (Benson  1969b). 

Mammillaria  tetrancistra  Engelm.,  Amer. 
J.  Sci.  Arts  II,  14:  337.  1852. 

Status:  Restricted  and  rare,  possibly 
threatened. 

Distribution:  Washington  Co., 
Utah;  southern  Nevada,  California, 
and  Arizona   (Benson  1969b). 

Opuntia  pulchella  Engelm.,  Trans.  Acad. 
Sci.  St.  Louis  2:  201.  1863. 

Status:  Rare,  restricted  and  possibly 
threatened. 

Distribution:  Millard  Co.,  Utah 
(bry);  Nevada  and  Arizona  (Benson 
1957).  This  species  includes  those  en- 
tities included  in  Micropuntia  by  Das- 
ton  (1946). 

Opuntia  whipplei  Engelm.  &  Bigelow  var. 
multigeniculata  L.  Benson,  Cact.  Ariz. 
ed.  3,  38.  1969. 


Status:  Restricted  and  rare. 

Distribution:  Washington  Co., 
Utah,  and  adjacent  Nevada  and  Arizona 
(Benson  1969b). 

Pediocactus  silcri  (Engelm.)  L.  Benson, 
Cact.  Succ.  .1.  (Los  Angeles)  33:  53. 
1961,  based  on  Echinocactus  sileri  En- 
gelm. ex  Coulter,  Contr.  U.S.  Natl. 
Herb.  3:  376.  1896. 

Type:  Cottonwood  Springs  and  Pipe 
Springs,  supposedly  from  southern 
Utah  but  more  likely  from  Arizona, 
1883,  Siler  s.n.  {mo)/ 

Status:  Arizona  Strip  endemic,  rare, 
local  and  endangered   (Ripley,  E). 

Distribution:  Washington  Co., 
Utah  {Welsh  12712  [bry]),  and 
northern  Arizona  (bry;  Benson  1969b). 

Sclerocactus  glaucus  (K.  Schum.)  L.  Ben- 
son, Cact.  Succ.  J.  (Los  Angeles)  38:  53. 
1966,  based  on  Echinocactus  glaucus 
K.  Schum.,  Gessammtb.  Kakt.  438. 
1898. 

Status:  Rare,  possibly  extirpated 
from  Utah  (Ripley,  E). 

Distribution:  Duchesne  and  Uintah 
counties,  Utah,  and  Delta  Co.,  Colorado 
(Benson  1966). 

Sclerocactus  pubispinus  (Engelm.)  L.Ben- 
son, Cact.  Succ.  J.  (Los  Angeles)  38: 
103.  1966,  based  on  Echinocactus  pubi- 
spinus Engelm.,  Trans.  Acad.  Sci.  St. 
Louis  2:  199.  1863. 

Status:  Rare  and  threatened  (Rip- 
ley,T). 

Distribution:  Box  Elder,  Beaver, 
and  Sevier  counties,  Utah,  and  Elko 
Co.,  Nevada  (Benson  1966). 

Sclerocactus  pubispinus  (Engelm.)  L. 
Benson  var.  sileri  L.  Benson,  Cact. 
Ariz.  ed.  3,  23.  1969. 

Type:  Southern  Utah,  1888,  Siler  s.n. 

(PH). 

St.\tus:  Rare  and  now  extirpated 
from  Utah. 

Distribution:  Kane  Co.,  Utah,  and 
Coconino  Co.,  Arizona  (Benson  1969b). 

Sclerocactus  whipplei  (Engelm.  &  Bige- 
low) Britton  &  Rose  var.  spinosior 
(Engelm.)  Boissev.  in  Boissev.  &  Da- 
vidson, Colorado  Cacti  51.  1940,  based 
on  Echinocactus  whipplei  var.  spinosior 
Engelm.,  Trans.  Acad.  Sci.  St.  Louis 
2:   199.  1863. 


344 


GREAT   BASIN    NATURALIST 


Vol.  35,  No.  4 


Type:  South  of  the  Great  Sah  Lake 
Desert,  19  Jul  1859,  H.  Engelmann  s.n. 
(mo). 

Status:  Rare  to  locally  common, 
neither  threatened  nor  endangered. 
<Ripley,T).^ 

Distribution:  Widespread  in  much 
of  the  southern  two-thirds  of  Utah. 

Sclerocactus  ivrightiae  L.  Benson,  Cact. 
Succ.  J.   (Los  Angeles)  38:  55.  1966. 

Type:  Near  San  Rafael  Ridge, 
Emery  Co.,  Utah,  5,000  feet,  Benson 
&  Benson  16595  (pom). 

Status:  Endemic,  restricted  and  rare, 
endangered   (Ripley,  T). 

Distribution:  Emery  and  Wayne 
counties,  Utah. 

Capparidaceae 

Cleomella  palmerana  M.  E.  Jones,  Zoe 
2:  236.  1891. 

Type:  Green  River,  Emery  Co., 
Utah,  9  May  1890,  M.  E.  Jones  s.n. 
(pom). 

Status:  Endemic  to  the  Navajo  Ba- 
sin of  Utah  and  Colorado,  edaphically 
restricted  but  neither  threatened  nor 
endangered. 

Distribution:  Duchesne,  Emery, 
Garfield,  Grand,  Kane,  and  Wayne 
counties,  Utah  (bry),  and  Colorado 
(Harrington  1954). 

Cleomella  placosperma  S.  Wats,  in  King, 
Rep.  Geol.  Explor.  40th  Parallel  5:  33. 
1871. 

Status:  Restricted  and  rare,  possibly 
threatened. 

Distribution:  Iron  Co.,  Utah  (bry), 
and  Nevada  (Shaw  1970). 

Caryophyllaceae 

Arenaria  kingii  (S.  Wats.)  M.  E.  Jones 
var.  plateauensis  (Maguire)  Reveal, 
Stat,  nov.,  based  on  A.  kingii  ssp. 
plateauensis  Maguire,  Bull.  Torrey  Bot. 
Club  74:  54.  1947. 

Type:  Cedar  Breaks  Rim,  Iron  Co., 
Utah,  23  Jun  1940,  10.400  feet,  Ma- 
guire 19024  (uTc). 

Status:  Endemic,  locally  common, 
neither  threatened   nor  endangered. 

Distribution:       Beaver,       Garfield, 


Kane,  Iron,  and  Wayne  counties,  Utah 
(Maguire  1947b). 

Silene  petersonii  Maguire,  Madroiio  6: 
24.  1941. 

Type:  Skyline  Drive,  1  mile  above 
Baldy  Ranger  Station,  Sanpete  Co., 
Utah,  8  Aug  1940,  10,900  feet,  Ma- 
guire 20000  (uTc). 

Status:  Endemic,  locally  common 
but  possibly  threatened    (Ripley,  T). 

Distribution:  Iron,  Garfield  and 
Sanpete  counties,  Utah  (Maguire  1941; 
Hitchcock  &  Maguire   1951). 

Silene  petersonii  Maguire  var.  minor 
Hitchc.  &  Maguire,  Univ.  Wash.  Publ. 
Biol.   13:    38.   1947. 

Type:  Red  Canyon,  5  miles  east  of 
U.S.  Highway  89^  Garfield  Co.,  Utah, 
16  Jul  1940,  Maguire  19550   (utc). 

Status:  Endemic,  rare  and  threat- 
ened. 

Distribution:  Garfield  Co.,  Utah; 
known  only  from  the  type  area. 

Chenopodiaceae 

Atriplex  bonnevillensis  C.  A.  Hanson, 
Stud.  Syst.  Bot.  1:  2.  1962. 

Type:  Dry  lake  bed  1.5  miles  north- 
east of  Desert  Range  Experiment  Sta- 
tion headquarters,  Millard  Co.,  12  Jul 
1961,  Hanson  554   (bry). 

Status:  Restricted,  though  locally 
abundant,  possibly  threatened. 

Distribution:  Juab  and  Millard 
counties,  Utah  (bry),  and  Nevada 
(Hanson  1962b). 

Atriplex  garrettii  Rydb.,  Bull.  Torrev  Bot. 
Club  39:  312.  1912. 

Type:  Vicinity  of  Moab,  Grand  Co., 
Utah,  1  Jul  1911,  Rydberg  &  Garrett 
8465   (ny). 

St.a,tus:  Colorado  River  drainage,  en- 
demic, restricted  but  locally  common. 

Distribution:  Garfield,  Grand, 
Kane,  and  San  Juan  counties,  Utah 
(bry),  and  reported  from  Colorado 
(Harrington  1954). 

Atriplex  obovata  Moq.,  Chenop.  Enum. 
61.   1840. 

Status:  Restricted,  rare  and  possibly 
threatened. 

Distribution:    San  Juan  Co.,  Utah; 


<Citc(l    by    Ripley    (1975)    as   S.    spinosior.    without    author;    \v<>    lau    fliul    no    indication    that    such    a 
made.   According  to   Benson    (1966),   this  name   is  a   synon.^^n   of    .V.    inibispiiuis    (Engohii.i    I..    Benson 


combination    h 
var.    pubspinus. 


Dec.  1975 


WELSH,   ET  AL.:    ENDANGERED    UTAH    PLANTS 


345 


rather  common  in  Arizona,  New  Mexi- 
co and  Mexico  (Hanson  1692b). 

Atriplex  welshii  C.  A.  Hanson,  Stud. 
Syst.  Bot.  1:   1.  1962. 

Type:  4  miles  south  of  Cisco  along 
Utah  Highway  128,  Grand  Co.,  Utah, 
5  Jul  1961,  Hanson  522  (bry). 

Status:  Endemic,  restricted  and  pos- 
sibly threatened. 

Distribution:  Carbon  and  Grand 
counties,  Utah   (bry)  . 

Cucurbitaceae 

Cucurbita  foetidissima  H.B.K.,  Nov.  Gen. 
&Sp.  2:  123.  1817. 

Status:  Restricted,  but  locally  com- 
mon and  often  weedy,  neither  threat- 
ened nor  endangered. 

Distribution:   San  Juan  {Welsh  et 
al.  2992  [bry]),  and  Washington  (bry) 
counties,    Utah,    and    widespread    else- 
where. 

CUSCUTACEAE 

Cuscuta  warneri  Yuncker,  Brittonia  12: 
38.  1960. 

Type:  Vicinity  of  Powell,  15  miles 
west  of  Fillmore,  Millard  Co.,  Utah, 
10  Sep  1957,  on  Phyla  cuneifolia,  War- 
ner s.n.  (uTc). 

Status:  Endemic,  possibly  extinct 
(Ripley,  PoEx). 

Distribution:  Millard  Co.,  Utah; 
kno\vn  only  from  the  type  collection 
(Yuncker  1960,  1965). 


Cyperaceae 

Car  ex  curatorum  Stacey,  Leafl.  W.  Bot. 
2:   213.   1937. 

Status:  Rare  and  restricted  to  spec- 
ialized habitats;  threatened. 

Distribution:  San  Juan  Co.,  Utah 
{Welsh  12425  [bry]),  and  in  Grand 
Canyon,  Arizona  (Stacey  1937). 

Cladium  californicum  (S.  Wats.)  O'Neill 
in  Tidestr.  &  Kittell,  Fl.  Arizona  &  N. 
Mex.  773.  1941,  based  on  C.  mariscus 
var.  californicum  S.  Wats.,  Bot.  Calif. 
2:  224.  1880. 

Status:  Rare,  restricted  and  endan- 
gered. 

Distribution:  Kane  {Welsh  &  At- 
wood  11709;  Welsh  &  Moore  11780; 
Welsh  &  Toft  11870   [bry])   and  San 


Juan  {Welsh  &  Toft  11874  [bry]),  and 
from  northern  Arizona,  southern  Ne- 
vada, and  southern  California  to  Cen- 
tral America  (Kearney  &  Peebles  1951). 


Ela 


EAGNACEAE 


Elaeagnus  commutata  Bemh.,  Allg.  Thij- 
ring.   Gartenzeitung  2:    137.    1843. 

Status: Restricted,  local  and  possibly 
extirpated  from  Utah. 

Distribution:  Daggett  {Julander 
J56  [bry])  and  Summit  {Welsh  & 
Moore  6724  [bry]  )  counties,  Utah; 
widespread  northward  to  Alaska 
(Welsh  1974b). 

Ericaceae 

Arctostaphylos     pringlei     Parry,      Bull. 
Calif.  Acad.  Sci.  2:  494.  1887. 
Status:   Local,  rare,  possibly  threat- 
ened. 

Distribution:  Washington  Co.,  Utah 
{Higgins  665  and  3414  [bry];  Higgins 
1972b). 

EUPHORBIACEAE 

Euphorbia  nephradenia  Barneby,  Leafl. 
W.  Bot.  10:  314.  1966. 

Type:  Lower  Cottonwood  Canyon, 
about  41  miles  southeast  of  Cannonville, 
Kane  Co.,  Utah,  12  Jun  1966,  4,500 
feet,  Barneby  14434  (ny). 

Status:  Endemic,  restricted  edaph- 
ically,  threatened  (Ripley,  T). 

Distribution:  Emery,  Kane,  and 
Wayne  counties,  Utah   (bry). 

Fabaceae 

Astragalus  adanus  A.  Nels.,  Bot.  Gaz.  53: 
222.  1912. 

Status:  Rare  and  possibly  extirpated 
from  Utah. 

Distribution:  Juab  Co.,  Utah;  wide- 
spread in  Idaho  (Barneby  1964). 

Astragalus  adsurgens  Pallas  ssp.  robus- 
tior  (Hook.)  Welsh,  Iowa  State  Coll. 
J.  Sci.  37:  357.  1963,  based  on  A.  ad- 
surgens var.  robustior  Hook.,  Fl.  Bo- 
reali-Amer.    1:    149.   1831. 

Status:  Restricted,  rare  and  possibly 
threatened. 

Distribution:  Daggett  Co.,  Utah 
{Hanson  283a  [bry]);  widespread 
east  of  the  continental  divide  north- 
ward to  Alaska  (Welsh  1974b). 


346 


GREAT   BASIN   NATURALIST 


Vol.  35,  No.  4 


Astragalus  alpinus  L.,  Sp.  PI.  760.  1753. 

Status:  Rare  and  local,  possibly 
threatened. 

Distribution:  Grand  {Holmgren  et 
al.  232 J  [bry,  ny,  utc]  )  and  Salt  Lake 
(ut,  without  collector)  counties;  wide- 
spread in  North  America  and  Eurasia 
(Welsh  1974b). 

Astragalus  ampullarius  S.  Wats.,  Amer. 
Naturalist  7:  300.  1873. 

Type:  Kanab,  Kane  Co.,  Utah,  1872, 
Thompson  s.n.  (gh). 

Status:  Arizona  strip  endemic, 
edaphically  restricted  and  though 
locally  common,  probably  threatened 
(Ripley,  T). 

Distribution:  Kane  and  Washington 
counties,  Utah,  and  northernmost  Co- 
conino and  Mohave  counties,  Arizona 
(bry,-  Rarneby  1964). 

Astragalus  asclepiadoides  M.  E.  Jones, 
Zoe2:  238.  1891. 

Type:  Cisco,  Grand  Co.,  Utah,  21  Jun 
1889,  M.  E.  Jones  s.n.  (pom) 

Status:  Edaphically  restricted  al- 
though locall}'  common,  neither  threat- 
ened nor  endangered. 

Distribution:  Carbon,  Duchesne, 
Emery,  Garfield,  Grand,  Sanpete,  Se- 
vier (ut),  Uintah,  and  Wayne  coun- 
ties, Utah  (bry,  utc),  and  western 
Colorado  (Rarneby  1964). 

Astragalus  aus trails  Lam.,  Fl.  Frang.  2: 
637.  1778. 

Status:  Possibly  extirpated  from 
Utah. 

Distribution:  Piute  (?)  Co.,  Utah 
(Rarneby  1964). 

Astragalus  barneby  Welsh  &  Atwood, 
nom.  nov.,  based  on  A.  desperatus  M. 
E.  Jones  var.  conspectus  Rarneby, 
Leafl.  W.  Rot.  5:  87.  1948. 

Status:  Rare,  restricted  and  threat- 
ened (Ripley,  T). 

Distribution:  Garfield  Co.,  Utah 
(bry),  and  in  Navajo  and  Coconino 
counties,  Arizona    (Rarneby   1964). 

This  remarkable  milkvetch  is  adequate- 
ly distinct  on  account  of  size  of  flower  and 
parts,  and  because  of  stature  to  segregate 
it  from  its  near  and  mirror-imaged  cog- 
ener  A.  desperatus.  The  name  change 
honors  the  author  of  the  monumental 
"Atlas   of  North   American   Astragalus," 


Rupert  C.  Rarneby  of  the  New  York  Ro- 
tanical  Garden. 

Astragalus  bodinii  Sheldon,  Minnesota 
Rot.  Stud.  1:   122.  1894. 

Status:  Rare  and  endangered. 

Distribution:  Wayne  Co.,  Utah 
(bry);  widespread  in  Alaska,  Canada, 
Colorado,  Wyoming,  and  one  record 
from  Nebraska  (Rarneby  1964). 

Astragalus  brandegei  Porter  in  Porter  & 
Coulter,  Syn.  Fl.  Colo.  24.  1872. 

Status:  Rare  and  obscure,  probably 
threatened. 

Distribution:  Carbon,  Emery,  Gar- 
field, Piute,  and  Wayne  counties,  Utah 
(bry)  ;  also  in  Colorado,  New  Mexico, 
and  Arizona  (Rarneby  1964). 

Astragalus  bryantii  Rarneby,  Proc.  Calif. 
Acad.  Sci.  IV,  25:  156.  1944. 

Status:  Probably  extirpated  from 
Utah. 

Distribution:  Kane  Co.,  Utah,  in 
Glen  Canyon,  the  collection  area  inun- 
dated by  Lake  Powell;  also  in  northern 
Arizona   (Rarneby  1964). 

Astragalus  callithrix  Rarneby,  Leafl.  W. 
Rot.  3:  102.  1942. 

Status:  Rare  and  restricted,  possibly 
threatened  (Ripley,  T) . 

Distribution:  Millard  Co.,  Utah 
(bry);  otherwise  known  only  from  two 
locations  in  Nye  Co.,  Nevada. 

Astragalus  castaneiformis  S.  Wats.  var. 
consobrinus  Rarneby,  Amer.  Midi. 
Naturalist  41:  496.   1949. 

Type:  Southeast  of  Ricknell,  Wayne 
Co.,  Utah,  10  Jun  1947,  7,600  feet, 
Ripley  &  Barneby  8605   (cas). 

Status:  Endemic,  rare  and  obscure, 
possibly  threatened. 

Distribution:     Garfield,    Piute,    Se- 
vier, and  Wayne  counties,  Utah   (Rar- 
neby 1964). 
Astragalus   chloodes   Rarneby,    Leafl.    W. 
Rot.  5:  6.  1947. 

Type:  Ca  6  miles  southeast  of  Jen- 
sen, Uintah  Co.,  Utah,  7  Jun  1946, 
Ripley  &  Barneby  7797  (g.\s). 

Status:  Endemic,  rare  and  threat- 
ened (Ripley,  T). 

Distribution:      Uintah     Co.,     Utah 
(BRY.  utc). 
Astragalus  coltonii  M.   E.   Jones,   Zoe  2: 
237.  1891. 


Dec.  1975 


WELSH,    ET   AL.:     ENDANGERED    UTAH    PLANTS 


347 


TYPE:  Canyons  of  the  Coal  Range 
east  of  Castle  Gate,  27  .Tun  1889,  M. 
E.  Jones  s.n.  (pom)  . 

Status:  Endemic,  locally  abundant, 
neither  threatened  nor  endangered.  Our 
plant  is  var.  coltonii. 

Distribution:  Carbon,  Emery,  Gar- 
field, Kane,  Sevier,  and  Wayne  coun- 
ties, Utah  (bry,  ut.  utc)  . 

Astragalus  convaUarius  Greene  var.  finiti- 
mus  Barneby,  Leafl.  W.  Bot.  7:  192. 
1954. 

Type:  Ca  3  miles  south  of  Enterprise, 
Washington  Co.,  Utah,  11  Jun  1942, 
Ripley  &  Barneby  4767  (gas). 

Status:  Rare  and  local,  threatened 
^(Ripley,  T). 

Distribution:  Washington  Co.,  Utah, 
and  adjacent  Nevada    (Barneby  1964). 

Astragalus  cottamii  Welsh,  Rhodora  72: 
189.  1970. 

Type:  Ca  4  miles  east  of  Clay  Hills 
divide,  San  Juan  Co.,  Utah,  1  May 
1966,  Welsh  5207  (bry). 

Status:  Restricted  and  local,  possibly 
threatened  (Ripley,  T). 

Distribution:  San  Juan  Co.,  Utah, 
and  adjacent  Monument  Valley,  Ari- 
zona (bry). 

Astragalus  cronquistii  Barneby,  Mem. 
New  York  Bot.  Gard.  13:  258.  1964. 

Type:  In  desert  along  west  side  of 
Comb  Wash,  9  miles  west  of  Bluff, 
San  Juan  Co.,  Utah,  27  May  1961, 
Cronquist  9123  (ny). 

Status:  Endemic  and  very  restricted, 
endangered  (Ripley,  E). 

Distribution:  San  Juan  Co.,  Utah 
(bry,  utc). 


Astragalus  cymboides  M.  E.  Jones,  Proc. 
Calif.  Acad.  Sci.  II,  5:   650.  1895. 

Type:  Huntington,  Emery  Co.,  Utah, 
16  Jun  1894,  at  5,000  feet,  M.  E.  Jones 
5464]  (pom). 

Status:  Endemic,  locally  common 
to  abundant,  neither  threatened  nor 
endangered. 

Distribution:  Carbon,  Emery, 
Grand,  and  San  Juan  counties,  Utah 
(bry). 

Astragalus  deserticus  Barneby,  Mem.  New 
York  Bot.  Gard.  13:  634.  1964. 

Type:  Common  on  slopes  near  In- 
dianola,  17  Jun  1909,  Tidestrom  2249 

(GH). 


Status:  Endemic  and  possibly  ex- 
tinct (Ripley,  PrEx). 

Distribution:  Sanpete  Co.,  Utah 
(Barneby  1964). 

Astragalus  detri talis  M.  E.  Jones,  Contr. 
W.  Bot.  13:  9.  1910. 

Type:  Ca  4  miles  above  Theodore 
[  Duchesne]  on  the  Colton  road,  Du- 
chesne Co.,  Utah,  11  May  1908,  M. 
E.  Jones  s.n.  (pom). 

Status:  Uinta  Basin  endemic,  local 
and  possibly  endangered   (Ripley,  E) . 

Distribution:  Duchesne  and  Uintah 
counties,  Utah,  and  Rio  Blanco  Co., 
Colorado   (bry). 

Astragalus  diversifolius  A.  Gray,  Proc. 
Amer.  Acad.  Arts  6:  230.  1864. 

Status:  Rare,  chsjunct  and  possibly 
extirpated  from  Utah. 

Distribution:  Juab  and  Tooele  coun- 
ties; also  in  southern  Idaho  where  rare 
(Barneby  1964). 

Astragalus  duchesnensis  M.  E.  Jones, 
Contr.  W.  Bot.  13:  9.  1910. 

Type:  Ca  1 3  miles  below  Theodore 
[Duchesne]  toward  Chepeta  Well,  Du- 
chesne Co.,  Utah,  23  May  1908,  M. 
E.  Jones  s.n.  (pom). 

Status:  Endemic,  restricted  and 
threatened  (Ripley,  T). 

Distribution:  Duchesne  and  Uintah 
counties,  Utah  (bry). 

Astragalus  eastwoodae  M.  E.  Jones,  Zoe 
4:  368.  1894,  based  on  A.  preussii  var. 
sulcatus  M.  E.  Jones,  Zoe  4:  37.  1893. 

Type:  Westwater,  Grand  Co.,  Utah 
(incorrectly  given  as  "Colorado"),  6 
May  1891^  M.  E.  Jones  s.n.    (pom). 

St.^tus:  Rare,  disjunct  and  possibly 
threatened. 

Distribution:  Emery,  Grand,  and 
San  Juan  counties,  Utah;  also  in  west 
central  Colorado    (Barneby  1964). 

Astragalus  emoryanus  (Rydb.)  Cory, 
Rhodora  38:  406.  1936,  based  on 
Hamosa  emoryana  Rydb.,  Bull.  Torrey 
Bot.  Club  54:  237.  1927. 

Status:  Rare  and  endangered. 

Distribution:  Kane  {Atwood  4629 
[bry]  )  Co.,  Utah;  Arizona,  New  Mex- 
ico, Texas,  and  Mexico  (Barneby 
1964). 

Astragalus  ensiformis  M.  E.  Jones,  Rev. 
Astrag.  226.  1923. 


348 


GREAT   BASIN   NATURALIST 


Vol.  35,  No.  4 


Status:  Rare  and  local;  threatened 
(Ripley,  T). 

Distribution:  Washington  Co., 
Utah,  and  Mohave  Co.,  Arizona  (bry). 

Astragalus  eucosmus  B.  L.  Robinson,  Rho- 
dora  10:   33.  1908. 

Status:  Possibly  extirpated. 

Distribution:  Summit  Co.  ,  Utah; 
widespread  in  northern  North  America 
(Barneby    1964;    Welsh    1974b). 

Astragalus  eurekensis  M.  E.  Jones,  Contr. 
W.  Bot.  8:   12.  1898. 

Type:  Eureka,  Juab  Co.,  Utah,  1891, 
M.  E.  Jones  s.n.  (pom). 

Status:  Endemic,  locall}'  common 
to  abundant,  neither  threatened  nor 
endangered. 

Distribution:  Juab,  Iron,  Millard, 
Sanpete,  Tooele,  and  Utah  counties, 
Utah  (bry,-  Barneby  1964). 

Astragalus  flavus  Nutt.  ex  Torr.  &  Gray 
var.  argillosus  (M.  E.  Jones)  Barneby, 
Mem.  New  York  Bot.  Card.  13:  401. 
1964,  based  on  A.  argillosus  M.  E. 
Jones,  Zoe2:  241.  1891. 

Type:  Green  River,  Emery  Co.,  Utah, 
7  May  1890,  M.  E.  Jones  s.n.  (pom). 

Status:  Endemic,  restricted  edaph- 
ically  but  locally  common,  neither 
threatened  nor  endangered. 

Distribution:  Emer\ ,  Garfield, 
Grand,  and  Wayne  counties,  Utah 
(bry). 

Astragalus  gilviflorus  Sheldon,  Minne- 
sota Bot.  Stud.  1:  21.  1894,  based  on 
A.  triphyllus  Pursh,  Fl.  Amer.  Sept.  2: 
740.  1814,  not  Pallas. 

Status:  Rare  or  possibly  extirpated 
from  Utah. 

Distribution:  Summit  Co.,  Utah 
(uTc);  widespread  east  of  the  conti- 
nental  divide    (Barneby    1964). 

Astragalus  hallii  A.  Grav  var.  fallax  (S. 
Wats.)  Barneby,  Leaff.  W.  Bot.  9:  91. 
1960,  based  on  A.  fallax  S.  Wats.,  Proc. 
Amer.  Acad.  Arts  20:    362.  1885. 

Status:  Rare  and  local,  possibly 
threatened. 

Distribution:  Garfield  and  Kane 
counties,  Utah  (bry). 

Astragalus  hamiltonii  C.  L.  Porter,  Rho- 
dora  54:  159.  1952. 

Type:  On  the  Wasatch  formation  5 
miles    south    of    Vernal,    Uintah    Co., 


Utah,  24  May  1950,  Hamilton  &■  Beath 
s.n.   (rm). 

Status:  Endemic,  rare  and  local,  en- 
dangered (Ripley,  E). 

Distribution:  Uintah  Co.,  Utah 
(bry). 

Astragalus  harrisonii  Barneby,  Mem.  New 
York  Bot.  Card.  13:  271.  1964. 

Type:  Wash  below  the  Natural 
Bridge,  near  Fruita,  Wayne  Co.,  Utah, 
8  Jun  1961,  Barneby  15131  (cas). 

Status:  Endemic,  rare  and  endan- 
gered (Ripley,  E). 

Distribution:  Wa3'ne  Co.,  Utah; 
known  only  from  the  type  area  (bry)  . 

Astragalus  iodanthus  S.  Wats,  in  King, 
Rep.  Geol.  Explor.  40th  Parallel  5:  70. 
1871. 

Status:  Rare  and  possibly  extirpated 
from  Utah. 

Distribution:  Tooele  Co.,  Utah; 
widespread  to  the  west  of  Utah  (Bar- 
neby 1964). 

Astragalus  iselyi  Welsh,  Great  Basin  Nat. 
34:  305.  1974. 

Type:  Brumley  Bridge,  ca  1.5  miles 
north  of  Pack  Creek  Ranch,  San  Juan 
Co.,  Utah,  5  May  1971,  Welsh  10970 
(bry). 

Status:  Endemic,  edaphically  re- 
stricted, endangered. 

Distribution:  Grand  and  San  Juan 
counties,  Utah   (bry,-  Welsh  1974a). 

Astragalus  jejunus  S.  Wats,  in  King,  Rep. 
Geol.  Explor.  40th  Parallel  5:  73.  1871. 

Status:  Rare  and  restricted,  possibly 
threatened. 

Distribution:  Rich  Co.,  Utah  (Bar- 
neby 1964);  southwestern  Wyoming 
and  in  an  isolated  location  in  White 
Pine  Co.,  Nevada. 

Astragalus  kentrophyta  A.  Gra}  var. 
coloradoensis  M.  E.  Jones,  Contr.  W. 
Bot.  10:  63.  1902. 

Status:  Navajo  Basin  endemic;  rare 
to  locally  common  but  probably  threat- 
ened . 

Distribution:  Garfield,  Kane,  San 
Juan,  and  Wayne  counties,  Utah  (bry), 
and  adjacent  northern  Arizona  (Bar- 
neby 1964). 

Astragalus  lance  arius  A.  Gra>-,  Proc. 
Amer.  Acad.  Arts  13:   370.  1878. 

Status:  Rare  and  local  in  generalized 


Dec.  1975 


WELSH,   ET   AL.:    ENDANGERED    UTAH    PLANTS 


349 


habitats;    possibly   threatened    (Ripley, 
T). 

Distribution:  Kane  (bry)  and 
Washington  counties,  Utah  and  in  Co- 
conino and  MohaA'e  counties,  Arizona 
(bry,-    Barneby    1964). 

Astragalus  lentiginosus  Dougl.  ex  Hook, 
var.  chartaceous  M.  E.  Jones,  Proc. 
Calif.  Acad.  Sci.  II,  5:  673.  1895. 

Type:  Ephraim,  Sanpete  Co.,  Utah, 
13  Jul  1894,  6,000  feet,  M.  E.  Jones 
5627m  (pom). 

Status:  Rare  and  local,  possibly 
threatened   (Ripley,  PoEx,  Tax?).^ 

Distribution:  Daggett,  Juab,  San- 
pete, Sevier,  Summit,  and  Tooele  coun- 
ties, Utah,  and  widespread  from  Colo- 
rado, Wyoming,  Idaho,  Oregon,  Cali- 
fornia,  and  Nevada    (Schoener   1975). 

Astragalus  lentiginosus  Dougl.  ex  Hook, 
var.  fremontii  (Gray  ex  Torr.)  S.  Wats, 
in  King,  Rep.  Geol.  Explor.  40th  Paral- 
lel 5:  66.  1871,  based  on  A.  fremontii 
Gray  ex  Torr.  in  Whipple,  Explor.  & 
Surv.  Railroad  Route  from  Mississippi 
River  to  Pacific  Ocean  4(5):  80.  1857. 

Status:  Rare  and  ephemeral,  pos- 
sibly threatened. 

Distribution:  Washington  Co.,  Utah 
(bry,-  Schoener  1974);  widespread  in 
southern  Nevada  and  adjacent  Cali- 
fornia (Barneby  1964). 

Astragalus  lentiginosus  Dougl.  ex  Hook, 
var.  scorpionis  M.  E.  Jones,  Rev.  Astrag. 
124.  1923. 

Status:  Rare  and  local;  possibly 
threatened. 

Distribution:  Juab  Co.,  Utah 
(Schoener  1975),  and  Nevada  (Barne- 
by 1964). 

Astragalus  lentiginosus  Dougl.  ex  Hook, 
var.  stramineus  (Rydb.)  Barneby,  Leafl. 
W.  Bot.  4:  122.  1945,  based  on  Cystium 
stramineum  Rydb.,  N.  Amer.  Fl.  24: 
409.  1929. 

Type:  Southern  Utah,  1870,  Palmer 
s.n.  (ny). 

Status:  Locally  common,  restricted, 
neither  threatened  nor  endangered. 

Distribution:  Washington  Co.,  Utah 


(bry),     Mohave     Co.,     Arizona,     and 
Clark  Co.,  Nevada   (Schoener  1975). 

Astragalus  lentiginosus  Dougl.  ex  Hook, 
var.  ursinus  (A.  Gray)  Barneby,  Leafl. 
W.  Bot.  4:  133.  1945,  based  on  A. 
ursinus  A.  Gray,  Proc.  Amer.  Acad.  Art 
13:  367.  1878. 

Type:  Bear  Valley,  in  south  central 
Utah,  1877,  Palmer  s.n.  (gh). 

Status:  Endemic,  possibly  extinct. 

Distribution:  Iron  (or  possibly  Se- 
vier) Co.,  Utah;  perhaps  the  specimens 
are  mislabeled." 

Astragalus  limnocharis  Barneby,  Leafl. 
W.  Bot.  4:  236.  1946. 

Type:  Gravelly  beach  of  Navajo 
Lake,  at  Spruce  Forest  Camp,  Iron  Co., 
Utah,  13  Jul  1940,  Maguire  19474 
(ny). 

Status:  Endemic,  rare  and  restricted; 
threatened. 

Distribution:  Iron  and  Kane  (bry) 
counties,   Utah    (Barneby   1964). 

Astragalus  loanus  Barneby,  Mem.  New 
York  Bot.  Gard.  13:  661.  1964,  based 
on  A.  newberryi  var.  wardianus  Bar- 
neby, Amer.  Midi.  Naturalist  37:  481. 
1947. 

Type:  Canyon  east  of  Glen  wood,  Se- 
vier Co.,  Utah,  1875,  Ward  223   (gh). 

Status:  Endemic,  rare  and  threat- 
ened (Ripley,  E) . 

Distribution:  Garfield,  Piute,  Sevier 
and  Wayne  counties,  Utah  (bry). 

Astragalus  lutosus  M.  E.  Jones,  Contr.  W. 
Bot.  13:  7.  1910. 

Type:  White  River,  Uintah  Co.,  Utah, 
25  May  1958,  5,200  feet,  M.  E.  Jones 
s.n.  (pom). 

Status:  Uinta  Basin  endemic,  rare 
and  edaphically  restricted;  endangered 
(Ripley,  E). 

Distribution:  Uintah  Co.,  Utah,  and 
Rio  Blanco  Co.  Colorado  (Barneby 
1964). 

Astragalus  malacoides  Barneby,  Mem. 
New  York  Bot.  Gard.  13:  500.  1964. 

Type:  Northeast  slope  of  Kaiparowits 
Plateau,  south  of  Willow  Tank,  Kane 
Co.,  Utah,  9  May  1939,  fJarrison  9069 
(us). 


'Schooner  (1975)  has  deteirmined  that  this  name  includes  var.  platyphrllidius  (Rydb.)  M.  E.  Peck,  and  therefore  de- 
fines   the    taxon    differently    from    Ripley     (1975). 

"This  variety  was  reduced  to  sj-nonjnny  luider  var.  palans  (M.  E.  .Tones)  M  E.  Jones  bj-  Schoener  (1975),  but  that 
taxon  is  currentlj'  known  only  from  the  valleys  of  the  Colorado  and  Virgin  rivers.  The  exaction  location  of  "Bear  Valley" 
has   not    been   determined    (McVaugh    1956). 


350 


GREAT  BASIN    NATURALIST 


Vol.  35,  No.  4 


Status:  Endemic,  restricted  though 
locallv  common;  threatened  (Riplev, 
E).    "  '  ,  ^ 

Distribution:  Kane  Co.,  Utah  (bry). 

Astragalus  minthorniae  (Rydb.)  Jeps.  var. 
gracilior  (Barneby)  Barneby,  Amer. 
Midi.  Naturalist  55:  493.  1956,  based 
on  A.  ensiformis  var.  gracilior  Barne- 
by Proc.  Calif.  Acad.  Sci.  IV.  25:  158. 
1944. 

Type:  Ca  5  miles  south  of  Veyo, 
Washington  Co.,  Utah,  4,900  feet,  Rip- 
ley &  Barneby  4951   (cas). 

Status:  Endemic  (?),  locally  com- 
mon in  disjunct  populations,  possibly 
threatened  (Ripley,  E). 

Distribution:  Washington  Co., 
Utah  (bry),  and  possibly  in  Mohave 
Co.,  Arizona. 

Astragalus  monumentalis  Barneb^'.  Leafl. 
W.  Bot.  7:  35.  1953. 

Type:  White  Canyon,  25  miles  south- 
east of  Hite,  San  Juan  Co.,  Utah,  18 
May  1950.  Harrison  11595  (us). 

Status:  Endemic,  rare  and  restricted 
edaphically,  possibly  threatened. 

Distribution:  Garfield  and  San  Juan 
counties,  Utah    (bry)  . 

Astragalus  musiniensis  M.  E.  Jones,  Proc. 
Calif.  Acad.  Sci.  II,  5:  671.  1895. 

Type:  San  Rafael  Swell.  Emery  (?) 
Co.,  Utah,  12  May  1914,  \1.  E.  Jones 
s.n.  (pom). 

Status:  Endemic,  rare  to  common, 
neither   threatened    nor   endangered. 

Distribution:  Carbon.  Emery,  Gar- 
field, Grand,  and  Wayne  counties, 
Utah  (bry). 

Astragalus  nidularius  Barnebv.  Leafl.  W. 
Bot.  8:  16.  1956. 

Type:  Near  the  head  of  White  Can- 
yon, 2  miles  below  the  Kachina  Bridge 
of  Natural  Bridges  N.M..  Barneby 
12778  (cas). 

Status:  Endemic,  restricted  and  lo- 
cal, disjunct  in  several  main  locations, 
neither  threatened  nor  endangered. 
(Ripley,  T) . 

Distribution:  Ciarfield,  San  Juan, 
and  Wayne  counties,  Utah   (bry). 

Astragalus  oophorus  S.  Wats.  var.  loncho- 
caly.r  Barneby.  Leafl.  W.  Bot.  7:  194. 
1954. 


Status:  Rare  and  threatened  (Rip- 
ley,T). 

Distribution:  Iron  Co.,  Utah  (bry), 
and  adjacent  Nevada   (Barneby  1964). 

Astragalus  pardalinus  (Rydb.)  Barneby, 
Mem.  New  York  Bot.  Gard.  13:  883. 
1964,  based  on  Phaca  pardalina  Rvdb.. 
N.  Amer.  Fl.  24:  352.  1929. 

Type:  Cedar  Mtn.,  Emery  Co.,  Utah, 
20  May  1915,  M.  E.  Jones  s.n.    (ny). 

Status:  Endemic,  restricted  though 
locally  common,  neither  threatened  nor 
endangered  (Ripley,  E). 

Distribution:  Emery,  Garfield,  and 
Wayne  counties,  Utah  (bry). 

Astragalus  perianus  Barneby,  Mem.  New 
York  Bot.  Gard.  13:  973.  1964. 

Type:  Mountains  north  of  Bullion 
Creek,  near  Marvsvale,  Piute  Co.,  Utah, 
23  Jul  1905,  Rydberg  &  Carlton  7104 
(ny). 

Status:  Endemic  and  threatened 
(Ripley,  PoEx). 

Distribution:  Piute  (Barneby  1964) 
and  Garfield    (bry)    counties.  Utah." 

Astragalus  pinonis  M.  E.  Jones,  Contr.  W. 
Bot.  8:   14.  1898. 

Type:  Frisco,  Beaver  Co.,  Utah,  22 
Jun  1880,  ca  8,000  feet.  M.  E.  Jones 
s.n.  (pom). 

Status:  Rare  and  possibly  extirpated. 

Distribution:  Beaver  and  Juab 
counties.  Utah,  and  in  east  central  Ne- 
vada (Barneby  1964). 

Astragalus  platytropis  A.  Gray.  Proc. 
Amer.  Acad.  Arts  6:  526.  1865. 

Status:  Rare  and  local,  both  edaph- 
ically and  altitudinally  restricted. 

Distribution:  Western  Beaver 
(bry)  and  Tooele  counties.  Utah,  and 
much  of  Nevada  and  adjacent  Cali- 
fornia (Barneby  1964). 

Astragalus  rafaelensis  M.  E.  Jones,  Rev. 
Astrag.  146.  1923. 

Type:  Near  Cedar  Mtn.,  Emery  Co., 
Utah,  19  May  1915,  M.  E.  Jones  s.n. 
(pom). 

Status:  Endemic,  restricted  edaph- 
ically, threatened  (Ripley,  T). 

Distribution:  San  Rafael  Swell, 
Emery  Co.,  Utah  (bry).. 


"This   species,   previously  known   only   from   the   type  coUeclioii.    was   rediscovered    in    1073;    one  of   many   species   though 
be  extinct  but  found  diunng  1975  as  a  result  of  the  interest  generated    by    the    Smithsonian    Report. 


Dec.  1975 


WELSH,   ET   AL.:    ENDANGERED    UTAH    PLANTS 


351 


Astragalus  sabulosus  M.  E.  Jones,  Zoe  2: 
239.  1891. 

Type:  Cisco,  Grand  Co.,  Utah,  2  May 
1890,  M.  E.  Jones  s.n.   (pom). 

Status:  Endemic,  rare  and  edaph- 
ically  restricted;   threatened. 

Distribution:  Grand  Co.,  Utah  (bry; 
Welsh  1974a). 

Astragalus  saurinus  Barneby,  Leafl.  W. 
Bot.  8:   17.  1956. 

Type:  Dinosaur  N.M.,  6  miles  north 
of  Jensen,  Uintah  Co.,  Utah,  26  Jun 
1953,  Holmgren  &  Tillett  9527    (ny). 

Status:  Endemic,  rare  and  edaph- 
ically  restricted;  threatened  (Ripley, 
E). 

Distribution:  Uintah  Co.,  Utah 
(bry.  utc). 

Astragalus  serpens  M.  E.  Jones.  Proc. 
Calif.  Acad.  Sci.  II,  5:  641.  1895. 

Type:  Loa  Pass,  Wayne  Co.,  Utah, 
17  Jul  1894,  M.  E.  Jones  56391  (pom). 

Status:  Endemic,  local  and  period- 
ically abundant  in  disjunct  populations, 
neither  threatened  nor  endangered 
(Ripley,  E). 

Distribution:  Garfield,  Piute  and 
Wayne  counties,  Utah   (bry). 

Astragalus  stocksii  Welsh,  Great  Basin 
Nat.  34:  307.  1974. 

Type:  Henry  Mts.,  Penellen  Pass, 
Garfield  Co.,  Utah,  30  May  1972, 
Welsh  11740  (bry). 

Status:  Endemic,  rare  and  local; 
threatened. 

Distribution:  Garfield  Co.,  Utah; 
known  only  from  the  type  area  (bry). 

Astragalus  striatiflorus  M.  E.  Jones,  Proc. 
Calif.  Acad.  Sci.  II,  5:   643.  1895. 

Type:  Above  Springdale,  Washing- 
ton Co.,  Utah,  25  Sep  1894,  4,000  feet, 
M.  E.  Jones  6080k  (pom). 

Status:  Rare  and  edaphically  re- 
stricted;   endangered    (Ripley,   T). 

Distribution:  Kane  and  Washing- 
ton counties,  Utah,  and  Coconino  Co., 
Arizona   (bry)  . 

A  main  locality  of  distribution  for 
this  plant  is  the  Coral  Pink  Dunes  region 
where  the  plants  grow  in  the  interdune 
valleys.  The  use  of  that  region  for  rec- 
reation possibly  spells  the  doom  of  this 
plant   in   that   section. 

Astragalus  tephrodes  A.  Gray  var.  brachy- 
lobus  (A.  Gray)  Barneby,  Amer.  Midi. 


Naturalist  M -.  466.  1947,  based  on  A. 
shortianus  var.  brachylobus  A.  Gray, 
Proc.  Amer.  Acad.  Arts  13:  367.  1878. 

Status:  Rare  and  possibly  extirpated 
from  Utah. 

Distribution:  Washington  Co.,  Utah 
(Barneby  1964). 

Astragalus  tctrapterus  A.  Gray,  Proc. 
Amer.  Acad.  Arts  13:  369.  1878. 

Type:  Ca  25  miles  north  of  St. 
George,  Washington  Co.,  Utah,  1877, 
Palmer  111  (gh). 

Status:  Restricted  but  locally  com- 
mon to  abundant,  possibly  threatened. 

Distribution:  Beaver,  Iron,  Kane, 
and  Washington  counties,  Utah  (bry)  ; 
also  in  northwestern  Arizona  (bry), 
Nevada,   and  Oregon    (Barneby   1964). 

Astragalus  wardii  A.  Gray,  Proc.  Amer. 
Acad.  Arts  12:   55.  1876. 

Type:  On  the  edge  of  the  Aquarius 
Plateau,  Sevier  Co.,  Utah,  1875,  8,700 
feet.  Ward  424  (gh). 

Status:  Endemic,  local  and  disjunct, 
neither  threatened  nor  endangered. 

Distribution:  Garfield,  Kane,  Piute, 
and  Sevier  counties,  Utah   (bry). 

Astragalus  wetherillii  M.  E.  Jones,  Zoe 
4:  34.  1893. 

Status:  Possible  extirpated  from 
Utah  (Ripley,  T). 

Distribution:  Grand  Co.,  Utah,  and 
west  central  Colorado  (Barneby  1964). 

Astragalus  woodruffii  M.  E.  Jones,  Rev. 
Astrag.  77.  1923. 

Type:  On  the  sandy  foot  of  the  San 
Rafael  Swell,  Emery  (?)  Co.,  Utah, 
17-18  May  1914,  M.  E.  Jones  s.n. 
(pom). 

Status:  Endemic,  Local  and  edaph- 
ically  restricted,   possibly   threatened. 

Distribution:  Emery,  Wayne,  and 
possibly  Garfield  counties,  Utah  (bry). 

Dalea  epica  Welsh,  Great  Basin  Nat.  31: 
90.  1971. 

Type:  Ca  10  miles  east  of  Halls 
Crossing,  San  Juan  Co.,  Utah,  30  Apr 
1966,  Welsh  5205   (bry). 

Status:  Endemic,  restricted  and  rare, 
and  possibly  threatened. 

Distribution:  San  Juan  Co.,  Utah 
(bry);  known  only  from  the  type  lo- 
cality. 


352 


GREAT   BASIN    NATURALIST 


Vol.  35,  No.  4 


Hedysarum  boreale  Nutt.  var.  gremiale 
(Rollins)  Northstrom  &  Welsh,  Great 
Basin  Nat.  30:  125.  1970,  based  on  H. 
gremiale  Rollins,  Rhodora  42;  230. 
1940. 

Type:  West  of  Vernal,  Uintah  Co., 
Utah,  16  Jun  1937,  Rollins  1735  (gh). 

Status:  Endemic,  local  and  restric- 
ted, possibly  threatened. 

Distribution:  Duchesne  and  Uintah 
counties,  Utah  (bry). 

Hoffmanseggia  repens  (Eastw.)  Cocker- 
ell,  Muhlenbergia  4:  68.  1908,  based  on 
Caesalpinia  repens  Eastw.,  Zoe  4:  116. 
1893. 

Type:  Court  House  Wash,  near 
where  it  comes  into  the  Grand  [Colo- 
rado] River,  near  Moab,  Grand  Co., 
Utah,    26    May    1892,    Eastwood    s.n. 

(CAs). 

Status:  Endemic,  edaphically  re- 
stricted but  locally  common,  neither 
threatened  nor  endangered,  but  appar- 
ently extirpated  from  the  type  area 
(Harrison  et  al.  1964). 

Distribution:  Emery  (bry),  Grand, 
and  Wayne  (bry)   counties,  Utah. 

Lathyrus  brachy calyx  Rydb.  ssp.  zionis 
(C.  L.  Hitchc.)  Welsh,  Proc.  Utah 
Acad.  Sci.  52:  217.  1966,  based  on  L. 
zionis  C.  L.  Hitchc,  Univ.  Wash.  Publ. 
Biol.  15:  36.  1952. 

Type:  Ten  miles  east  of  the  east  en- 
trance of  Zion  N.P.,  Kane  (?)  Co., 
Utah,  30  May  1949,  Hitchcock  19013 

(WTU). 

Status:  Rare  to  locally  abundant  in 
disjunct  populations,  neither  threatened 
nor  endangered. 

Distribution:  Grand,  Kane,  San 
Juan,  and  Washington  counties,  Utah, 
and  Coconino  Co.,  Arizona  (bry)  ;  re- 
portedly more  widespread  in  Arizona 
(McDougall  1973)   . 

Lotus  longebracteatus  Rydb.,  Bull.  Tor- 
rey  Bot.  Club  30:  254.  1903. 

Type:  Southern  Utah,  possibly  Wash- 
ington Co.,  Utah,  1877,  Palmer  94 
(us). 

Status:  Endemic  (?),  restricted  but 
locally  abundant,  neither  threatened 
nor  endangered. 

Distribution:  Kane  and  Washing- 
ton counties,  Utah  (bry,-  Ottley  1944). 


Lupinus  jonesii  Rydb.,  Bull.  Torrey  Bot. 
Club  30:   256.   i903. 

Type:  Silver  Reef,  Washington  Co., 
Utah,  3  May  1894,  M.  E.  Jones  5143 
(us). 

Status:  Endemic,  rare  and  threat- 
ened. 

Distribution:  Washington  Co.,  Utah 
(bry). 

Lupinus  marianus  Rj-^db.,  Bull.  Torrey 
Bot.  Club  34:  41.  1907. 

Type:  Along  Bullion  Creek  above 
Marysvale,  Piute  Co.,  Utah,  21  Jul 
1905,  Rydberg  &  Carlton  7024  (ny). 

Status:  Endemic,  restricted  and 
threatened  (Ripley,  T). 

Distribution:  Piute  Co.,  Utah 
(bry). 

Oxytropis  jonesii  Barneby,  Proc.  Calif. 
Acad.  Sci.  IV,  27:  215.  1952. 

Type:  Red  Canyon,  Garfield  Co., 
Utah,  7  Jun  1947,  7,150  feet,  Ripley  & 
Barneby  8550  (gas). 

Status:  Endemic,  edaphically  re- 
stricted and  threatened. 

Distribution:  Emery  (us),  Garfield 
(bry),  Iron  (bry,  us)  and  Uintah 
(bry,  NY,  UTc)  counties,  Utah. 

Oxytropis  multiceps  Torr.  &  Gray,  Syn. 
Fl.  N.  Amer.  1:   341.  1838. 

Status:  Rare  and  local,  possibly  en- 
dangered. 

Distribution:  Daggett  Co.,  Utah, 
and  in  Wyoming  and  Colorado  (Barne- 
by 1952).' 

Oxytropis  obnapiformis  C.  L.  Porter,  Ma- 
drono 9:  133.  1947. 

Status:  Rare  and  local,  possibly 
threatened  (Ripley,  E). 

Distribution:  Daggett  Co.,  Utah; 
also  in  Wyoming  and  Colorado  (bry). 

Peter ia  thompsonae  S.  Wats.,  Amer.  Nat- 
uralist 7:   300.  1873. 

Type:  Kanab,  Kane  Co.,  Utah,  1872, 
Thompson  s.n.  (gh). 

St.\tus:  Restricted  in  disjunct  pop- 
ulations, neither  threatened  nor  endan- 
gered (Ripley,  T). 

Distribution:  Emery,  Grand,  Kane, 
San  Juan,  and  Washington  counties, 
Utah;  Mohave  Co.,  Arizona,  Nevada, 
and  Idaho  (bry,-  Porter  1956). 

Psoralea  aromatica  Pavson,  Bot.  Gaz.  60: 
379.  1915. 


Dec.  1975 


WELSH.    ET    AL.:     ENDANGERED    UTAH    PLANTS 


353 


Status:  Edaphically  restricted,  local 
and  rare;  possibly  threatened. 

Distribution:  Emery  and  Grand 
counties,  Utah,  and  Montrose  Co., 
Colorado  (bry;  Ockendon  1965). 

Psoralea  castorea  S.  Wats.,  Proc.  Amer. 
Acad.  Arts  14:  291.  1879. 

Type:  Near  Beaver  City,  Beayer  Co., 
Utah,  1877,  Palmer  96  (gh).'^ 

Status:  Unknown  from  Utah  in 
contemporary  collection,  although  a 
specimen  without  collector  or  date 
labeled  P.  castoria  is  on  file  at  ut;  either 
extirpated  from  the  state,  or,  and  more 
likely,  never  from  Utah. 

Distribution:  Arizona  and  adjacent 
southern  California  (Kearney  &  Peebles 
1951;  Munz  &Keck  1959).' 

Psoralea  epipsila  Barneby,  Leafl.  W.  Bot. 
3:    193.   1943. 

Type:  Ca  17  miles  east  of  Kanab  to- 
ward Jepson  Springs,  Kane  Co.,  Utah, 
6  Jun  1942,  Ripley  &  Barneby  4832 
(gas). 

Status:  Mohave  corridor  endemic, 
rare  and  endangered   (Ripley,  T). 

Distribution:  Kane  Co.,  Utah,  and 
adjacent  Coconino  Co.,  Arizona  (Bar- 
neby 1943). 

Psoralea  juncea  Eastw.,  Proc.  Calif.  Acad. 
Sci.  II,  6:  286.  1896. 

Type:  Epsom  Creek,  San  Juan  Co., 
Utah,  13  Jul  1895,  Eastwood  21   (gas). 

Status:  Navajo  Basin  endemic, 
edaphically  restricted  but  abundant  to 
common,  neither  threatened  nor  endan- 
gered. 

Distribution:  Garfield,  Kane,  and 
San  Juan  counties,  Utah  (bry-),  and 
Coconino  Co.,  Arizona. 

Psoralea  lanceolata  Pursh  var.  steno- 
phylla  (Rydb.)  Toft  &  Welsh,  Great 
Basin  Nat.  32:  85.  1972,  based  on  P. 
stenophylla  Rydb.,  Bull.  Torrey  Bot. 
Club  40:  46.  1913. 

Type:  Proposed  dam  site  near  Wil- 
son Mesa,  Grand  Co.,  Utah,  1  Jul  1911, 
Rydberg  &  Garrett  8367  (ny). 

Status:  Endemic,  locally  common 
and  neither  threatened  nor  endangered. 

Distribution:  Garfield,  Grand, 
Kane,  San  Juan  and  Wayne  counties, 
Utah  (Toft  &  Welsh  1972). 

'It  seems  likely  that   Palmer  mislabled  the  type  material,   a 
Arizona.    The   species   is   not   known    from   Utah. 


Psoralea  pariensis  Welsh  &  Atwood,  spec, 
nov.  P.  megalanthae  Wooton  &  Stand- 
ley  proxime  affinis  sed  floribus  brevior- 
ibus  et  venas  supra  valde  albo-strigosas. 

Plantae  habentes  caudices  bene-evo- 
luti  et  radices  tuberosae,  2-8  cm  altae; 
caules  0.5-3  (4.5)  cm  longi,  internodiis 
1-3  elongatis,  strigosi;  stipulae  4-7  (10) 
mm  longae,  ovatae  vel  obovatae,  strigosae; 
foliola  3-5,  9-23  (25)  mm  longa,  7-22 
mm  lata,  obovata  vel  orbicularia,  cuneata, 
rotundata  ad  truncata  vel  emarginata 
apicaliter,  utrinque  glandulifera;  petioli 
1.3-6.3  (7)  cm  longi,  strigosi,  pilis  ap- 
pressis  vel  adscendentibus;  pedunculi  0.5- 
2.2  (2.8)  cm  longi,  pilis  appressis  vel  ad- 
scendentibus; bracteae  4-6  (8)  mm  longae, 
ovatae  vel  obovatae,  acuminatae  abrupte, 
pilosae  rigide;  pedicelli  (1)  2-3.8  mm 
longi,  pilis  adscendentibus;  calyces  plus 
minusve  gibbosi  basim,  tubus  3.3-4  (4.6) 
mm  longus,  dentes  inaequales,  infimis  5.3- 
6.8  mm  longis,  circa  duplo  latioribus  quam 
lateralibus;  corolla  8.8-10.5  (12)  mm  lon- 
ga, dente  infimo  calycis  leviter  longiori- 
bus;  alae  vexillo  subaequalis,  purpureo- 
maculatae  ad  apicem;  fructus  usque  ad  9 
mm  longum  et  seminum  usque  ad  5.2 
mm  longum. 

Type:  UTAH:  Garfield  Co.:  Bryce 
Canyon  National  Park,  in  ponderosa 
pine  woods,  as  ground  layer,  at  ca 
8,000  feet  elevation,  26  Jun  1975, 
Welsh  &  Murdoch  12859.  Holotype, 
bry,  Isotypes  to  be  distributed. 

Additional  spegimens  examined: 
UTAH:  Garfield  Co.:  East  Creek,  3 
miles  south  of  Inspiration  Point,  Bryce 
Canyon  N.P.,  in  black  sagebrush  area, 
11  Jun  1970,  Buchanan  1494  (bry, 
wsGo);  Paria  View,  Bryce  Canyon 
N.P.,  in  open  ponderosa  pine  woods,  ca 
8,000  feet  elevation,  12  Jun  1975,  S. 
L.  &  S.  L.  Welsh  12810  (bry);  East 
Creek,  9  Jun  1931,  Weight  B-31/ 6-305 
(bcnp,  us).  Kane  Co.:  Hackberry  Can- 
yon in  Cottonwood  Wash,  ca  10  miles 
north  of  U.S.  Highway  89,  sec.  31, 
T.  40S.,  R.  IW.,  26  Apr  1972,  Atwood 
3684  (bry). 

Status:  Endemic,  rare  and  threat- 
ened. 

Distribution:  Kane  and  Garfield 
counties,  Utah. 

I    tliat    it    was    in    leality    from     "Beaverdam."    Mohave    Co.. 


354 


GREAT   BASIN   NATURALIST 


Vol.  35,  No.  4 


Parryella  filifolia  Torr.  &  Gray  in  Gray, 
Proc.  Amer.  Acad.  Arts  7:  397.  1868. 

Status:  Restricted  and  rare;  threat- 
ened. 

Distribution:  Grand  and  San  Juan 
counties,  Utah  (bry);  also  in  northern 
Arizona  and  New  Mexico. 

Psorothamnus  thompsonae  (Vail)  Welsh 
&  Atwood,  comb,  nov.,  based  on  Paro- 
sela  thompsonae  Vail,  Bull.  Torrey  Bot. 
Club  24:  18.  1897. 

Type:  Kanab,  Kane  Co.,  Utah,  or  pos- 
sibly Mohave  Co.,  Arizona,  1872, 
Thompson  s.n."  (ny). 

Status:  Endemic  (?);  restricted  but 
locally  common  and  neither  threatened 
nor  endangered. 

Distribution:  Emery,  Garfield, 
Kane,  San  Juan,  and  Wayne  counties, 
Utah  (bry  and  personal  observations). 

Robinia  neomexicana  A.  Gray,  Mem. 
Amer.  Acad.  Arts  II,  5:  314.  1855. 

Status:  Restricted  and  rare,  possibly 
threatened. 

Distribution:  Garfield  (ut)  and 
Washington  counties,  Utah  (bry)  ; 
widespread  elsewhere. 

Gentianaceae 

Gentianella  tortuosa  (M.  E.  Jones)  Gil- 
lett,  Ann.  Missouri  Bot.  Gard.  44:  248. 
1957,  based  on  Gentiana  tortuosa  M. 
E.  Jones,  Proc.  Calif.  Acad.  Sci.  II,  5: 
707.  1895. 

Type:  Panguitch  Lake,  Garfield  Co., 
Utah,  7  Sep  1894,  8,400  feet,  M.  E. 
Jones  6008  (pom). 

Status:  Restricted  but  locally  com- 
mon, neither  threatened  nor  endan- 
gered. 

Distribution:  Garfield,  Iron,  Kane, 
Sanpete  counties,  Utah  (bry),  and 
Clark  Co.,  Nevada  (Gillett  1957). 

Geraniaceae 

Geranium  marginale  Rydb.  ex  Hanks  & 
Small,  N.  Amer.  Fl.  25:  16.  1907. 

Type:  Aquarius  Plateau  at  the  head 
of  Poison  Creek,  Garfield  Co.,  Utah,  4 
Aug  1905,  Rydberg  &  Carlton  7401 
(ny). 


Status:  Endemic,  restricted  and 
possibly  threatened  (Ripley,  T). 

Distribution:  Garfield,  Sevier,  and 
Wayne  counties,  Utah  (Nebeker  1974). 

Hydrophyllaceae 

Hydrophyllum  fendleri  (A.  Gray)  A.  A. 
Heller,'  PI.  World  1:  23.  1897,  based 
on  H.  occidentale  var.  fendleri  A.  Gray, 
Proc.  Amer.  Acad.  Arts  10:  314.  1875. 

Status:  Restricted  and  rare,  possibly 
threatened. 

Distribution:  San  Juan  Co.,  Utah 
(uT.  UTc);  widespread  from  Washing- 
ton and  Oregon  to  Wyoming,  Colorado 
and  New  Mexico,  with  var.  fendleri 
restricted  to  the  Rocky  Mountain  area 
(Constance  1942). 

Nama  retrorsum  J.  T.  Howell,  Leafl.  W. 
Bot.  5:   149.  1949. 

Status:  Restricted  and  local,  possibly 
threatened   (Ripley,  T) . 

Distribution:  Garfield,  Grand,  and 
Kane  counties,  Utah  (bry),  and  north- 
eastern Arizona    (Atwood,  in  press). 

Nemophila  parviflora  Dougl.  ex  Benth. 
var.  austinae  (Eastw.)  A.  Brand, 
Pflanzenr.  IV  (Heft  251):  55.  1913, 
based  on  A^.  austinae  Eastw.,  Bull.  Tor- 
rey Bot.  Club  28:  143.  1901. 

Status:  Restricted  and  local,  possibly 
threatened. 

Distribution:  Weber  Co.,  Utah; 
widespread  in  Idaho,  Nevada,  Califor- 
nia, Oregon,  and  Washington  (Atwood, 
in  press) . 

Phacelia  anelsonii  J.  F.  Macbride,  Contr. 
Gray  Herb.  49:  26.  1917. 

Status:  Restricted  and  local;  threat- 
ened  (Ripley,  T) . 

Distribution:  Washington  Co.,  Utah 
(bry)  ;  southern  Nevada  and  adjacent 
California   (Atwood,  in  press). 

Phacelia  argillacea  Atwood,  Phvtologia 
26:  437.  1973. 

Type:  Clear  Creek,  in  Spanish  Fork 
Canyon  along  the  railroad  on  the  east 
side  of  U.S.  Highway  50-6,  Utah  Co., 
Utah,  Atwood  et  al.  309i   (bry). 

Status:  Endemic,  rare  and  endan- 
gered   (Ripley,  E). 

'It  is  likely  that  sonic  of  llic  collections  sent  to  (iiay  at  Haivaid  University  iwho,  in  turn,  i^avd  tlicm  to  \\ntson)  niay 
not  liavc  been  collected  by  Mrs.  Thompson,  hut  others  with  John  Wesley  Powell.  It  is  known  that  l)otii  Powell  and  Bishoj) 
collected  plants  and  gave  them  to  Ellen  Thompson  who  did  not  collect  outside  of  the  Kanab  area  of  Kane  Co.,  Utah  (Cron- 
quist    ct    al.     1972). 


Dec.  1975 


WELSH,   ET   AL.:    ENDANGERED    UTAH    PEANTS 


355 


Distribution:  Utah  Co.,  Utah  (At- 
wood  1973,  in  press). 

Phacelia  cephalotes  A.  Gray,  Proc.  Amer. 
Acad.  Arts  10:  325.  1875. 

Type:  Valley  of  the  Virgin  River, 
Washington  Co.,  Utah,  May  1874, 
Parry  179  (gh). 

Status:  Restricted  and  local,  pos- 
sibly threatened   (Ripley,  T). 

Distribution:  Kane  and  Washing- 
ton counties,  Utah,  and  Mohave  and 
Navajo  counties,  Arizona  (Atwood,  in 
press). 

Phacelia  constancei  Atwood,  Rhodora  74: 
451.  1972. 

Status:  Edaphically  restricted  and 
local;  threatened  (Ripley,  T). 

Distribution:  Emery,  Garfield, 
Kane,  and  San  Juan  counties,  Utah, 
and  Mohave  Co.,  Arizona  (Atwood 
1975,  in  press). 

Phacelia  crenulata  Torr.  ex  Wats,  in  King 
var.  angustifolia  Atwood,  Great  Basin 
Nat.  35:   158.  1975. 

Status:  Restricted  and  in  disjunct 
populations,  neither  threatened  nor 
endangered. 

Distribution:  Beaver,  Garfield,  and 
Kane  counties,  Utah,  and  in  Coconino 
and  Mohave  counties,  Arizona  (At- 
wood 1975,  in  press). 

Phacelia  demissa  A.  Gray  var.  heterotri- 
cha  J.  T.  Howell,  Amer.  Midi.  Nat- 
uralist 29:  8.  1943. 

Type:  Marysvale,  Piute  Co.,  Utah, 
4  Jun  1894,  M.  E.  Jones  5388a  (pom). 

Status:  Endemic,  restricted  and  pos- 
sibly threatened   (Ripley,  T). 

Distribution:  Piute,  Sevier,  and 
Wayne  counties,  Utah  (Atwood,  in 
press). 

Phacelia  howelliana  Atwood,  Rhodora 
74:  456.  1972. 

Type:  Ca  0.4  miles  north  of  Bluff 
along  Utah  Highway  163,  San  Juan 
Co.,  Utah,  13  May  1970,  Atwood  2454 
(bry) . 

Status:  Endemic,  restricted  but  lo- 
cally abundant,  neither  threatened 
nor  endangered. 

Distribution:  Grand  and  San  Juan 
counties,  Utah  (bry;  Atwood  1975,  in 
press). 


Phacelia  indecora  J.  T.  Howell,  Amer. 
Midi.  Naturalist  29:  12.  1943. 

Type:  Bluff,  San  Juan  Co..  Utah, 
24  May  1919,  M.  E.  Jones  s.n.   (gas). 

Status:  Endemic,  rare  and  endan- 
gered (Ripley,  T). 

Distribution:  Wayne  and  San  Juan 
counties,  Utah   (Atwood,  in  press). 

Phacelia  mammillarensis  Atwood,  Phyto- 
logia  26:  437.  1973. 

Type:  Ca  6  miles  east  of  Glen  Can- 
yon City,  along  the  road  to  Warm 
Creek,  Kane  Co.,  Utah,  6  May  1970, 
Welsh   &   Atwood   9809    (bry). 

Status:  Endemic,  edaphically  restric- 
ted and  endangered    (Ripley,  E). 

Distribution:  Garfield  and  Kane 
counties,  Utah  (Atwood  1975,  in  press). 

Phacelia  rafaelensis  Atwood,  Rhodora  74: 
454.  1972. 

Type:  Capitol  Reef  N.M.,  Wayne 
Co.,  Utah,  12  Jun  1969,  Atwood  &  Hig- 
gins  1834  (bry). 

Status:  Restricted  and  local,  threat- 
ened (Ripley,  T). 

Distribution:  Emery,  Washington, 
and  Wayne  counties,  Utah,  and  Coco- 
nino Co.,  Arizona  (Atwood  1975,  in 
press). 

Phacelia  salina  (A.  Nels.)  J.  T.  Howell, 
Leafl.  W.  Bot.  4:  16.  1944,  based  on 
Emmenanthe  salina  A.  Nels.,  Bull. 
Torrey  Bot.  Club  24:  381.  1898,  a  sub- 
stitute name  for  E.  foliosa  M.  E.  Jones, 
Zoe  4:  278.  1893,  not  P.  foliosa  Phillip. 

Type:  Deep  Creek  Valley,  above 
Furber,  Tooele  Co.,  Utah,  8  Jun  1891, 
M.  E.  Jones  s.n.   (pom). 

Status:  Rare  and  possibly  extirpated 
from  Utah. 

Distribution:  Tooele  Co.,  Utah,  and 
adjacent  Nevada  and  southern  Wyo- 
ming  (Atwood,  in  press). 

Phacelia  tetramera  J.  T.  Howell,  Leafl. 
W.  Bot.  4:  16.  1944,  based  on  Emme- 
nanthe pusilla  A.  Gray,  Proc.  Amer. 
Acad.  Arts  11:  87.  1876,  not  P.  pusilla 
Buckley  nor  Torr. 

Status:  Rare  and  possibly  threatened. 

Distribution:  Weber  Co.,  Utah 
(Arnow  3985  [bry,  ut]);  also  in  Ne- 
vada, California  and  Oregon  (Howell 
1944). 

Phacelia  utahensis  E.  G.  Voss,  Bull.  Tor- 
rey Bot.  Club  64:   135.  1937. 


356 


GREAT   BASIN   NATURALIST 


Vol.  35,  No.  4 


Type:  Gunnison,  Sanpete  Co.,  Utah, 
7  Jun  1910,  M.  E.  Jones  s.n.  (pom). 

Status:  Endemic,  edaphically  re- 
stricted and  threatened   (Ripley,  T). 

Distribution:  Carbon  (ut),  Sanpete, 
and  Sevier  counties,  Utah  (Atwood 
1975,  in  press). 

JUNCACEAE 

Juncus  bryoides  F.  J.  Hermann,  Leafl.  W. 
Bot.  5:  117.  1948. 

Status:  Restricted,  rare  and  possibly 
threatened. 

Distribution:  Salt  Lake  and  Wash- 
ington counties,  Utah,  and  in  Califor- 
nia (Hermann  1948). 

Lamiaceae 

Stachys  rothrockii  A.  Gray,  Proc.  Amer. 
Acad.  Arts  12:  82.1876. 

Status:  Restricted,  rare  and  possibly 
endangered. 

Distribution:  Near  Orderville,  Kane 
Co.,  Utah  (bry,-  Barneby  1944),  north- 
ern New  Mexico  and  Arizona. 

LiLIACEAE 

Agave  utahensis  Engelm.  in  King,  Rep. 
Geol.  Explor.  40th  Parallel  5:  497. 
1871. 

Type:  About  St.  George,  Washing- 
ton Co.,  Utah,  Palmer  s.n.  (mo). 

Status:  Restricted  and  rare  except 
locally;  commercially  exploited  and 
possibly  endangered.  Our  plant  is  var. 
utahensis. 

Distribution:  Washington  Co.,  Utah 
(bry)  ;  also  in  Coconino  and  Mohave 
counties,  Arizona  (Breitung  1960). 

Allium  passeyi  Holmgren  &  Holmgren, 
Brittonia  26:   309.  1974. 

Type:  Southwest  part  of  Howell 
Valley,  sec.  7,  T.llN.,  R.5W.,  Box 
Elder  Co.,  Utah,  13  Jun  1960,  4,800 
feet,  Holmgren  et  al.  15125   (ny). 

Status:  Endemic,  rare  and  endan- 
gered (Ripley,  E). 

Distribution:  Box  Elder  Co.,  Utah; 
known  only  from  the  type  locality 
(Holmgren  &  Holmgren  1974). 

Eremocrinum  albomarginatum  (M.  E. 
Jones)  M.  E.  Jones,  Zoe  4:  53.  1893, 
based  on  Hesperanthes  albomarginata 
M.  E.  Jones,  Zoe  2:  251.  1891. 


Type:  Green  River,  Emery  Co.,  Utah, 
9  May  1890,  M.  E.  Jones  s.n.    (pom). 

Status:  Navajo  Basin  endemic, 
edaphically  restricted  but  locally  com- 
mon to  abundant  and  neither  threat- 
ened nor  endangered. 

Distribution:  Emery,  Garfield, 
Grand,  Kane,  San  Juan,  and  Wayne 
counties,  Utah,  and  Apache  and  Nava- 
jo counties,  Arizona  (MacDougall 
1973). 

Nolina  microcarpa  S.  Wats.,  Proc.  Amer. 
Acad.  Arts  14:  247.  1879. 

Status:  Apparently  extirpated  from 
Utah;  last  seen  in  Zion  N.P.  in  1925 
(Cottam  1974). 

Distribution:  Washington  Co., 
Utah,  to  northern  Mexico,  eastward  to 
western  Texas. 

This  plant  has  mistakenly  been  called 
Nolina  parry  i  (Kearney  &  Peebles 
1951)  or  A^.  bigelovii  (Cottam  1974). 

Yucca  brevifolia  Engelm.  in  King,  Rep. 
Geol.  Explor.  40th  Parallel  5:  496. 
1871. 

Status:  Restricted  but  locally  abun- 
dant; possibly  threatened  by  commer- 
cial exploitation. 

Distribution:  Southwestern  Wash- 
ington Co.,  Utah;  western  Arizona, 
southern  Nevada  and  adjacent  Califor- 
nia  (McKelvey  1938;  Weber  1953). 

Yucca  toftiae  Welsh,  Great  Basin  Nat. 
34:  308.  1974. 

Type:  Three  Garden,  ca  1  mile  north 
of  the  confluence  of  San  Juan  and  Colo- 
rado rivers.  Lake  Powell,  San  Juan 
Co.,  Utah,  4  Jun  1973,  Welsh  11955a 
(bry) . 

Status:  Endemic,  local  and  uncom- 
mon;  possibly  threatened. 

Distribution:  Kane  and  San  Juan 
counties,  Utah  (bry);  much  of  the 
known  range  of  this  entity  has  been 
inundated  by  the  water  of  Lake  Powell. 

Zigadenus  vaginatus  (Rvdh.")  J.  F.  Mac- 
bride,  Contr.  Gray  Herb.  53:  4.  1918, 
based  on  Anticlea  vaginata  Rvdb.,  Bull. 
Torrey  Bot.  Club  39:    108.   1912. 

Type:  Armstrong  Canyon,  near  the 
Natural  Bridges,  San  Juan  Co.,  Utah, 
4-6  Aug  1911.  Rydberg  &  Garrett  9407 
(ny). 

Status:  Endemic,  restricted  and  pos- 
sibly endangered. 


Dec.  1975 


WELSH,   ET   AL.:    ENDANGERED    UTAH   PLANTS 


357 


Distribution:  Grand,  Kane,  and  San 
Juan  counties,  Utah  (bry). 

This  distinctive  species  was  placed 
in  synonymy  under  the  Mexican  species, 
Zigadenus  volcanicus  Benth.,  PI.  Hart- 
wegianus  96.  1840,  by  I'idestrom  (1925), 
Tidestrom  and  Kittell  (1941),  and  fol- 
lowed by  Holmgren  and  Reveal  (1967). 
This  plant  is  related  to  Z.  elegans  Pursh, 
but  differs  in  having  an  elongated  panicu- 
late inflorescence  and  smaller  white  to 
greenish  flowers  borne  in  the  late  summer 
and  early  fall. 

LOASACEAE 

Mentzelia  argillosa  J.  Darlington,  Ann. 
Missouri  Bot.  Gard.  21:  153.  1934. 

Type:  Vermillion,  Sevier  Co.,  Utah, 
16  Jul  1894,  5,600  feet,  M.  E.  Jones 
563^  (mo). 

Status:  Endemic,  edaphically  re- 
stricted and  threatened. 

Distribution:  Sevier  Co.,  Utah 
(bry). 

Mentzelia  obscura  Thompson  &  Roberts, 
Phytologia  21:    284.   1971. 

Status:  Local  and  infrequent. 
Distribution:        Washington       Co., 
Utah;    southern    Nevada    and    western 
Arizona    westward    to    California    and 
Baja  California,  Mexico. 

Petalonyx  parry i  A.  Gray,  Proc.  Amer. 
Acad.  Arts  10:  72.  1874. 

Type:  St.  George,  Washington  Co., 
Utah,  "within  a  stone's  throw  of  the 
great  Mormon  Temple"  (see  Parry 
1875),  1874,  Parry  75   (gh). 

Status:  Rare  and  possibly  endan- 
gered. 

Distribution:  Washington  Co.,  Utah 
(bry),  northern  Arizona  and  southern 
Nevada  (Davis  &  Thompson  1967). 

LOGANIACEAE 

Buddie j a  utahensis  Coville,  Proc.  Biol. 
Soc.  Wash.  7:  69.  1892. 

Type:  Near  St.  George,  Washington 
Co.,  Utah,  1877,  Palmer  s.n.  (us). 

Status:  Restricted,  rare  and  possibly 
threatened. 

Distribution:  Washington  Co., 
Utah;  also  in  northwestern  Arizona, 
southern  Nevada  and  southeastern  Cali- 
fornia (Munz  1974). 


Loranthaceae 

Phorandendron  californicum  Nutt.,  J. 
Acad.  Nat.  Sci.  Philadelphia  II,  1:  185. 
1848. 

Status:  Restricted  but  locally  abun- 
dant; neither  threatened  nor  endan- 
gered. 

Distribution:  Washington  Co., 
Utah;  widespread  in  Arizona,  Nevada, 
southern  California,  and  northern  Mex- 
ico (Kearney  &  Peebles  1951). 

Malvaceae 

Sphaeralcea  caespitosa  M.  E.  Jones,  Contr. 
W.  Bot.  12:  4.  1908. 

Type:  Wah  Wah,  Beaver  Co.,  Utah, 
1906,  6,000  feet,  M.  E.  Jones  s.n.  (pom)  . 

Status:  Endemic,  rare  and  threat- 
ened. 

Distribution:  Beaver  and  Millard 
counties,  Utah  (bry;  Jefferies  1972). 

Martyniaceae 

Proboscidea  louisiana  (Miller)  ThelL, 
Mem.  Soc.  Sci.  Nat.  Cherbourg  3:  480. 
1912,  based  on  Martynia  louisiana  Mil- 
ler, Gard.  Diet,  ed  8,  Martynia  No.  3. 
1768. 

Status:  Restricted  and  uncommon, 
but  neither  rare  nor  endangered. 

Distribution:  Washington  Co.,  Utah 
(bry);  widespread  in  the  southwestern 
states  and  elsewhere. 

Najadaceae 

Najas  caespitosus  (Maguire)  Reveal,  stat. 
&  comb,  nov.,  based  on  Najas  flexilis 
ssp.  caespitosus  Maguire  in  Maguire  & 
Jensen,  Rhodora  44:   7.  1942. 

Type:  Common  in  shallow  water. 
Pelican  Point,  Fish  Lake,  Sevier  Co., 
Utah,  3  Aug  1940,  8,600  feet,  Maguire 
19888  (ny). 

Status:    Endemic   and  endangered. 

Distribution:  Sevier  Co.,  Utah; 
known  onl}'  from  the  type  area  (Ma- 
guire &  Jensen  1942). 

Najas  guadalupensis  Morong,  Mem.  Tor- 
rey  Bot.  Club  3:   60.  1893. 

Status:  Rare  and  restricted. 

Distribution:  Cache  Co.,  Utah 
(uTC;  Maguire  &  Jensen  1942);  wide- 
spread elsewhere. 


358 


GREAT   BASIN   NATURALIST 


Vol.  35,  No.  4 


Nyctaginaceae 

Boerhaavia  torreyana  (S.  Wats.)  Stand- 
ley,  Contr.  U.S.  Natl.  Herb.  12:  385. 
1909,  based  on  B.  spicata  var.  torrey- 
ana S.  Wats.,  Proc.  Amer.  Acad.  Arts 
24:  70.  1889. 

Status:  Local,  rare  and  highly  re- 
stricted; possibly  threatened. 

Distribution:  Kane  Co.,  Utah  {At- 
wood  3389  [bry,  wts]). 

Hermidium  alipes  S.  Wats.  var.  pallidum 
C.   L.   Porter,  Rhodora   54:    158.    1952. 

Type:  Ca  5  miles  south  of  Vernal, 
Uintah  Co.,  Utah,  3  Jun  1950,  5,200 
feet,  Porter  5308  (rm). 

Status:  Endemic,  rare  and  endan- 
gered. 

Distribution:  Uintah  Co.,  Utah 
(Porter  1952). 

Oleaceae 

Menodora  scabra  A.  Gray,  Amer.  J.  Sci. 
Arts  II,  14:  44.  1852. 

Status:  Rare,  in  disjunct  populations, 
and  possibly  threatened. 

Distribution:  Garfield  and  Wash- 
ington counties,  Utah  (bry)  ;  wide- 
spread in  the  southwestern  states  and 
northern  Mexico  (Kearney  &  Peebles 
1951). 

Onagraceae 

Epilobium  nevadense  Munz,  Bull.  Torrey 
Bot.  Club  56:   166.  1929. 

Status:  Rare  and  threatened  (Rip- 
ley,  T). 

Distribution:  Washington  Co., 
Utah  (Higgins  1972b),  and  Clark  Co., 
Nevada. 

Oenothera  brevipes  A.  Gray  var.  pallidula 
Munz,  Amer.  J.  Bot.  15:  229.  1928. 

Status:  Restricted  and  local,  possibly 
threatened. 

Distribution:  Washington  Co.,  Utah 
(bry)  ;  also  in  northwestern  Arizona, 
southern  Nevada  and  southern  Califor- 
nia (Raven  1969). 

Oenothera  gouldii  (Raven)  Welsh  &  At- 
wood,  comb,  nov.,  based  on  Camissonia 
gouldii  Raven,  Contr.  U.S.  Natl.  Herb. 
37:  368.  1969. 

Type:  Steep  slope  of  volcanic  cone 
among  loose  cinders,  Diamond  Valley, 
12  miles  north   of   St.    George,   Wash- 


ington Co.,  Utah,  15  Oct  1941,  3,500 
feet,  Gould  1423  (pom). 

Status:  Restricted  and  rare,  pos- 
sibly threatened. 

Distribution:  Washington  Co., 
Utah,  and  Coconino  Co.,  Arizona  (Ra- 
ven 1969). 

Oenothera  megalantha  (Munz)  Raven, 
Univ.  Calif.  Publ.  Bot.  34:  111.  1962,. 
based  on  O.  heterochroma  var.  mega- 
lantha Munz,  Leafl.  W.  Bot.  3:  52. 
1941. 

Status:  Restricted  and  rare,  endan- 
gered (Ripley,  E). 

Distribution:  Kane  Co.,  Utah  (bry), 
and  Nye  Co.,  Utah  (Raven  1969;  Beat- 
ley    1973). 

Oenothera  parryi  S.  Wats.,  Amer.  Nat- 
uralist 9:  270.  1875. 

Type:  Near  St.  George,  Washington 
Co.,  Utah,  1874,  Parry  72  (gh). 

Status:  Restricted  but  locallv  abun- 
dant (Ripley,  T). 

Distribution:  Washington  Co., 
Utah,  and  adjacent  Arizona    (bry). 

Ophioglossaceae 

Botrichium  boreale  (Fries)  Milde  var. 
obtusilobum  (Rupr.)  M.  Braun,  Index 
N.  Amer.  Ferns  33.  1938.  Based  on  B. 
crassinervum  var.  obtusilobum  Rupr., 
Beitr.  Pflanzenk.  Russ.  Reiches  11:  42. 
1959. 

Status:    Rare;    possibly    threatened. 

Distribution:  Summit  Co.,  Utah 
according  to  Flowers  1944;  not  reported 
from  Utah  by  Cronquist  et  al.  1972); 
widespread  in  northwestern  America. 

Botrychium  lanceolatum  (S.  G.  Gmelin) 
Angstrom,  Bot.  Not.  1854:  68.  1854, 
based  on  Osmunda  lanceolata  S.  G. 
Gmelin,  Novi  Comment.  Acad.  Petrop. 
12:  516-1768. 

St.\tus:  Rare;  possibly  threatened. 

Distribution:  Juab  Co.,  Utah 
(Flowers  1944;  Maguire  &  Holmgren 
1946);  widely  distributed  in  northern 
North  America    (Welsh  1974b). 

Botrichium  matricariifolium  A.  Braun  in 
Koch  var.  hesperium  (Maxon  &  Clau- 
sen) M.  Braun,  Index  N.  Amer.  Ferns 
39.  1938,  based  on  B.  matricariifolium 
ssp.  herperium  Maxon  &  Clausen, 
Mem.  Torrey  Bot.  Club  19:  38.  1938. 
Status:    Rare;   possibly  threatened. 


Dec.  1975 


WELSH,   ET   AL.:    ENDANGERED   UTAH    PL.'\NTS 


359 


Distribution:  Salt  Lake  Co.,  Utah 
(Flowers  1944;  not  reported  by  Cron- 
quist  et  al.  1972  as,  in  their  opinion, 
Flowers'  identification  was  in  error); 
also  in  Colorado. 

Orchidaceae 

Cypripcdiurn  calceolus  L.  xar.  parviflo- 
rum  (Salisb.)  Fern.,  Rhodora  48:  4. 
1946,  based  on  C.  parviflorum  Salisb., 
Trans.  Linn.  Soc.  London  1:   77.  1791. 

Status:    Extirpated  or  endangered. 

Distribution:  Cache  (utc)  and 
Utah  (bry)  counties,  LTtah;  widespread 
in  northern  North  America. 

Cypridedium  fasciculatum  Kellogg  ex 
Wats.,  Proc.  Amer.  Acad.  Arts  17:  380. 
1882. 

Status:  Rare  and  endangered. 

Distribution:  Salt  Lake  (bry)  and 
Summit  (Tidestrom  1925)  counties, 
Utah;  widespread  and  rare  in  Wash- 
ington, California,  Idaho,  Montana, 
Wyoming,  and  Colorado. 

Spiranthes  cernua  (L.)  Rich.,  Mem.  Mus. 
Hist.  Nta.  4:  59.  1818,  based  on  Ophrys 
cernua  I..,  Sp.  PI.  946.  1753. 

Status:  Extirpated  from  Utah. 

Distribution:  Salt  Lake  Co.,  Utah 
(M.  E.  Jones  1908  [pom]);  widespread 
in  the  eastern  United  States  (Correll 
1950). 

Papaveraceae 

Papaver  radicatum  Rottb.,  Skr.  Ki0ben- 
havnske  Selsk.  Laerd.  Elsk.  10:  455. 
1770. 

Status:  Rare  and  restricted. 

Distribution:  Duchesne  Co,,  Utah 
(bry)  ;  \\ddespread  in  northern  North 
America,  circumboreal  (Welsh  1974b). 

Arcotmecon  humilis  Coville,  Proc.  Biol. 
Soc.    Wash.    7:    67.    1892. 

Type:  St.  George,  Washington  Co., 
Utah,  1874,  Parry  s.n.   (gh). 

Status:  Restricted  and  rare,  endan- 
gered (Ripley,  E) . 

Distribution:  Washington  Co., 
Utah,  and  Mohave  Co.,  Arizona  (bry). 

PiNACEAE 

Pinus  longaeva  D.  K.  Bailey,  Ann.  Mis- 
souri Rot.  Card.  57:  243.  1970. 


Status:  Restricted  but  hardly  rare; 
possibly  threatened. 

Distribution:  Beaver  (personal  ob- 
servations). Carbon,  Duchesne,  Gar- 
field, Iron  (personal  observations), 
Kane,  Millard,  and  Washington  coun- 
ties, Utah;  also  in  Nevada  and  eastern 
California   (Bailey  1970). 

Poaceae 

Andropogon  glomeratus  (Walter)  Britton, 
Sterns  &  Poggenb.,  Prelim.  Cat.  Antho. 
&  Pterid.  within  100  mi.  New  York  City 
67.  1888,  based  on  Cinna  glomerata 
Walter,  Fl.  Carol.  59.  1788. 

Status:  Rare  and  possibly  extirpated 
from  Utah,  previously  known  only 
along  Lake  Powell. 

Distribution:  Kane  and  San  Juan 
counties,  Utah  (bry);  widely  distrib- 
uted in  southern  and  southeastern 
United  States,  West  Indies,  Yucatan 
and  Central  America. 

Blepharidachne  kingii  (S.  Wats.)  Hackel 
in  DC,  Monogr.  Phanerog.  6:  261. 
1889,  based  on  Eremochloe  kingii  S. 
Wats,  in  King,  Rep.  Geol.  Explor.  40th 
Parallel  5:  382.  1871. 
Status:  Rare  and  local. 
Distribution:  Beaver,  Millard,  and 
Tooele  counties,  Utah,  and  adjacent 
Nevada  (bry). 

Festuca  dasyclada  Hackel  ex.  Beal, 
Grasses  N.  Amer.  2:  602.  1896. 

Type:  Joe's  Valley,  Emery  Co., 
LTtah,  1875,  Parry  s.n.   (us). 

Status:  Endemic  and  possibly  en- 
dangered (Ripley,  PoEx). 

Distribution:  Emery  and  Sanpete 
counties,  Utah  (N.  H.  Holmgren,  per- 
sonal communication),  and  Colorado 
(N.   West,  personal   communication).^" 

Festuca  sororia  Piper,  Contr.  U.S.  Natl. 
Herb.  16:   197.  1913. 

Status:   Rare  and  restricted. 
Distribution:      Southeastern      Utah 
(Hitchcock  &  Chase  1950). 

Helictotrichon  mortonianum  (Lams.- 
Scribn.)  Henrard,  Blumea  3:  429. 
1940,  based  on  Avena  mortonianum 
Lams.-Scribn.,  Bot.  Gaz.  21:   133.  1896. 

Status:  Rare  and  endangered. 

Distribution:    Utah    (Hitchcock    & 


^"Until    1975    this    species   was   known    only    from    the    type. 


360 


GREAT   BASIN    NATURALIST 


Vol.  35,  No.  4 


Chase  1950) ;  also  in  Colorado  and  New 
Mexico. 

Imperata  brevifolia  Vasey,  Bull.  Torrey 
Bot.  Club  13:  26.  1886. 

Status:  Rare,  restricted  and  endan- 
gered or  possibly  extirpated  from  Utah. 

Distribution:  San  Juan  Co.,  Utah 
(bry)  ;  California,  Nevada,  and  Mexico 
(Hitchcock  &  Chase  1950). 

The  known  localities  in  Utah  where 
this  grass  occurred  have  been  inundated 
by  Lake  Powell. 

Muhlenbergia  arsenei  Hitchc,  Proc.  Biol. 
Soc.  Wash.  41:  161.  1928. 

Status:  Rare  and  local. 

Distribution:  Garfield  (bry),  Kane 
(bry),  San  Juan  (ut),  and  Washing- 
ton (utc)  counties,  Utah;  New  Mexico 
and  Cahfomia  (Cottam  et  al.  1940). 

Muhlenbergia  curtifolia  Lams.-Scribn., 
Bull.  Torrey  Bot.  Club  38:  328.  1911. 

Type:  Between  Kanab  and  Carmel, 
Kane  Co.,  Utah,  M.  E.  Jones  6047  (us). 

Status:  Restricted  and  local,  pos- 
sibly threatened. 

Distribution:  Garfield,  Kane,  San 
Juan,  and  Washington  (ut)  counties, 
Utah  (bry,  utc)  ;  southern  Nevada 
and  northern  Arizona  (Hitchcock  & 
Chase  1950). 

Muhlenbergia  minutissima  i(Steudel) 
Swallen,  Contr.  U.S.  Natl.  Herb.  29: 
207.  1947,  based  on  Agrostis  minutis- 
sima Steudel,  Syn.  PI.  Glum.  1:  171. 
1854. 

Status:  Local  and  rare. 
Distribution:  Washington  Co.,  Utah 
(Atwood  &  Higgins  5528  [bry]). 

Muhlenbergia  wrightii  Vasey  in  Coult., 
Manual  Bot.  Rocky  Mt.  Region  409. 
1885. 

Status:  Rare  and  local. 
Distribution:    Utah    (Hitchcock    & 
Chase  1950);  Arizona,  Colorado,  Okla- 
homa,    New     Mexico,     and     northern 
Mexico. 

Puccinellia  simplex  Lams.-Scribn., 
U.S.D.A.  Div.  Agrostol.  Circ.  16:  1. 
1899. 

Status:  Rare  and  possibly  threatened. 

Distribution:  Weber  Co.,  Utah 
{Arnow  3986  [bry]);  also  in  Califor- 
nia  (Hitchcock  &  Chase  1950). 


Sporobolus  pulvinatus  Swallen,  J.  Wash. 
Acad.  Sci.  31:  351.  1941. 

Status:  Rare  and  possibly  threat- 
ened. 

Distribution:  San  Juan  Co.,  Utah 
(bry);  New  Mexico,  Arizona,  Texas 
and  northern  Mexico  (Hitchcock  & 
Chase  1950). 

POLEMONIACEAE 

Gilia  caespitosa  A.  Gray,  Proc.  Amer. 
Acad.  Arts  12:  80.  1876. 

Type:  Rabbit  Valley  on  barren  cliffs 
of  sandstone,  Wayrte  Co.,  Utah,  1875, 
7,000  feet,  Ward  s.n.  (gh). 

Status:  Endemic,  rare  and  endan- 
gered (Ripley,  E). 

Distribution:  Wayne  Co.,  Utah 
(bry,  utc). 

Gilia  lad  folia  S.  Wats,  ex  Parry,  Amer. 
Naturalist  9:  347.  1875. 

Type:  Valley  of  the  Virgin,  near  St. 
George,  Washington  Co.,  Utah,  1874, 
Parry  188  (gh). 

Status:  Rare  and  local;  possibly 
threatened. 

Distribution:  Kane,  Wayne,  and 
Washington  counties,  Utah  (bry)  ;  Ari- 
zona, southern  Nevada  and  southern 
California  (Matthews  1971). 

Gilia  mcvickerae  M.  E.  Jones,  Proc.  Calif. 
Acad.  Sci.  II,  5:   712.  1895. 

Type:  Marysvale,  Piute  Co.,  Utah, 
7,000  feet,  M.  E.  Jones  5378  (pom). 

Status:  Endemic,  rare  and  local; 
threatened  (Ripley,  T). 

Distribution:  Piute,  Sevier,  and 
Uintah   counties,   Utah    (bry). 

Gilia  stenothyrsa  A.  Gray,  Proc.  Amer. 
Acad.  Arts  8:  276.  1870. 

Type:  In  a  clear  forest,  Uinta  Mts., 
Duchesne  or  Uintah  counties,  Utah, 
1844,  Fremont  s.n.  (gh). 

Status:  Endemic,  locally  common 
and  neither  threatened  nor  endangered. 

Distribution:  Carbon  (ut),  Emery, 
Duchesne  and  Uintah  counties,  Utah 
(bry,  utc). 

Phlox  cluteana  A.  Nels.,  Amer.  J.  Bot.  28: 
24.  1922. 

St.a>tus:  Rare  and  local;  possibly 
threatened  (Ripley,  T). 

Distribution:  San  .Juan  Co.,  Utah 
(bryj,  and  northern  Arizona  (Kearney 
&  Peebles  1951). 


Dec.  1975 


WELSH,   ET   AL.:    ENDANGERED    UTAH    PLANTS 


361 


Phlox  gladiformis  (M.  E.  Jones)  E.  Nels., 
Rev.  West.  N.  Amer.  Phloxes  21.  1899, 
based  on  P.  longifolia  var.  gladiformis 
M.  E.  Jones,  Proc.  Calif.  Acad.  Sci.  II, 
5:  711.  1895. 

Type:  Cedar  City,  Iron  Co.,  Utah, 
11  May  1894,  6,500  feet,  M.  E.  Jones 
5208c  (pom). 

Status:  Rare  and  local;  possibly 
threatened  (Ripley,  T). 

Distribution:  Garfield,  Iron,  and 
Washington  counties,  Utah  (bry),  and 
adjacent  Nevada. 

Phlox  grahomii  WherrA',  Brittonia  5:  63. 
1943. 

Type:  Talus  slopes  on  west  side  of 
Green  River,  south  of  the  mouth  of 
Sand  Wash.  Uintah  Co.,  Utah,  27  May 
1933,  Graham  7884   (cm). 

Status:  Endemic,  rare  and  local; 
threatened  (Ripley,  T). 

Distribution:  Uintah  Co.,  Utah; 
known  only  from  the  type  locality. 

Phlox  jonesii  Wherry,  Notul.,  Nat.  Acad. 
Nat.  Sci.  Philadelphia  146:   8.   1944. 

Type:  Zion  Canyon,  Washington 
Co.,  Utah,  7  May  1923,  M.  E.  Jones 
s.n.  (us). 

Status:  Endemic,  rare  and  threat- 
ened   (Ripley,  T) . 

Distribution:  Washington  Co., 
Utah;  known  only  from  the  type  lo- 
cality (Wherry  1955). 

Polygonaceae 

Eriogonum  ammophilum  Reveal,  Phyto- 
logia  23:  163.  1972. 

Type:  Ca  1.3  miles  northwest  of 
Ibex  Warm  Point,  on  a  dry  sandy  flat, 
Millard  Co.,  Utah,  4  Aug  1970,"  5,270 
feet,  Holmgren  &  Holmgren  4650  (us). 

Status:  Endemic,  rare  and  local; 
endangered  (Ripley,  E). 

Distribution:     Millard     Co.,     Utah 

(bry,    DERM,    UTC). 

Eriogonum  aretioides  Barneby,  Leafl.  W. 
Bot.  5:  154.  1949. 

Type:  Bare  limestone  gravel  benches 
in  the  foothills  of  the  Escalante  Range 
at  Widtsoe,  Garfield  Co.,  Utah,  8  Jun 
1947,  7,750  feet,  Ripley  &  Barneby 
8570  (c.-^s). 

Status:  Endemic,  edaphically  re- 
stricted and  endangered  (Ripley,  E) . 


Distribution:     Garfield     Co.,    Utah 

(BRY,    UT,    UTC). 

Eriogonum  hatemanii  M.  E.  Jones,  Contr. 
W.  Bot.  11:  11.  1903. 

Type:  Price,  Carbon  Co.,  Utah,  29 
Jun  1898,  M.  E.  Jones  s.n.    {pom). 

Status:  Restricted  and  local,  neither 
threatened  nor  endangered. 

Distribution:  Carbon,  Duchesne, 
Emery,  Garfield,  and  Uintah  counties, 
Utah,  and  Rio  Blanco  Co.,  Colorado 
(Reveal  1973a). 

Eriogonum  brevicaule  Nutt.  var.  cottamii 
(S.  Stokes)  Reveal,  Great  Basin  Nat. 
32:  113.  1972,  based  on  E.  tenellum 
ssp.  cottamii  S.  Stokes,  Gen.  Eriog.  70. 
1936. 

Type:  Canyons  in  bottoms  of  the 
slopes  of  West  Mtn.,  Utah  Co.,  Utah, 
20  Aug  1925,  Cottam  411   (bry). 

Status:  Endemic,  restricted  and  rare. 

Distribution:  Juab,  Millard,  and 
Utah   counties,  Utah    (bry.   ny,   utc). 

Eriogonum  brevicaule  Nutt.  var.  wa- 
satchense  (M.  E.  Jones)  Reveal,  Great 
Basin  Nat.  32:  113.  1972,  based  on  E. 
wasatchense  M.  E.  Jones,  Contr.  W. 
Bot.   11:    11.   1903. 

Type:  American  Fork  Canyon,  Utah 
Co.,  Utah,  27  Jul  1880,  M.  E.  Jones 
1877  (pom). 

Status:  Endemic,  restricted  and 
rare. 

Distribution:  Davis,  Juab,  Millard, 
Salt  Lake,  Utah,  and  Weber  counties, 
Utah  (bry,  ds,  gh,  ut,  utc). 

Eriogonum  clavellatum  Small,  Bull.  Tor- 
rey  Bot.  Club  25:  48.  1898. 

Type:  Barton  Range,  San  Juan  Co., 
Utah,  13  Jul  1895,  Eastwood  132  (ny). 

Status:  Rare  and  highly  restricted; 
threatened  (Ripley,  T). 

Distribution:  San  Juan  Co.,  Utah 
(bry,  utc),  and  Montezuma  Co.,  Colo- 
rado (cs). 

Eriogonum  contortum  Small  ex  Rydb., 
Agric.  Exp.  Sta.  Agric.  Coll.  Colorado 
Bull.  (Fl.  Colorado)  100:   107.  1906. 

Status:  Infrequent  but  neither 
threatened  nor  endangered. 

Distribution:  Grand  Valley  en- 
demic in  Grand  Co.,  Utah  (bry,  utc), 
and  Garfield  and  Mesa  counties,  Colo- 
rado. 


362 


GREAT   BASIN   NATURALIST 


Vol.  35,  No.  4 


Eriogonum  corymbosum  Benth.  in  DC. 
var.  davidsei  Reveal,  Great  Basin  Nat. 
27:  216.  1968. 

Type:  Ca  0.7  miles  south  of  U.S. 
Highway  50-6  at  Wellington,  just  south 
of  the  Price  River  bridge,  9  Sep  1967, 
Reveal  &  Davidse  956  (utc). 

Status:  Endemic,  restricted  and  lo- 
cal;  endangered    (Ripley,  E). 

Distribution:  Carbon  Co.,  Utah; 
known  only  from  type  locality. 

Eriogonum  corymbosum  Benth.  in  DC. 
var.  divaricatum  Torr.  &  Gray  in  Beck- 
with,  Explor.  &  Surv.  Railroad  Route 
from  Mississippi  River  to  Pacific  Ocean 
2:  29.  1857. 

Type:  Near  Green  River,  Emery  Co.. 
Utah,  1  Oct  1853,  Creutzfeldt  s.n.  (ny). 

Status:  Endemic,  local  and  restric- 
ted; neither  endangered  nor  threatened. 

Distribution:  Emery,  Garfield,  and 
Grand  counties,  Utah. 

This  form  of  Eriogonum  corymbosum 
differs  from  var.  corymbosum  in  having 
small,  usually  crenulate  leaves,  smaller 
more  compact  stature,  and  compact  inflo- 
rescences. 

Eriogonum  corymbosum  Benth.  in  DC. 
var.  revealianum  (Welsh)  Reveal,  stat. 
&  comb,  nov.,  based  on  E.  revealianum 
Welsh,  Great  Basin  Nat.  30:   17.  1970. 

Type:  Gravelly,  boulder-strewn,  east- 
facing  slope  near  the  head  of  the  can- 
yon at  milepost  26  south  of  Antimony 
along  Utah  Highway  22,  Garfield  Co., 
Utah,  4  Sep  1969,  S.  L.  &  S.  L.  Welsh 
9389  (bry). 

Status:  Endemic,  rare  and  local;  en- 
dangered. 

Distribution:  Garfield  Co.,  Utah; 
known  only  from  the  type  locality. 

This  form  of  Eriogonum  corymbosum 
differs  from  var.  corymbosum  in  having 
elongated,  entire  leaves  concentrated 
near  the  base  of  elongated  flowering 
stems    and    open,     spreading    inflores- 


Eriogonum    cronquistii   Reveal,    Madrono 
19:  289.  1969. 

Type:  Loose  decomposed  granite  talus 
slopes  on  the  west  side  of  Bull  Mtn., 
Henry  Mts.,  Garfield  Co.,  Utah.  14  Aug 
1967,  8,300  feet,  Holmgren  &  Reveal 
3010  (utc). 


Status:  Endemic,  restricted  and 
local;  endangered  (Ripley,  E). 

Distribution:  Garfield  Co.,  Utah; 
known  only  from  the  type  locality 
(bry,  NY,  utc). 

Eriogonum  desertorum  (Maguire)  R.  J. 
Davis,  Fl.  Idaho  246.  1952,  based  on 
E.  chrysocephalum  ssp.  desertorum  Ma- 
guire, Leafl.  W.  Bot.  3:   11.  1941. 

Status:  Restricted  and  rare;  possibly 
threatened. 

Distribution:  Box  Elder  and  Tooele 
counties,  Utah;  northeastern  Nevada 
and  southern  Idaho  (Reveal  1973a). 

Eriogonum  ephedroides  Reveal,  Madroiio 
19:  295.  1969. 

Type:  Ca  10  miles  south  of  Bonanza 
along  Utah  Highway  45  south  of  the 
White  River,  25  Jul  1965,  Holmgren 
et  al.  2265   (utc). 

Status:  Uinta  Basin  endemic,  re- 
stricted and  rare;  endangered  (Ripley, 
E). 

Distribution:  Uintah  Co.,  Utah 
(bry,  utc),  and  Rio  Blanco  Co.,  Colo- 
rado (ny). 

Eriogonum  eremicum  Reveal,  Ph-s^tologia 
23:  165.  1972. 

Type:  Ca  17  miles  southeast  of  Gar- 
rison along  Utah  Highway  21,  Millard 
Co.,  Utah,  23  Jul  1965,  Holmgren  et 
al.  2247  (utc). 

Status:  Endemic,  restricted  and 
rare;  threatened  (Ripley,  T). 

Distribution:  Millard  Co.,  Utah 
(bry,  utc). 

Eriogonum  fasciculatum  Benth.  var.  poli- 
folium  (Benth.  in  DC.)  Torr.  &  Gray, 
Proc.  Amer.  Acad.  Arts  8:  169.  1870, 
based  on  E.  polifolium  Benth.  in  DC, 
iProdr.  14:   12.  1856. 

Status:  Local  and  common  to  abun- 
dant, neither  threatened  nor  endan- 
gered. 

Distribution:  Emery  (bry,  ut,- 
where  rare)  and  Washington  (where 
common)  counties,  Utah;  widespread 
and  common  in  Arizona,  Nevada,  Cali- 
fornia, and  Baja  California,  Mexico. 

Eriogonum  grayi  Reveal,  Phvtologia  25: 
193.  1973. 

Type:  Lake  Blanche,  Salt  Lake  Co., 
Utah,  15  Aug  1947.  Holmgren  et  al. 
7121   (uTC). 


Dec.  1975 


WELSH,   ET   AL.:    ENDANGERED   UTAH   PLANTS 


363 


Status:  Endemic,  disjunct  and  infre- 
quent. 

Distribution:  Juab,  Salt  Lake,  Utah, 
and  Weber  counties,  Utah  (bry,  wsco. 
UTC;  Reveal  1973a). 

Eriogonum  humivagans  Reveal,  Madrono 
19:  219.  1969. 

Type:  Ca  13.5  miles  east  of  Monti- 
cello,  13  Aug  1966,  6,800  feet,  Holm- 
gren &  Reveal  3001  (utc). 

Status:  Endemic,  restricted  and  rare; 
endangered  (Ripley,  E). 

Distribution:  San  Juan  Co.,  Utah; 
known    on!}'    from    the    type    locality. 

Eriogonum  hylophilum  Reveal  &  Brother- 
son,  Great  Basin  Nat.  27:   190.  1968. 

Type:  Along  Utah  Highway  53  in 
Gate  Canyon,  2.7  miles  southwest  of 
the  summit  of  the  Badlands  Cliffs,  Du- 
chesne Co.,  Utah,  15  Aug  1966,  6,500 
feet,  Holmgren  &  Reveal  3017    (utc). 

Status:  Endemic,  rare  and  restricted; 
endangered  (Ripley,  E) . 

Distribution:  Duchesne  Co.,  Utah 
(Reveal  1968a;  1973a). 

Eriogonum  intermontanum  Reveal,  Ma- 
drono 19:  293.  1969. 

Type:  Ca  1.5  miles  south  of  the  Uin- 
tah Co.  line  at  the  head  of  Middle 
Canyon  of  West  Water  Creek  drain- 
age in  Roan  Cliffs,  Grand  Co.,  Utah, 
27  Jul  1965,  8,400  feet,  Holmgren  et 
al.  2278  (uTc). 

Status:  Endemic,  rare  and  local;  en- 
dangered  (Ripley,  E). 

Distribution:  Grand  Co.,  Utah 
(bry,  utc). 

Eriogonum  jamesii  Benth.  in  DC.  var. 
rupicola  Reveal,  Phytologia  25:  202. 
1973. 

Type:  Along  Utah  Highway  15,  4.9 
miles  west  of  the  east  entrance  to  the 
park  on  Checkerboard  Mesa,  Zion  N.P., 
Washington  Co.,  Utah.  12  Aug  1972, 
Reveal  &  Reveal  2874  (us). 

Status:  Endemic,  restricted  and  rare; 
threatened   (Ripley,  T) . 

Distribution:  Kane  and  Washing- 
ton counties,  Utah   (Reveal  1973a). 

Eriogonum  lancifolium  Reveal  &  Brother- 
son,  Great  Basin  Nat.  27:   188.  1968. 

Type:  On  low  hills  5  miles  east  of 
Wellington,  Carbon  Co.,  Utah,  9  Sep 
1967,  Reveal  &  Davidse  957   (utc). 


Status:  Endemic,  restricted  and  lo- 
cal; threatened  (Ripley,  E). 

Distribution:  Carbon  Co.,  Utah 
(bry,  us,  utc). 

Eriogonum  leptocladon  Torr.  &  Gray  in 
Beckwith,  Explor.  &  Surv.  Railroad 
Route  from  Mississippi  River  to  Pacific 
Ocean  2:  129.  1857. 

Type:  Near  the  Green  River,  Emery 
Co.,  Utah,  1  Oct  1853,  Creutzfeldt  s.n. 
(ny). 

Status:  Endemic,  edaphically  re- 
stricted but  locally  abundant  and 
neither  threatened  nor  endangered.  Our 
plant  is  vai .  leptocladon. 

Distribution:  Emery,  Garfield, 
Grand,  San  Juan,  and  Wayne  counties, 
Utah  (bry.  UTC;  Reveal  1966). 

Eriogonum  leptophyllum  (Torr.  &  Gray) 
Wooton  &  Standley,  Contr.  U.S.  Natl. 
Herb.  16:  118.  1913,  based  on  E.  ef- 
fusum  var.  leptophyllum  Torr.  in  Sitgr., 
Rep.  Exped.  Down  Zuni  &  Colorado 
rivers  168.  1853. 

Status:   Local  and  rare;  threatened. 

Distribution:  San  Juan  Co.,  Utah 
{Harrison  12163  [bry]  ) ;  southwestern 
Colorado,  northeastern  Arizona  and  ad- 
jacent New  Mexico   (Reveal  1968a). 

Eriogonum  loganum  A.  Nels.,  Bot.  Gaz. 
54:   149.  1912. 

Type:  Logan,  Cache  Co.,  Utah,  26 
Jun  1909,  Smith  1704   (rm). 

Status:  Endemic,  extremel}^  restric- 
ted and  rare;  endangered    (Ripley,  E). 

Distribution:  Cache  Co.,  Utah 
(bry.  utc). 

Eriogonum  nanum  Reveal,  Phytologia  25: 
194.  1973. 

Type:  Talus  slopes  and  limestone 
outcrops  south  of  Willard  Peak,  Box 
Elder  Co.,  Utah,  31  Aug  1964,  9,500 
feet.  Reveal  &  Holmgren  665   (us). 

St.\tus:  Endemic,  restricted  and  rare; 
threatened  (Ripley,  T). 

Distribution:  Box  Elder  and  Weber 
counties,  Utah   (bry,  utc,  wsco). 

Eriogonum   natum   Reveal,   spec.   nov. 

A  Eriogono  hrevicaule  Nutt.  differt 
foliis  ellipticis,  2-2.5  (3)  cm  longis  et  (8) 
10-13  (15)  mm  latis,  lanatis,  inflores- 
centiis  cymoso-umbellatis,  floribus  flavis, 
2-2.5   (3)  mm  longis,  glabris. 

Spreading  herbaceous  perennials  1-3.5 


364 


GREAT   BASIN    NATURALIST 


Vol.  35,  No.  4 


dm  high,  1-4  dm  across,  with  a  short 
woody  caudex  arising  from  a  stoutish, 
woody  taproot;  leaves  essentially  basal, 
the  leaf-blade  elliptic,  2-2.5  (3)  cm  long, 
(8)  10-13  (15)  mm  wide,  densely  to- 
mentose  below,  somewhat  less  so  and 
greenish-tomentose  above,  the  petiole  (1) 
2-3  cm  long,  tomentose;  flowering  stems 
erect  to  spreading,  slender,  1-2  (2.5)  dm 
long,  white  to  greenish-tomentose;  inflo- 
rescences cymose-umbellate.  3-10  (15)  cm 
long,  3-5  (8)  cm  wide,  trichotomously 
branched  throughout,  tomentose;  bracts 
scalelike  to  foliaceous,  ternate,  the  former 
1-3  mm  long,  tomentose  to  floccose  with- 
out, tomentose  within,  the  latter  1-3  per 
node,  linear-lanceolate  to  lanceolate,  5-10 
(12)  mm  long,  (1.5)  2-4  (5)  mm  wide, 
tomentose;  peduncles  lacking;  involucres 
solitary  or  infrequently  in  groups  of  2, 
turbinate-campanulate,  2.5-4  mm  long,  2-3 
mm  wide,  thinly  to  densely  tomentose 
without,  glabrous  within,  the  5  acute 
teeth  0.5-0.8  mm  long,  usually  with  a 
membranaceous  margin,  the  bractlets 
linear-oblanceolate,  1.5-3  mm  long, 
fringed  with  gland-tipped  cells,  the  pedi- 
cels 2.5-5  mm  long,  glabrous;  flowers 
bright  yellow  with  golden  yellow  bases 
and  golden  to  greenish  midribs,  2-2.5  (3) 
mm  long,  glabrous,  the  tepals  oblong  to 
obovate,  distinctly  keeled  at  the  base  and 
along  the  midrib  of  each  tepal,  united 
about  14  to  1/5  the  length  of  the  flower; 
stamens  exserted,  2.5-4  mm  long,  the  fila- 
ments sparsely  pilose  basally,  the  anthers 
yellow,  0.3-0.5  mm  long,  oblong  to  oval; 
achenes  light  brown,  2-3  mm  long,  the 
globose  base  tapering  to  a  long,  3-angled, 
slightly  roughened  beak. 

Type:  UTAH:  Millard  Co.:  Along 
U.S.  Highway  50-6,  46.2  miles  east  of  the 
Nevada  state  line  and  about  43  miles  west 
of  Delta,  on  low  white  alkaline  clay  out- 
crops 50-300  meters  north  of  the  highway, 
ca  0.2  miles  east  of  the  (hrt  road  junction 
to  the  Antelope  Spring-Black  Hill  Well 
roads,  north-northwest  of  Sevier  Lake.  1 3 
Aug  1975,  Reveal  &  Reveal  3924.  Holo- 
type,    us!    Isotypes,  ariz,   asu,   bry,   cas, 

COLO,     GH,     ISC,     K,     MARY,     MO,     NY,     OKL, 

osc.  ph.  rm,  rsa,  smu,  tex,  uc,  utc,  wtu! 

Additional    specimens    examined: 

UTAH:  Millard  Co.:  Ca  43  miles  west  of 

Delta,    30    Aug    1975,    Reveal   &    Reveal 

3999     (bry,     CAS,     GH,     NY,     OKL,     RSA,     US, 

UTc) ;  ca  29.8  miles  west  of  Delta,  30  Aug 


1975,  Reveal  &  Reveal  4000    (ariz,  asu, 

BRY,  CAS,  GH,  ISC,  MARY,  MO,  NY,  OKL,  OSC, 
RM,   RSA,   SMU,   TEX,    US,   UTC,   WTu)  . 

Eriogonum  natum  belongs  to  the  large 
species  group  t}pified  by  E.  brevicaule 
and  is  seemingly  most  closely  related  to 
E.  brevicaule  var.  cottamii  (S.  Stokes)  Re- 
veal, a  narrowly  restricted  variant  of  the 
pinyon-juniper  woodlands  of  north  central 
Utah.  The  new  species  differs  from  var. 
cottamii  in  having  longer  and  broader  el- 
liptical leaves,  a  longer  but  less  branched 
inflorescence,  and  smaller  flowers.  The 
new  species  is  restricted  to  the  white  alka- 
line beaches  of  Sevier  Lake  and  is  cur- 
rently known  for  the  two  locations  cited 
above. 

Eriogonum  natum  is  named  to  honor 
its  discoverer,  Mark  L.  Reveal 
(1961-         ). 

Status:  Endemic,  rare  and  threat- 
ened. 

Distribution:  Millard  Co.,  Utah. 

Eriogonum  nummulare  M.  E.  Jones, 
Contr.  W.  Bot.  11.  13.  1903. 

Type:  Dutch  Mtn.,  Tooele  Co.,  Utah, 
15  Jun  1900,  M.  E.  Jones  s.n.   (pom). 

Status:  Endemic,  rare  and  seeming- 
ly local. 

Distribution:  Juab,  Millard,  and 
Tooele  counties,  Utah   (Reveal  1973a). 

Eriogonum  ostlundii  M.  E.  Jones,  Contr. 
W.  Bot.  11:  12.  1903. 

Type:  Near  Joseph  City,  Sevier  Co., 
Utah,  13  Jun  1898,  M.  E.  Jones  s.n. 
(pom). 

Status:  Endemic,  restricted  and  lo- 
cal; threatened    (Ripley,  T). 

Distribution:  Piute  and  Sevier 
counties,  Utah  (Reveal  1973a). 

Eriogonum  panguicense  (M.  E.  Jones) 
Reveal,  Proc.  Utah  Acad.  Sci.  42:  291. 
1966,  based  on  E.  pauciflorum  var. 
panguicense  M.  E.  Jones.  Contr.  W. 
Bot.  11:  9.  1903. 

Type:  Panguitch,  Garfield  Co.,  Utah, 
24  Jun  1890,  M.  E.  Jones  s.n.   (pom). 

Status:  Endemic,  restricted  and  lo- 
cal. 

Distribution:  Garfield,  Iron,  Kane, 
Sevier,  and  Washington  counties,  Utah 
(Reveal   1966;   1973a). 

Eriogonum  panguicense  (M.  E.  Jones) 
Reveal  var.  alpestre  (S.  Stokes)  Reveal, 
Proc.   Utah  Acad.   Sci.   42:    292.    1966, 


Dec.   1975 


WELSH,   ET   AL.:    ENDANGERED   UTAH   PLANTS 


365 


based  on  E.  chrysocephalum  ssp.  al- 
pestre  S.  Stokes,  Gen.  Eriog.  93.  1936. 

Type:  Cedar  Breaks  N.M.,  Iron  Co., 
Utah,  18  Jul  1930,  Goodman  &  Hitch- 
cock 1601  (cAs). 

Status:  Endemic,  rare  and  local; 
threatened  (Ripley,  T). 

Distribution:   Iron  Co.,  Utah   (bry, 

us,  UTC). 

Eriogonum  pharnaceoides  Torr.  in  Sitgr. 
var.  cervinum  Reveal,  Great  Basin  Nat. 
34:  245.  1974. 

Type:  Foothills  south  of  Pinto  on  the 
north  slope  of  the  Pine  Valley  Mts.,  18 
Aug    1973,    Atwood    &    Higgins    5895 

(us)- 

Status:  Rare,  in  disjunct  populations. 

Distribution:  Millard  and  Washing- 
ton counties,  Utah,  and  in  Mohave  Co., 
Arizona,  and  Lincoln  Co.,  Nevada  (Re- 
veal 1974). 

Eriogonum  plumatella  Dur.  &  Hilg.,  J. 
Acad.  Nat.  Sci.  Philadelphia  II,  3:  45. 
1855. 

Status:  Probably  extirpated  from 
Utah. 

Distribution:  "Utah"  {Palmer  s.n. 
[gh]  ) ;  infrequent  and  widely  scat- 
tered in  northwestern  Arizona,  south- 
ern Nevada  and  southeastern  California. 

Eriogonum  saurinum  Reveal,  Great  Basin 
Nat.  27:  197.  1968. 

Type:  Along  the  Island  Park  road, 
10  miles  east  of  Vernal  along  Brush 
Creek  on  steep  hillsides  on  the  ridges, 
Uintah  Co.,  Utah,  15  Aug  1966,  5,200 
feet,  Holmgren  &  Reveal  3019   (utc). 

Status:  Edaphically  restricted; 
threatened  (Ripley,  T). 

Distribution:  Uintah  Co.,  Utah,  and 
adjacent  northwestern  Colorado  (Re- 
veal 1973a). 

Eriogonum  smithii  Reveal,  Great  Basin 
Nat.  24:  202.  1968. 

Type:  Between  Little  Flat  Top  and 
Big  Flat  Top,  San  Rafael  Desert,  ca  10 
miles  southeast  of  Utah  Highway  24, 
Emery  Co.,  Utah,  14  Aug  1966,  5,500 
feet,  Holmgren  &  Reveal  3012    (utc). 

Status:  Endemic,  edaphically  re- 
stricted and  local;  threatened  (Ripley, 
T). 

Distribution:  Emery  Co.,  Utah 
(bry,  utc)  . 


Eriogonum  spathulatum  A.  (jray,  Proc. 
Amer.  Acad.  Arts  10:   76.  1874. 

Type:  Lower  Valley  of  the  Sevier 
River,  Sevier  Co.,  Utah,  Jul  1874, 
Parry  245   (gh). 

Status:  Endemic  and  scattered  in 
isolated  populations,  but  neither  threat- 
ened nor  endangered. 

Distribution:  Beaver,  Iron,  Millard, 
Piute,  Sanpete,  and  Sevier  counties, 
Utah  (Reveal  1973a). 

Eriogonum  thompsonae  S.  Wats.,  Amer. 
Naturalist  7:  302.  1873. 

Type:  Sandstone  cliffs  near  Kanab, 
Kane  Co..  Utah,  1872.  Thompson  s.n. 
(gh). 

Status:  Arizona  stri])  endemic,  re- 
stricted and  rare;  threatened  (Ripley, 
T). 

Distribution:  Kane  and  Washington 
counties,  Utah  (bry,  mary,  utc),  and 
Mohave  Co.,  Arizona  (bry.  cas,  us, 
utc). 

Eriogonum  thompsonae  S.  Wats.  var. 
albiflorum  Reveal,  Madrono  19:  299. 
1969. 

Type:  Ca  3  miles  west  of  Virgin, 
Washington  Co.,  Utah,  11  Aug  1966, 
3,700  feet,  Holmgren  &  Reveal  2991 
(utc). 

Status:  Endemic,  edaphically  re- 
stricted and  threatened   (Ripley,  T). 

Distribution:  Washington  Co., 
Utah  (bry,  utc). 

Eriogonum  tumulosum  (Barneby)  Reveal, 
Phytologia  23:  173.  1972,  based  on  E. 
villiflorum  A.  Gray  var.  tumulosum 
Barneby,  Leafl.  W.  Bot.  5:    153.  1949. 

Type:  Sandstone  ledges  and  rock- 
pavement  on  Red  Plateau,  southwest  of 
Woodside,  Emery  Co.,  Utah,  13  Jun 
1947,  Ripley  &  Barneby  8678  (cas). 

Status:  Restricted  and  very  local 
in  disjunct  populations. 

Distribution:  Duchesne  and  Emery 
counties,  Utah,  and  in  Moffat  Co., 
Colorado  (cs). 

Eriogonum  umbeUatum  Torr.  Var.  deser- 
eticum  Reveal,  var.  nov.  A  var.  umbel- 
lato  foliis  glabris  et  floribus  stramineis 
differt. 

Type:  UTAH:  Utah  Co.:  Along  the 
Timpooneke  Road,  1  mile  northwest  of 
Utah  Highway  80,  near  Timpooneke 
Campground,  east  of  Mt.  Timpanogos, 


366 


GREAT   BASIN   NATURALIST 


Vol.  35,  No.  4 


associated  with  Quercus,  Populus  and 
Artemisia  at  about  7,600  feet,  10  Jul 
1974,  Reveal  3702.  Holotype,  us!  Iso- 

typeS,  BRY,  CAS,  GH.  MARY,  MO,  NY,  OKL, 
UTC! 

Status:  Endemic,  locally  common, 
but  neither  threatened  nor  endangered. 

Distribution:  Juab,  Salt  Lake,  San- 
pete, Tooele,  Utah,  and  Wasatch 
counties,  Utah. 

This  form  of  Eriogonum  umbellatum 
has  been  confused  with  var.  dichro- 
cephalum  Gandoger  which  has  leaves 
pubescent  at  least  on  the  lower  surface. 

Eriogonum  viridulum  Reveal,  Proc.  Utah 
Acad.  Sci.  42:  287.  1966. 

Type:  Ca  8  miles  east  of  Duchesne 
along  U.S.  Highway  40,  Duchesne  Co., 
Utah,  2  Sep  1964,  Reveal  675    (utc). 

Status:  Uinta  Basin  endemic,  re- 
stricted and  local;  threatened  (Ripley, 
T). 

Distribution:  Duchesne  and  Uintah 
counties,  Utah,  and  Moffat  Co.,  Colo- 
rado (Reveal  1973a). 

Eriogonum  zionis  J.  T.  Howell,  Leafl.  W. 
Bot.  2:  253.  1940. 

Type:  Zion  N.P.  along  the  Mt.  Car- 
mel  highway  in  the  canyon  of  Clear 
Creek,  Washington  Co.,  Utah,  8  Sep 
1938,  Eastwood  &  Howell  6344   (cas). 

Status:  Endemic,  rare  and  local;  en- 
dangered (Ripley,  E). 

Distribution:  Kane  and  Washington 
counties,  Utah  (for  var.  zionis),  with 
var.  coccineum  J.  T.  Howell  restricted 
to  northern  Arizona. 

Polygonum  utahense  Brenckle  &  Cottam, 
Bull.  Univ.  Utah,  Biol.  Ser.  4 (4): 3. 
1940. 

Type:  Ca  6  miles  north  of  Escalante, 
Garfield  Co.,  Utah,  17  Sep  1935,  Cottam 
6507  (ut). 

Status:  Endemic;  species  of  uncer- 
tain taxonomic  status. 

Distribution:  Garfield  Co.,  Utah 
(bry,  ut)  ;  known  only  from  the  type 
locality. 

POLYPODIACEAE 

Asplenium  adiantum-nigrum  L.,  Sp.  PI. 
1081.  1753. 

Status:  Rare  and  local;  status  un- 
certain within  Utah  as  not  collected 
since  the  1930s  (Flowers  1944). 


Distribution:  Washington  Co., 
Utah;  widespread  in  Eurasia,  known 
only  from  three  locations  in  the  United 
States  (Cronquist  et  al.  1972). 

Asplenium  resi liens  Kunze,  Linnaea  18: 
331.  1844,  based  on  A.  parvulum  Mar- 
tens &  Galeotti,  Mem.  Foug.  Mex.  60. 
1842,  not  Hook. 

Status:  Local  and  rare;  possibly 
threatened. 

Distribution:  San  Juan  Co.,  Utah 
(Flowers  1965);  widespread  in  North 
and   South  America. 

Asplenium  septentrionale  L.,  Sp.  PL  1068. 
1753. 

Status:  Rare  and  obscure;  possibly 
endangered. 

Distribution:  Daggett  iWieholdt 
1460 A  [utc])  and  Grand  (Maguire 
1935)    counties,  Utah;   circumboreal. 

Notholaena  jonesii  Maxon,  Amer.  Fern. 
J.  7:    108.  1917. 

Status:  Restricted  and  rare;  possibly 
threatened. 

Distribution:  Washington  Co.,  Utah 
(Maxon  1917);  Arizona  and  southern 
California  (Flowers  1944;  Cronquist  et 
al.  1972). 

Portulacaceae 

Calyptridium  monandrum  Nutt.  ex  Torr. 
&  Gray,  Fl.  N.  Amer.  1:   198.  1838. 

Status:  Rare  and  restricted;  possibly 
threatened. 

Distribution:  Washington  Co.,  Utah 
(bry);  also  Arizona  and  California, 
and  Baja  California,  Mexico. 

Talinum  valid ulum  Greene,  Leafl.  Bot. 
Observ.  Crit.  2:  270.  1912. 

Status:  Rare  and  obscure;  possibly 
threatened. 

Distribution:  Emery  Co.,  Utah 
(bry);   Coconino  Co.,  Arizona. 

Primulaceae 

Primula  incana  M.  E.  Jones,  Proc.  Calif. 
Acad.  Sci.  II,  5:  706.  1895. 

Type:  Beaver  Co-op  Ranch,  at  the 
head  of  the  South  Fork  of  the  East  Fork 
of  the  Sevier  Ri^cr,  Garfield  Co.,  Utah, 
in  cold  bogs,  7,000  feet,  M.  E.  Jones 
531 2av  (i'om). 

Status:  Rare  and  possibly  extir- 
pated in  the  type  area;  threatened. 


Dec.  1975 


WELSH,   ET   AL.:    ENDANGERED   UTAH   PLANTS 


367 


Distribution:  Daggett  and  Garfield 
counties,  Utah  (Cosgriff  1968);  wide- 
spread in  northwestern  North  America 

(Welsh  1974b). 

Primula  maguirei  L.  0.  Williams,  Amer. 
Midi.  Naturalist  17:    747.   1936. 

Type:  Damp  overhanging  rock 
ledges  and  cracks,  5  miles  up  Logan 
Canyon,  Cache  Co.,  Utah,  19  Apr  1932, 
Maguirc  &  Maguire  3650   (mo). 

Status:  Endemic,  rare  and  threat- 
ened (Riple}^  T) . 

Distribution:      Cache      Co.,      Utah 

(UTC). 

Primula  specuicola  Rydb.,  Bull.  Torrey 
Bot.  Club  40:  461.  1913. 

Type:  Along  the  San  Juan  River  near 
Bluff,  San  Juan  Co.,  Utah,  25-29  Aug 
1911,  Rydberg  9882  (ny). 

Status:  Restricted  habitatwise,  local 
and  threatened   (Ripley,  T). 

Distribution:  Garfield  (ut).  Grand, 
Kane,  San  Juan,  and  Wayne  counties, 
Utah,  and  in  northern  Arizona  (Cos- 
griff 1968;  McDougall  1973). 

Ranunculaceae 

Aquilegia  micrantha  Eastw.,  Proc.  Calif. 
Acad.  Sci.  II,  4:  559.  1895. 

Type:  Near  Bluff,  San  Juan  Co., 
Utah,  Jul   1894,   Wetherill  s.n.    (cas). 

Status:  Restricted  habitatwise,  local 
but  not  threatened  nor  endangered. 

Distribution:  Emery,  Garfield, 
Grand,  Kane,  and  San  Juan  counties, 
Utah  (bry)  ;  also  in  Arizona  and  Colo- 
rado. 

Ranunculus  acriformis  A.  Gray  var.  aesti- 
valis L.  Benson,  Amer.  Midi.  Natural- 
ist 40:  43.  1948. 

Type:  Meadow  at  springs  just  east 
of  U.S.  Highway  89  and  300  yards  west 
of  the  Sevier  River,  8.3  miles  north  of 
the  principal  intersection  in  Panguitch 
and  about  1.5  miles  south  of  the  inter- 
section with  Utah  Highway  20  leading 
to  Parowan,  Garfield  Co.,  Utah,  29  Aug 
1948,  6,400  feet,  Benson  13420  (pom). 

Status:  Endemic  and  presumed  to  be 
extinct  (Ripley,  PoEx). 

Distribution:  Garfield  Co.,  Utah 
(bry)  ;  known  only  from  the  type  lo- 
cality. 


ROSACEAE 

(Viamacrhodos  erecta  (L).  Bunge  in 
Ledcb.,  Fl.  Altaica  1:  430.  1829,  based 
on  Sihbaldia  erecta  L.,  Sp.  PI.  1:  284. 
1753. 

Status:  Rare  and  local,  altitudinally 
restricted. 

Distribution:  Piute  and  Wayne 
counties,  Utah  (bry)  ;  also  from  Colo- 
rado, North  Dakota  and  Michigan, 
north  to  Yukon  and  Alaska;  Asia 
(Welsh  1974b). 

Crataegus  chrysocarpa  Ashe,  North  Caro- 
lina Agric.  Exp.  Sta.  Techn.  Publ.  175: 
110.  1900,  based  on  C.  rotundifolia 
Moench,  Verz.  Ausl.  Baume  Stand 
Weiss  29.  1785,  not  Lam. 

Status:  Rare  and  threatened. 

Distribution:  Cache  Co.,  Utah 
(Maguire  1937);  widespread  elsewhere. 

Crataegus  succulenta  Schrader  ex  Link, 
Handbuch2:  78.  1831. 

Status:   Local  and  rare;  threatened. 
Distribution:       Utah      Co.,      Utah 
(bry,   UTC;   Barnes   1943);   widespread 
to  the  east  of  Utah  (Little  1953). 

Ivesia  sabulosa  (M.  E.  Jones)  Keck, 
Lloydia  1:  124.  1938,  based  on  Ponten- 
tilla  sabulosa  M.  E.  Jones,  Proc.  Calif. 
Acad.  Sci.  II,  5:  680.  1895. 

Type:  Head  of  the  Sevier  River,  prob- 
ably in  Garfield  Co.,  Utah,  11  Sep 
1894,  8,000  feet,  M.  E.  Jones  6032 
(pom). 

Status:  Rare  and  local,  possibly 
threatened. 

Distribution:  Garfield  and  Wash- 
ington counties,  Utah  (bry),  and  Nye 
Co.,  Nevada  (Keck  1938b). 

Ivesia  utahensis  S.  Wats.,  Proc.  Amer. 
Acad.  Arts  17:   371.  1882. 

Type:  On  the  summit  of  Bald  Moun- 
tain, in  Wasatch  Range,  above  Alta, 
Salt  Lake  Co.,  Utah,  Aug  1879,  over 
12,000  feet,  M.  E.  Jones  1231   (gh). 

Status:    Endemic,  rare  and  local. 

Distribution:  Salt  Lake,  Summit, 
and  Utah  counties,  Utah  (Keck  1938b). 

Rubus  neomexicanus  A.  Gray,  Smithso- 
nian Contr.  Knowl.  5:  55.  1853. 

Status:   Local,  rare  and  threatened. 

Distribution:  San  Juan  Co.,  Utah 
(bry)  ;  New  Mexico,  Arizona,  and 
northern  Mexico. 


368 


GREAT   BASIN   NATURALIST 


Vol.  35,  No.  4 


Most  of  the  known  range  of  this 
species  has  been  destroyed  by  Lake 
Powell. 

RUBIACEAE 

Galium  multiflorum  Kellogg  var.  wat- 
sonii  A.  Gray,  Syn.  Fl.  N.  Amer.  1 :  40. 
1884. 

Type:  Wasatch  Mts.,  Utah,  1869, 
Watson  484  (gh)  . 

Status:  Endemic,  neither  threatened 
nor   endangered. 

Distribution:  Box  Elder,  Cache, 
Davis,  Tooele,  and  Utah  counties,  Utah 
(Dempster  &  Ehrendorfer  1965). 

Galium  scabruisculum.  (Ehrendorfer) 
Dempster  &  Ehrendorfer,  Brittonia  17: 
312.  1965,  based  on  G.  hypotrichium 
ssp.  scabjuisculum  Ehrendorfer,  Contr. 
Dudley  Herb.  5:   13.  1956. 

Type:  Calf  Springs  Wash,  San  Rafael 
Swell,  Emery  Co.,  Utah,  Maguire 
18457  (gh). 

Status:  Endemic,  neither  threatened 
nor  endangered. 

Distribution:  Carbon  and  Emery 
counties,  Utah. 

RUTACEAE 

Ptelea  trifoliata  L.  ssp.  pallida  (Greene) 
V.  L.  Bailey,  Brittonia  14:  23.  1962, 
based  on  P.  pallida  Greene,  Contr.  U.  S. 
Natl.  Herb.  10:  70.  1906. 

Status:  Rare  and  possibly  extirpated. 
Distribution:     Garfield    and    Kane 
counties,  Utah;   this  subspecies  also  in 
Arizona  and  Colorado  (Bailey  1962). 

SCROPHULARIACEAE 

Castilleja  aquariensis  N.  H.  Holmgren, 
Bull.  Torrey  Bot.  Club  100:  87.  1973. 

Type:  Aquaris  Plateau,  22  miles 
northwest-north  of  Escalante  on  the 
road  to  Bicknell,  0.5  mile  north  of  Clay- 
ton Guard  Station  turnoff,  Garfield  Co., 
Utah,  11  Aug  1970,  9,600  feet,  Holm- 
gren &  Holmgren  4726  (ny). 

Status:  Endemic,  rare  and  local;  en- 
dangered  (Ripley,  E). 

Distribution:  Garfield  Co.,  Utah; 
known  only  from  the  type  area. 

Castilleja  leonardii  Rydb.,  Bull.  7V)rrey 
Bot.  Club  34:  36.  1907. 

Type:   Head  of  American  Fork  Can- 


yon, Utah  Co.,  Utah,  1885,  Leonard 
151   (ny). 

Status:  Endemic,  locally  common 
but  neither  threatened  nor  endangered. 

Distribution:  Cache,  Daggett,  Davis, 
Duchesne,  Salt  Lake,  Sanpete,  Sununit, 
Tooele,  Utah,  and  Wasatch  counties, 
Utah  (bry,  ut)  . 

Castilleja  parvula  Rydb.,  Bull.  Torrey 
Bot.  Club  34:  40.  1907. 

Type:  Mountains  north  of  Bullion 
Creek  near  Marysvale,  Piute  Co.,  Utah, 
1905,  Rydberg  &  Carlton  7158  (ny). 

Status:  Endemic,  rare  and  local; 
threatened  (Ripley,  T). 

Distribution:  Piute  Co.,  Utah  (ny, 
utc). 

Castilleja  revealii  N.  H.  Holmgren,  Bull. 
Torrey  Bot.  Club  100:  87.  1973. 

Type:  Bryce  Canyon  N.P.,  along  the 
road  to  Bryce  Point,  0.5  mile  from  In- 
spiration Point  turnoff,  Garfield  Co., 
Utah,  24  Jun  1965,  8,000  feet,  Holm- 
gren &  Reveal  2017  (ny). 

Status:  Endemic,  rare  and  local;  en- 
dangered  (Ripley,  E). 

Distribution:  Garfield  Co.,  Utah; 
known  only  from  the  type  locality. 

Mimulus  eastwoodiae  Rydb.,  Bull.  Torrey 
Bot.  Club  40:   483.  1913. 

Type:  In  crevices  of  perpendicular 
or  overhanging  rocks  along  the  San 
Juan  River  near  Bluff,  25-29  Aug  1911, 
Rydberg  9883  (ny). 

Status:  Edaphically  restricted,  local 
and  disjunct;  not  threatened  nor  endan- 
gered. 

Distribution:  Grand,  Kane,  and  San 
Juan  counties,  Utah  (bry),  and  adja- 
cent northern  Arizona  (Kearney  & 
Peebles  1951). 

Penstemon  abietinus  Pennell,  Contr.  U.S. 
Natl.  Herb.  20:  276.  1920. 

Type:  Ireland  Ranch,  head  of  Salina 
Canyon,  Sevier  Co.,  Utah,  15  Jun  1894, 
2,400  m,  M.  E.  Jones  5440  (us). 

Status:  Endemic,  rare  and  local; 
threatened  (Ripley,  T). 

Distribution:  Iron  (ut),  Sevier,  and 
Utah  counties,  Utah  (bry;  Keck  1937a). 

Penstemon  acaulis  L.  O.  Williams,  Ann. 
Missouri  Bot.  Card.  21:  345.  1934. 

Status:  Restricted,  rare  and  threat- 
ened (Ripley,  T). 


Dec.    1975 


WELSH,   F,T   AL.:    ENDANGERED    UTAH    PLANTS 


369 


Distribution:  Daggett  Co.,  Utah 
(ny,  us,  utc)  and  adjacent  vSweetwater 
Co.,  Wyoming  (Keck  1937a). 

Penstemon  atwoodii  Welsh,  Great  Basin 
Nat.  35:  378.  1976. 

Type:  South  end  of  Horse  Mtn.,  ca 
10  miles  south-southeast  of  Canaan 
Peak,  Kane  Co.,  Utah,  14  Jun  1975, 
S.  L.  &  S.  L.  Welsh  12820  (bry). 

Status:  Endemic,  local  and  threat- 
ened. 

Distribution:  Garfield  and  Kane 
counties,  Utah  (bry,  ny). 

Penstemon  bracteatus  Keck,  Leafl.  W.  Bot. 
1:  82.  1934. 

Type:  Red  Canyon,  Garfield  Co., 
Utah,  20  Jun  1933,  Eastwood  &  Howell 
783   (cAs). 

Status:  Endemic,  restricted  and  rare; 
possibly  threatened. 

Distribution:  Garfield  Co.,  Utah 
(bry). 

Penstemon  caespitosus  Nutt.  var.  suffruti- 
cosus  A.  Gray,  Syn.  Fl.  N.  Amer.  2: 
270.  1878. 

Type:  Near  Beaver,  Beaver  Co.,  Utah, 
1877,  Palmer  s.n.   (gh). 

Status:  Endemic,  restricted  and  lo- 
cal; threatened   (Ripley,  T). 

Distribution:  Beaver,  Garfield  and 
Piute  counties,  Utah  (Keck  1937a). 

Penstemon  compactus  (Keck)  Crosswhite, 
Amer.  Midi.  Naturahst  77:  6.  1967, 
based  on  P.  cyananthus  ssp.  compactus 
Keck,  Amer.  Midi.  Naturalist  23:  615. 
1940. 

Type:  Stony  slopes  of  Mt.  Naomi, 
Cache  Co.,  Utah,  18  Aug  1938,  2,900 
'meters,  Maguire  16148  (utc). 

Status:  Endemic,  restricted  and 
threatened  (Ripley,  T). 

Distribution:  Cache  Co.,  Utah  (bry, 
UTC,  w^sco). 

Penstemon  concinnus  Keck,  Amer.  Midi. 
Naturalist  23:  608.  1940. 

Type:  Tunnel  Springs,  northwest 
corner  of  Desert  Range  Experiment  Sta- 
tion boundary,  about  10  miles  east  of 
Garrison,  Millard  Co.,  Utah,  28  Jun 
1933,  1,675  meters,  Cottam  5655   (ds). 

Status:  Endemic,  restricted  and  rare; 
endangered    (Ripley,  E). 

Distribution:  Beaver  and  Millard 
counties,  Utah  (bry,  ny,  us,  utc). 


Penstemon  garrettii  Pennell,  (^ontr.  U.S. 
Natl.  Herb.  20:  353.  1920. 

Type:  Crevices  in  travertine  rock, 
"Hot  Pots,"  near  Midwav,  Wasatch  Co., 
Utah,  6  Jul  1905,  Carlrton  &  Garrett 
6697  (ny). 

St.'Vtus:  Endemic,  restricted  and  pos- 
sibly extinct. 

Distribution:  Duchesne  (utj  and 
Wasatch    counties,    Utah. 

Penstemon  grahamii  Keck  in  Graham, 
Ann.  Carnegie  Mus.  26:   331.  1937. 

Type:  Talus  slope  on  the  west  side 
of  Green  River,  south  of  the  mouth  of 
Sand  Wasli,  Uintah  Co.,  Utah,  27  May 
1933,  Graham  7883   (cm). 

Status:  Endemic,  rare,  restricted 
and  endangered  (Ripley,  E) . 

Distribution:  Uintah  Co.,  Utah 
(bry,  utc;  Keck  1938a). 

Penstemon  humilis  Nutt.  ex  Gray  var. 
brevifolius  A.  Gray,  Syn.  Fl.  N.  Amer. 
2:  267.  1878. 

Type:  Cottonwood  Canyon,  Wasatch 
Mts.,  Salt  Lake  Co.,  Utah,  1869,  9,000- 
10,000  feet,  Watson  781  (gh). 

Status:  Endemic,  local  and  possibly 
threatened. 

Distribution:  Cache,  Juab,  Salt 
Lake,  Utah,  and  Weber  counties,  Utah 
(bry,  ut). 

Penstemon  humilis  Nutt.  ex  Gray  var. 
obtusifolius  (Pennell)  Reveal,  stat.  no  v., 
based  on  P.  obtusifolius  Pennell,  Contr. 
U.S.  Natl.  Herb.  20:   370.  1920. 

Type:  Springdale,  Washington  Co., 
Utah,  16  May  1894,  1,600  meters,  M. 
E.  Jones  5249am  (pom). 

Status:  Endemic,  rare  and  restricted; 
possibly  threatened. 

Distribution:  Beaver  and  Washing- 
ton counties,  Utah  (Keck  1945). 

Penstemon  jonesii  Pennell,  Contr.  U.S. 
Natl.  Herb.  20:  338.  1920. 

Type:  Springdale,  Washington  Co., 
Utah,  17  May  1894,  M.  E.  Jones  5250, 
in  part  (us). 

Status:  Endemic,  rare  and  obscure; 
taxonomic  status  questionable. 

Distribution:  'Washington  Co., 
Utah;  known  only  from  the  tj^e  lo- 
cality. 

Penstemon  laevis  Pennell,  Contr.  U.S. 
Natl.  Herb.  20:  347.  1920. 


370 


GREAT   BASIN    NATURALIST 


Vol.  35,  No.  4 


Type:  Red  sand  at  Springdale,  Wash- 
ington Co.,  Utah,  17  May  1894,  1,200 
meters,  M.  E.  Jones  5250^  in  part  (us). 

Status:  Endemic,  restricted  and  lo- 
cal. 

Distribution:  Kane,  Garfield,  and 
Washington  counties,  Utah  (bry,  utc)  . 

Penstemon  leiophyllus  Pennell,  Contr. 
U.S.  Natl.  Herb:  20:   346.  1920. 

Type:  Mammoth  Creek,  Garfield 
Co.,  Utah,  10  Sep  1894,  2,400  meters, 
M.  E.  Jones  6026b   (us). 

Status:  Endemic,  restricted  and  lo- 
cal; threatened    (Ripley,  T). 

Distribution:  Garfield,  Iron,  Kane, 
and  Washington  counties,  Utah   (bry). 

Penstemon  lentus  Pennell  var.  albiflorus 
(Keck)  Reveal,  stat.  nov.,  based  on  P. 
lentus  ssp.  albiflorus  Keck,  Amer.  Midi. 
Naturalist  23:  616.  1940. 

Type:  Abajo  Mts.,  ca  8  miles  west  of 
Blanding,  near  the  "Bear's  Ears,"  San 
Juan  Co.,  Utah,  9  Jun  1938,  2,400 
meters,  C.  L.  Porter  1801   (rm). 

Status:  Endemic,  locally  common 
and  not  threatened. 

Distribution:  San  Juan  Co.,  Utah 
(bry). 

Penstemon  leonardii  Rydb.,  Bull.  Torrev 
Bot.  Club  40:  483.  1913. 

Type:  Diehl's  Grove,  Wasatch  Mts., 
possibly  Utah  Co.,  Utah,  1  Aug  1884, 
Leonard  s.n.   (ny). 

Status:  Endemic,  locally  common 
and  not  threatened  nor  endangered. 

Distribution:  Cache,  Davis,  Rich, 
Salt  Lake,  Summit,  Uintah,  Utah,  Wa- 
satch, Washington,  and  Weber  counties, 
Utah  (bry.  ut). 

Penstemon  longiflorus  (Pennell)  S.  L. 
Clark,  Great  Basin  Nat.  35:  434.  1976, 
based  on  P.  cyananthus  ssp.  longiflorus 
Pennell,  Contr.  U.S.  Natl.  Herb.  20: 
353.  1920. 

Type:  Beaver,  Beaver  Co.,  Utah, 
Palmer  376  (ny). 

Status:  Endemic,  restricted  but  lo- 
cally common. 

Distribution:  Beaver,  Millard,  and 
Piute  counties,  Utah  (bry). 

Penstemon  nanus  Keck,  Amer.  Midi.  Nat- 
uralist 23:   607.  1940. 

Type:  Desert  Range  Experiment  Sta- 
tion, about   10  miles  east  of  Garrison, 


Millard  Co.,  Utah,  13  May  1939,  1,675 
meters,   Plummer  7313    (ds). 

Status:  Endemic,  restricted  and  lo- 
cal; threatened   (Ripley,  E). 

Distribution:  Beaver  and  Millard 
counties,  Utah   (bry,  utc). 

Penstemon  parvus  Pennell,  Contr.  U.S. 
Natl.  Herb.  20:    345.   1920. 

Type:  The  Button,  Aquarius  Plateau, 
Wayne  Co..  Utah,  11  Aug  1875,  Ward 
546  (us). 

Status:  Endemic,  restricted  and  rare; 
threatened  (Ripley,  T). 

Distribution:  Garfield  and  Wayne 
counties,  Utah,    (bry;   Pennell    1920). 

Penstemon  sepalulus  Rydb.,  Bull.  Torrey 
Bot.  Club  36:  690.  1909. 

Type:  Canyons  of  the  Wasatch  Mts., 
Provo  Canyon,  Utah  Co.,  Utah,  Jul 
1869,  Watson  786  (ny). 

Status:    Endemic,  locally  abundant. 

Distribution:  Utah  Co.,  Utah  (bry; 
Keck  1932). 

Penstemon  tidestromii  Pennell,  Contr. 
U.S.  Natl.  Herb.  20:   379.  1920. 

Type:  "XL"  Canyon,  San  Pitch  Mts., 
Sanpete  Co.,  Utah,  24  Jun  1908,  1,650 
meters,  Tidestrom  1296  (us). 

Status:  Endemic,  local  and  obscure; 
taxonomic  status  questionable. 

Distribution:  Sanpete  Co.,  Utah; 
known  only  from  the  type  locality. 

Penstemon  uintahensis  Pennell,  Contr. 
U.S.  Natl.  Herb.  20:   350.  1920. 

Type:  Dyer  Mine,  Uinta  Mts.,  Uin- 
tah Co.,  Utah,  50  Jun  1902,  Goodding 
1221   (ny). 

St.'vtus:  Endemic,  rare  and  restric- 
ted; threatened    (Ripley,  T). 

Distribution:  Daggett.  Duchesne, 
and  Uintah  counties,  Utah   (bry). 

Penstemon  wardii  A.  Gra^'.  Proc.  Amer. 
Acad.  Arts  12:    82.   1876. 

Type:  Near  Glenwood,  Sevier  Co., 
Utah,  4  Jun   1875,   Ward  162    (us). 

Status:  Endemic,  restricted  and  lo- 
cal;  threatened    (Ripley,  T). 

Distribution:  Sanpete  and  Sevier 
counties,  Utah   (bry.  utc). 

Synthris  laciniata  (A.  Gray)  Rydb.  ssp. 
ibapahensis  Pennell,  Proc.  Acad.  Nat. 
Sci.   Philadelphia   85:  -92.    1933. 

Type:  Wot  gravelly  slopes  near  ra- 
vine of  snow   on   Mount  Ibapah,  Juab 


Dec.  1975 


WELSH,   ET   AL.:    ENDANGERED    UTAH    PLANTS 


371 


Co.,  Utah,  5  Jul  1932,  9,500-10,000 
feet,  Stanton  1000  (ph). 

Status:  Endemic,  restricted  iind  ob- 
scure;   taxonomic   status   questionable. 

Distribution:  Juab  Co.,  Utah; 
known  only  from  the  type  locality. 

SELAGINELL.A.CEAE 

Selaginella  utahensis  Flowers,  Amer.  Fern 
J.  39:  83.  1949. 

Type:  South  of  St.  George,  Washing- 
ton Co.,  in  a  wash  bottom,  5  Apr  1931, 
Cottam  5644  (ut). 

Status:  Rare  and  local;  possibly 
threatened. 

Distribution:  Washington  Co., 
Utah,  and  Spring  Mts.,  Clark  Co.,  Ne- 
vada  (Cronquist  et  al.  1972). 

Verbenaceae 

Aloysia  wrightii  (A.  Gray)  A.  A.  Heller, 
Muhlenbergia  1:  147.  1906,  based  on 
Lippa  wrightii  A.  Gray,  Amer.  J.  Sci. 
II,  16:  98.  1853. 

Status:  Local  and  rare;  possibly  en- 
dangered. 

Distribution:  Washington  Co.,  Utah 
(Higgins  615  [bry];  Higgins  1972b); 
from  Texas  to  California  and  in  north- 
ern Mexico. 

Violaceae 

Viola  purpurea  Kellogg  var.  charlestonen- 
sis  (Baker  &  Clausen)  Welsh  &  Reveal, 
Stat.  &  comb,  nov.,  based  on  V .  charles- 
tonensis  Baker  &  Clausen  in  Clokey, 
Madrono  8:  58.  1945. 

Status:  Restricted,  local  and  rare; 
threatened  (Ripley,  T). 

Distribution:  Washington  Co., 
Utah,  and  Clark  Co.,  Nevada  (Clokey 
1945;  Eastmond  1969). 

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hitchcockii.  Great  Basin  Nat.   30:   94-98. 
.     1972a.     Five  new  varieties  of  Eriogonum 

(Polygonaceae).  Great  Basin  Nat.  32:  113-116. 


.      1972b.     Descriptions  of  new  species  and 

combinations   in   Eriogonum    (Polygonaceae). 

Phytologia  23:   163-178. 
.     11937a.     Eriogonum     (Polygonaceae)     of 

Utah.    Phytologia    25:     169-217. 
.     1973b.     Statement    of   James    L.    Reveal, 

assistant    professor    of    botany.    University    of 

Maryland.  College  Park,  Md.  National  Wild- 
life Refuges,  HR  7983.  Fish  &  Wildlife  Mis- 
cell.   Ser.   93-18(2):    406-416. 
.      1974.     Two  new  varieties  of  Eriogonum 

(Polygonaceae)    from   the   intermountain   re- 
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Ripley,  S.  D.     1975.     Report  on  endangered  and 

threatened  species  of  the  United  States.  House 

Document  94-51:   1-200. 
Rollins,     R.     C.     1938.     Glaucocarpum,     a     new 

genus  in  the  Cruciferae.  Madrono  4:  232-235. 
.     1940.     iStudies  in  the  genus  Hedysarum 

in  North  America.  Rhodora  42:   217-239. 
.     1941  a.     Monographic  study  of  Arabis  in 

western  North  America.  Rhodora  43:  289-485. 
.      1941b.     Some  new  or  noteworthy  North 

American  crucifers.   Contr.   Dudley   Herb.   3: 

174-184. 
.     1953.     Draba  on  Clay  Butte.  Wyoming. 

Rhodora  55:  229-235. 
.     1971.     Protogyny  in  the  Cruciferae  and 

notes  on  Arabis  and  Caulanthus.  Contr.  Gray 

Herb.  201:   3-10 
Rollins,    R.    C,   and    E.    A.    Shaw.     1973.     The 

genus     Lesquerella     (Cruciferae)     in     North 

America.     Harvard    University    Press,    Cam- 
bridge. 288  pp. 
Rydberg,    p.    a.      1900a.     Studies    on    the    Rocky 

Mountain    flora — I.    Bull.    Torrey    Bot.    Club 

27:    169-189. 
.     1900b.     Studies  on  the  Rocky  Mountain 

flora— III.   Bull.   Torrey  Club  27:    614-636. 
.     1902a.     Studies  on  the  Rocky  Mountain 

flora— VII.   Bull.   Torrey  Bot.   Club  29:    145- 

160. 
.     1902b.     Studies  on  the  Rocky  Mountain 

flora— IX.    Bull.    Torrey    Bot.    Club   29:    680- 

'693. 
.     1903.     Studies   on    the   Rocky   Mountain 

flora— X.  Bull.  Torrey  Bot.  Club  30:  247-262. 
.      1907a.     Studies  on  the  Rocky  Mountain 

flora— XVII.  Bull.  Torrey  Bot.  Club  34:   35- 

50. 
.     1907b.     Studies  on  the  Rocky  Mountain 

flora— XVIII.    Bull.    Torrey    Bot.    Club    34: 

417-437. 
.     1909.     Studies   on   the   Rocky   Mountain 

iflora— XX.   Bull.   Torrey   Bot.   Club   36:    675- 

698. 
.     1912a.     Studies  on  the  Rocky  Mountain 

flora— XXVI.  Bull.  Torrey  Bot.  Club  39:  99- 

111. 
.      1912b.     Studies  on  the  Rocky  Mountain 

flora— XXVII.    Bull.    Torrey    Bot.    Club    39: 

301-328. 
.     1913a.     'Studies  on  the  Rocky  Mountain 

flora— XXVIII.    Bull.    Torrey   Bot.    Club   40: 


43-74. 
— .     1913b. 


Studies  on  the  Rocky  Mountain 


flora— XXIX.  Bull.  Torrey  Bot.  Club  40:  461- 
485. 

— .     1915.     Gaillardia.     N.     Amer.     Fl.     34: 
131-140. 


376 


GREAT   BASIN    NATURALIST 


Vol.  35,  No.  4 


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ScHREiNER.  K.  M.  1975.  Threatened  or  endan- 
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UTAH   PLANT   NOVELTIES    IN 
CYMOPTERUS  AND  PENSTEMON 

Stanley     L.     Welshi 

Abstract. —  Cymopterus  higginsii  nnd  Penstetnon  atwoodii  are  named  and  described  from  ma- 
terials collected  in  the  Kaiparowits  Plateau  vicinity  of  eastern  Kane  County.  Utah.  Habitat,  distri- 
bution,  and  probable  affinities  are  outlined. 


Examination  of  specimens  obtained 
from  the  Kaiparowits  Plateau  region  of 
eastern  Garfield  and  Kane  counties  in 
southern  Utah  has  revealed  the  existence 
of  two  previously  undescribed  entities,  one 
each  in  Cymopterus  (Apiaceae)  and 
Penstemon  (Scrophulariaceae).  Existence 
of  these  taxa  is  not  surprising  when  one 
considers  the  historic  remoteness  of  much 
of  that  great  region.  The  discovery  and 
the  extent  of  the  range  of  each  taxon 
must  be  credited  to  the  extensive  field  work- 
allowed  under  the  baseline  studies  of  the 
Navajo-Kaiparowits  enA'ironmental  pro- 
ject directed  by  personnel  from  Brigham 
Young  University  (Welsh,  Murdock,  and 
Wood  1975). 

The  Cymopterus  species  is  known  from 
saline  soils  of  the  Tropic  Shale  formation 
and  associated  pedimental  gravels  on  fans 
and  bajadas  below  the  Straight  Cliffs  for- 
mation in  that  portion  of  Kane  County 
from  the  Paria  River  eastward  to  the  Last 
Chance  Creek  vicinity.  Apparent  rela- 
tionships of  C.  higginsii  seem  to  lie  with 
C.  fendleri  from  which  it  differs  inter  alia 
in  the  rose  to  purple  flowers  with  evident 
pedicels  and  wider  wings  on  the  fruit. 
The  pseudoscape  is  poorly  developed. 

The  corollas  in  the  Penstemon  species 
are  glandular  hairy  externally,  and  the 
taxon  seems  to  belong  with  those  species 
treated  by  Pennell  (1920)  as  Section 
Crista ti  and  by  Keck  (1938)  illegitimately 
as  Section  Aurator.  The  nearest  ally  ap- 
pears to  be  P.  jamesii  from  which  P.  at- 
woodii differs  as  noted  in  the  diagnosis. 
P.  atwoodii  is  known  only  from  middle 
elevations  of  the  Kaiparowits  region, 
where  it  grows  on  the  Cretaceous  forma- 
tions in  juniper-piny  on  woodland. 

Both  species,  the  Cymopterus  and  the 
Penstemon,  are  plants  of  very  restricted 
range.  They  are  in  areas  which  are  now 
subject   to   commercial   exploitation,    and 


both  should   be  considered  as  threatened 
species. 

Cymopterus  higginsii  Welsh  sp.  nov. 

Plantae  acaulescentes  non  caespitosae 
pseudoscapis  non  vel  non  nisi  evolutis  in- 
fermis,  pubescentes  parse  pili  complanti: 
folia  ovata  vel  subelliptica  in  circum- 
scriptem,  laminis  1.8-7.7  cm  longis  1.5- 
6.0  cm  latis  hi-  vel  tripinnatis  vi- 
ridibus  foliolosis  longior  quam  latis 
pinnatis  ad  bipinnata,  lobi  obtusi  ad  ro- 
tundatos  vel  acutos  raro,  petiolis  1.8-14 
cm  longis;  pedunculi  folia  longior  ad  ex- 
tremum  2-12  cm  longi,  purpura scentes; 
involucrum  vaginans  margine  scariosa; 
involucellum  bracteolarum  brevior  quam 
floras,  lobis  aliquot  dentatis  acutis  vel 
acuminatis;  umbellae  compactae,  radiis 
3-5,  1-10  mm  longis,  umbellula  centrali 
sessili;  pedicelli  1-6  mm  longi;  flores 
rosei  ad  purpurascens;  fructus  ovalis  ad 
ellipticum  7-10  mm  longus  5-8  mm  latus, 
alis  corpus  subaequalis  incrassatis  spon- 
giosis. 

C.  fendleri  affinis  sed  floribus  roseis  ad 
purpurascens  pedicellis  evidentibus  et  alis 
corpus  subaequalis. 

Habitat  and  Distribution. —  Tropic 
shale  and  pedimental  covering  derived 
from  Straight  Cliffs  and  other  formations, 
on  saline  soils,  from  East  Clark  Bench 
eastward  to  Last  Chance  Canyon,  at  least 

30  miles  east  of  Glen  Canyon  City,  east- 
ern Kane  County,  Utah. 

Type:  Utah:  Kane  Co.,  Shadscale  dom- 
inated bajada,  on  gravell}'  pedimental 
fan,  east  of  None  Butte,  ca  17  miles  east 
of  Glen  Canyon  Citv,  S.  L.  Welsh  12740, 

31  May  1975  (Holotype  BRY;  Isotypes  to 
be  distributed).  Paratypes:  Utah:  Kane 
Co.,  Site  9,  Navajo-Kaiparowits  Project, 
base  of  Smoky  Mt.,  2  miles  from  Ahl- 
strom     Point    junction,     Atriplex-Kochia- 


'Department    of    Botan.v    ;ind    Range    Science,    Brigham    Young    Univeisily. 


377 


378 


GREAT   BASIN    NATURALIST 


Vol.  35,  No.  4 


Artemisia  community,  N.  D.  Atwood 
3439,  March  1972  (BRY) ;  ca  2  miles 
north  of  Church  Wells,  on  bench  between 
Coyote  Creek  and  Wahweap  Creek,  N.  D. 
Atwood  et  al.  3493,  23  March  1972 
(BRY) ;  ca  30  miles  east  of  Glen  Canyon 
City,  on  Tropic  Shale  formation,  N.  D. 
Atwood  4549,  23  April  1973  (BRY). 

The  species  is  named  to  honor  Larry 
Charles  Higgins,  student  of  Boragmaceae, 
especially  of  Cryptantha.  and  specialist  in 
western  botany  generally. 

Penstemon  atwoodii  Welsh  sp.  nov. 

Herbae  perennes  1.4-5.3  cm  altae; 
caules  pauci  vel  multi  e  caudicibus  ramifi- 
cantibus  glabri  infra  medium  ])ilis  patulis 
glanduliferis  super;  folia  glabra,  basalia 
oblanceolata  ad  spathulata  vel  ovata  ad 
elliptica  raro  Integra  2.8-9.0  cm  longa 
(0.2)  0.4-0.8  (1.4)  cm  lata,  caulina  lanci- 
linearia  ad  oblonga  vel  spathulata,  ± 
auriculata  super  Integra  vel  serrata  re- 
mota  raro,  3.0-7.0  cm  longa  0.3-0.8  (1.4) 
cm  lata;  inflorescentia  verticillastorum 
distinctarum  plurium;  bracteae  foliaceae; 
calyces  6.5-8.5  mm  longi  lobis  lanceolatis 
herbaceis  purpureis  pilis  glanduliferis; 
corollae  pilis  glanduliferis  externis  cyanae 
vel  cyano-caesiae  13-16  mm  longae  ex- 
pansae  distales  5-6  mm  latae  glabrae  intus 
praeter  ad  orficium  labium  inferum;  an- 
therarum  thecae  glabrae  divaricatae  vel 
explanatae;  staminodium  barbatum  pilis 
luteo  lineare;  capsulae  glabrae. 

P.  jamesii  sensu  lato  affinis  sed  floribus 
parvioribus  et  glabris  intus  praeter  ad 
orficium  labium  inferum. 

Habitat  and  Distribution. —  Kaipar- 
owits,  Wahweap,  and  Straight  Cliffs  for- 


mation at  6,200  to  8,000  feet  elevation 
in  pinyon-juniper  woodland  on  the  Kai- 
parowits  Plateau  of  eastern  Garfield  and 
Kane  counties,  Utah. 

Type:  Utah:  Kane  Co.,  Gray  sand  of 
Kaiparowits  formation,  pinyon-juniper 
community,  south  end  of  Horse  Mountain, 
ca  10  miles  south-southeast  of  Canaan 
Peak,  S.  L.  &  S.  L.  Welsh  12820,  14  June 
1975  (Holotype;  BRY;  Isotypes  to  be 
distributed). 

Paratypes:  Utah:  Garfield  Co.,  Death 
Ridge  ca  16  miles  southwest  of  Escalante, 
N.  D.  Atwood  5177,  30  May  1973;  do  S. 
L.  Welsh  &  J.  R.  Murdock  12866,  28  June 
1975.  Kane  Co.,  4  miles  southeast  of  sum- 
mit of  Collets  Wash,  Kaiparowits  Plateau, 
N.  D.  Atwood  s.  n.,  19  June  1969;  Pin- 
yon-juniper woods  on  basal  Wahweap 
formation,  ca  6  miles  north  of  junction 
of  Escalante  road  with  head  of  Last 
Chance  Creek,  Kaiparowits  Plateau,  S.  L. 
Welsh  &  J.  R.  Murdock  12793,  4  June 
1975;  do  S.  L.  Welsh  &  J.  R.  Murdock 
12973a,  4  June  1975. 

This  species  is  named  to  honor  Nephi 
Duane  Atwood,  student  of  Hydrophyl- 
laceae,  field  botanist  extraordinary,  and 
first  to  recognize  the  unique  nature  of  this 
taxon. 

References 

Keck.    D.    D.     1938.     Studies    in   Penstemon    VI. 

The   section  Aurator.   Bull.   Torrev  Bot.   Club 

65:    233-255. 
Mathi.\s,     M.     E.     .\nd     L.     Constance.      1945. 

Cymopterus  Raf.  N.  Amer.  Fl.  28B:    170-183. 
Pennell,   F.   W.      1920.     Scrophulariaceac  of  the 

central   Rockv   Mountain   states.   Contr.   U.   S. 

Nat.  Herb.  20:    313-381. 
Welsh,  S.  L.,  J.  R.  Murdock,  and  B.  W.  Wood. 

1975.     Navajo-Kaiparowits  Environmental 

Baseline  Studies.  Unpublished  mss,  800  p. 


THE  ZYGOPTERA  (ODONATA)  OF  UTAH 
WITH  NOTES  ON  THEIR  BIOLOGY^ 

A.  V.  Provonsha^ 

Abstract. —  Detailed  distribution  maps  of  Utali  are  provided  for  each  of  the  33  state  species  of 
Zygoptera.  Notes  on  the  general  range,  habitat  preference,  reproductive  behavior,  emergence  data, 
and  general  biology  of  each  species  are  also  included. 


As  noted  by  Kormondy  (1957),  pub- 
lished data  on  the  geographical  distribu- 
tion of  western  Odonata  is  scarce.  Ken- 
nedy (1915)  gave  a  partial  list  of  the 
Odonata  of  Washington  and  Oregon,  and 
in  1917  he  published  records  from  central 
California  and  Nevada.  The  Washington 
list  has  recently  been  updated  by  Paulson 
(1970)  and  a  complete  list  and  keys  to 
the  California  Odonata  is  given  by 
Smith  and  Pritchard  (1956).  Bick  and 
Hornuff  (1972)  published  man}^  new 
Odonata  records  for  northwestern  Wyo- 
ming. Although  Larsen  (1952)  and  Mus- 
ser  (1962)  added  considerably  to  our 
knowledge  of  Utah  Anisoptera,  only  one 
paper  (Brown  1934)  dealing  exclusively 
with  the  Odonata  of  Utah  included  a 
treatment  of  the  Zygoptera.  Brown's  re- 
port consisted  of  an  annotated  checklist 
including  some  25  currently  valid  zygop- 
teran  species. 

During  the  summers  of  1970  and  1971 
the  state  of  Utah  was  extensively  collected 
in  an  effort  to  gain  an  increased  under- 
standing of  the  distribution  and  species 
composition  of  the  damselfly  fauna  of  that 
area.  Some  152  localities  throughout  the 
state  were  sampled,  and  the  collections  at 
the  University  of  Utah,  Utah  State  Uni- 
versity, Brigham  Young  University,  and 
Dixie  College  were  examined.  As  a  result, 
the  list  of  Zygoptera  known  to  occur  in 
Utah  has  been  expanded  to  33.  In  ad- 
dition, field  work  and  rearing  has  added 
considerably  to  our  knowledge  of  the  bi- 
ologv  of  many  species  (also  see  Provonsha 
and  McCafferty  1973). 

No  attempt  has  been  made  to  cite  in  de- 
tail all  collection  records.  Although  county 
records  may  be  adequate  for  some  states 
where  counties  are  numerous  and  rela- 
tively small,  they  are  meaningless  for 
Utah,  where  several  counties  contain  over 


Fig.    1.     County  map  of  Utah. 

5,000  square  miles  and  in  many  cases 
more  than  one  distinct  biotic  region.  In- 
stead, distributions  based  on  all  collecting 
localities  known  to  the  author  are  plotted 
for  each  species.^  For  those  instances 
where  counties  are  mentioned  in  the  text, 
the  reader  may  refer  to  Fig.  1  for  their 
specific  location.  Where  possible,  notes 
on  general  distribution,  habitat  prefer- 
ences, emergence  data,  and  reproductive 
behavior  are  included  in   the   text. 

For  keys  to  the  zygopteran  species 
known  to  occur  in  Utah,  the  author  re- 
commends the  following  publications: 
Johnson  (1972),  Smith  and  Pritchard 
(1956),   and  Walker    (1953). 

^Complete  collecting  data  may  be  obtained 
from  the  author  upon  request. 


^Published   with    the   approval    of   the    Director    of   the    Purdue     University     Agricultural      Experimeni 
Series  No.   6034. 

^Department   of   Entomology,    Purdue   University,    West   Lafayette,     Indiana    47907. 


Station     as     Journal 


379 


380 


GREAT   BASIN   NATURALIST 


Vol.  35,  No.  4 


Calopteryx  aequabilis  Say,  1839 
Fig.  2 

This  species  is  known  to  occur  through- 
out most  of  Canada  east  of  Saskatchewan 
and  the  north  central  and  northeastern 
United  States,  with  isolated  pockets  in 
Colorado,  California,  and  the  Columbia 
River  drainage  system.  The  major  popula- 
tion of  C.  aequibilis  in  Utah  is  found  at 
the  Raft  River,  Box  Elder  Co.,  which  is 
the  only  major  river  in  Utah  flowing 
north  as  part  of  the  Columbia  River  drain- 
age system.  One  other  small  isolated  pop- 
ulation occurs  at  Far  West,  Weber  Co. 
A  single  male  was  collected  at  Goshen, 
Utah  Co.,  on  19  May  1969,  by  P.  V. 
Winger.  Subsequent  collections  at  that 
locality  have  failed  to  provide  additional 
specimens,  and  it  is  doubtful  that  a  popu- 
lation is  established  there.  The  species 
is  restricted  to  streams  where  the  nymphs 
cling  to  debris  and  submerged  roots  along 
the  banks.  The  above  record  from  Goshen 
is  the  earliest  Utah  record  I  have;  species 
have  been  taken  at  the  Raft  River  through 
mid-September.  This  species  does  not  ovi- 
posit in  tandem;  however,  the  male  re- 
mains in  close  proximity  to  the  female 
and  actively  wards  off  other  intruding 
males.  Martin  (1939)  and  Walker  (1953) 
observed  females  descending  as  much  as 
one  foot  below  the  water  to  deposit  eggs. 
In  Jidy  1971  I  observed  several  females 
ovipositing  at  the  Raft  River.  On  this  oc- 
casion none  submerged  but  completed 
oviposition  just  below  the  water  line  in 
algal  mats  in  shallow  water  close  to  the 
bank.  It  must  be  noted,  however,  that 
the  water  level  was  lower  than  usual  and 
there  was  little  suitable  vegetation  in  the 
deeper  portions  of  the  river. 

Hetaerina  amcricana  (Fabricius,  1788) 
Fig.  2 

With  the  exception  of  the  far  eastern 
and  northwestern  states  and  Florida,  H. 
americana  has  a  general  distribution 
throughout  the  United  States  and  extends 
southward  through  Mexico  into  Guate- 
mala. It  is  found  throughout  Utah  in 
rivers  and  streams  below  5,800  feet  ele- 
\ation  which  have  a  moderate  flow  and 
sufficient  marginal  vegetation  for  ovipo- 
sition and  nymphal  development.  The 
earliest  emergence  date  I  have  for  the 
state  is  29  May.  The  nymphs  emerge  over 


most  of  the  summer  and  can  be  found  in 
the  adult  stage  to  the  end  of  September. 
The  female  oviposits  singularly  and  com- 
pletely submerge  to  deposit  their  eggs, 
while  the  male,  which  perches  nearby, 
faces  the  submerged  female  and  actively 
fends  off  intruding  males  (Johnson  1961 
and  Bick  and  Sulzback   1966). 

Hetaerina  vulnerata  Hagen,  1853 
Fig.  2 

H.  vulnerata  is  restricted  to  the  south- 
western United  States  and  Mexico,  enter- 
ing only  the  southwest  corner  of  Utah  in 
Washington  County,  which  is  part  of  the 
Mohave  Desert  Lower  Sonoran.  Like  H. 
americana,  the  nymphs  are  found  on 
roots,  vegetation,  and  debris  in  streams 
with  a  moderate  current.  Although  these 
two  species  were  found  at  nearby  streams, 
they  were  never  taken  at  the  same  lo- 
calities (Provonsha  and  McCaffertv  1973). 


FIG,  2 

Calopteryx  aequibile 
Hetaerina  americana 
Hetaerina  vulnerata 


]> 


Emergence  begins  around  the  first  of  June 
and  is  probably  completed  by  mid-July. 
I  am  not  aware  of  any  reproductive  stud- 
ies conducted  on  this  species,  and  I  have 
never  observed  any  in  copulation.  How- 
ever, it  is  assumed  that  .the  method  of 
oviposition  is  similar  to  that  of  H.  amer- 
icana. 


Dec.  1975 


PROVONSHA:    ZYGOPTERA   OF    UTAH 


381 


Archelestes  grandis  (Rambur,  1842) 
Fig.  3 

Although  this  species  ranges  over  much 
of  the  United  States,  in  Utah  it  is  restric- 
ted to  the  southern  deserts.  The  n5nnphs 
are  active  swimmers  and  can  be  found 
in  ponds  and  slow  desert  streams.  All 
Utah  collecting  sites  were  below  4,600 
feet.  A.  grandis  is  the  largest  of  all  Nearc- 
tic  damselflies,  the  females  having  a  wing 
span  of  approximately  40  mm.  Like  most 
other  lestids,  it  is  a  late  emerger.  The 
earliest  Utah  record  I  have  is  9  July.  Bick 
and  Bick  (1970)  reported  that  in  Okla- 
homa emergence  is  much  earlier  and  that 
oviposition  is  common  by  mid-June.  In 
that  area  the  eggs  reportedly  hatch  ap- 
proximately 16  days  after  oviposition,  and 
the  nymphs  overwinter  in  a  fairlj'  late 
stage  of  development.  However,  there  is 
some  evidence  that  in  the  western  des- 
erts, where  emergence  tends  to  be  later 
and  where  there  is  often  a  winter  drought, 
eclosion  may  be  delayed  until  the  fol- 
lowing spring.  Oviposition  takes  place  in 
tandem,  and  the  eggs  are  deposited  as 
high  as  13  m  above  water  in  woody  plants 
overhanging  the  water.  This  unique  re- 
productive behavior  has  enabled  this  spe- 
cies to  colonize  certain  habitats,  such  as 
desert  streams,  where  marginal  vegeta- 
tion may  be  plentiful  but  vegetation 
within  the  water  may  be  wanting. 


T^m 


Archelestes  grandis 
Lestes  congener 


■m- 


Lestes  congener  Hagen,  1861 
Fig.  3 

This  species  is  found  over  much  of  the 
United  States  and  is  by  far  the  most  com- 
mon and  wide-ranging  Lestes  in  Utah. 
All  collecting  sites  were  at  permanent  and 
semipermanent  ponds  or  "pondlike"  ex- 
pansions of  slow  streams  at  altitudes  rang- 
ing from  4,200  to  7,000  feet.  The  nymphs 
are  free  swimmers  and  seem  to  prefer 
shallow  ponds  with  some  areas  free  of 
vegetation.  Emergence  begins  in  early 
July  and  continues  through  late  August. 
As  in  most  other  Lestes  species,  the  eggs 
are  usually  deposited  some  distance  above 
water.  However,  on  one  occasion  during 
flood  conditions  I  observed  two  females 
submerge  their  abdomens  almost  to  the 
base  to  deposit  eggs  below  the  water  line. 

Lestes  disjunctus  disjunctus  Selys,  1862 
Fig.  4 

Although  primarily  found  in  Canada, 
Alaska,  and  the  northern  regions  of  the 
United  States,  this  species  does  follow  the 
mountains  south  through  Utah  and  Colo- 
rado into  Arizona.  In  Utah  it  was  found 
mainly  at  permanent  and  semipermanent, 
richly  vegetated  ponds  in  mountainous 
areas  between  5,000  and  7,000  feet.  Emer- 
gence begins  around  mid-July  and  con- 
tinues through  mid- August.  Although  ovi- 


FiG.  ^ 


Lestes  disjunctus 


^ 


382 


GREAT   BASIN   NATURALIST 


Vol.  35,  No.  4 


position  usually  takes  place  well  above 
water,  on  two  separate  occasions  in  1970 
I  observed  paired  adults  completely  sub- 
merge to  ovipost.  These  observations 
were  made  during  flood  conditions  when 
more  than  the  usual  amount  of  vertical 
stems  were  below  water.  These  observa- 
tions coincide  for  the  most  part  with  ob- 
senations  by  Bick  and  Bick  (1961)  for 
L.  d.  nustralis  Walker. 

Lestes  dry  as  Kirby,  1890 
Fig.  5 

This  species  is  Holarctic,  occurring  not 
onh'  in  North  America  but  Europe  and 
Asia  as  well.  It  is  found  most  commonly 
at  permanent  and  semipermanent  ponds 
and  less  frequently  in  marshy  areas.  It 
was  the  only  Lestes  species  taken  above 
8,000  feet  in  Utah.  Although  the  locality 
records  are  scattered,  L.  dryas  tended  to 
be  the  dominant  species  at  these  localities. 
Emergence  begins  about  the  last  week  in 
June  and  continues  through  mid-August. 

Lestes  unquiculatus  Hagen,  1861 
Fig.  6 

The  range  of  L.  unquiculatus  is  trans- 
continental in  the  northern  United  States 
and  southern  Canada.  It  was  found  at  only 
a  few  localities  in  Utah  and  never  in  any 


7W 


FIG,   6 
Lestes  unquiculatus 


l> 


large  numbers.  Specimens  were  collected 
primarily  at  temporary  and  semiperm- 
anent marshy  areas  between  5,000  and 
6,350  feet.  Emergence  data  for  the  state 
is  insufficient.  However,  Walker  (1953) 
reported  that  in  Canada  they  fly  mainly 
in  .Inly  and  August. 

Argia  alberta  Kennedy,  1918 

Fig.  7 

A.  alberta  is  restricted  to  the  western 
United  States.  Although  it  occurs  in  most 
regions  of  Utah  at  altitudes  between  4,000 
and  6,500  feet,  it  was  never  taken  in  any 
large  numbers.  The  nymph  has  not  been 
described,  and  none  were  taken  during 
this  study.  With  the  exception  of  the  San 
Juan  River  in  San  Juan  Co.,  all  adults 
were  taken  in  association  with  small, 
slow  flowing  streams  or  marshy  springs. 
The  earliest  collecting  record  I  have  for 
the  state  is  4  June  and  the  flight  period 
extends  through  the  end  of  August.  As 
with  most  Argia  species,  oviposition  usu- 
ally takes  place  in  tandem. 

Argia  emma  Kennedy,  1915 
Fig.  8 

This  species  is  found  in  the  western 
United  States  and  British  Columbia.  In 
Utah  it  is  found  in  the  mountainous  areas 


Dec.  1975 


PROVONSHA:    ZYGOPTERA   OF   UTAH 


383 


I^ 


FIG,  7 


•  Argia  alberta 

*  Argia  sedula 


'm- 


FIG,   8 


Argia  emma 


-^m 


^ 


in  the  northern  two-thirds  of  the  state  in 
rivers  and  streams  with  gentle  to  moderate 
currents  at  altitudes  between  4,200  and 
5,800  feet.  The  nymphs  may  be  found  on 
stones  but  were  most  frequently  taken  on 
roots  and  debris  near  the  bank.  Emergence 


begins  around  the  first  of  June  and  con- 
tinues through  mid-July  and  oviposition 
usually  takes  place  in  tandem. 

Argia  fumipennis  violacea  (Hagen,  1861) 

This  species  is  known  from  Guatemala 
north  through  the  southwestern  United 
States,  most  of  the  central  and  eastern 
United  States,  and  eastern  Canada.  How- 
ever, I  have  seen  only  one  specimen  from 
Utah  (Univ.  of  Mich.  Coll.).  The  label 
read:  "Utah  E.  M.  Legard  #55."  Although 
specific  local  information  was  lacking,  it 
is  speculated,  based  on  overall  distribu- 
tion, that  this  specimen  was  collected  in 
the  Lower  Sonoran  region  of  Washington 
Co.  It  is  reported  that  A.  f.  violacea  pre- 
fers small  lakes  and  shallow  streams  with 
moderate  currents  and  exposed  rocks 
(Walker  1953). 

Argia  lugens  (Selys,  1854) 
Fig.  9 
This  species  is  found  only  in  Mexico 
and  the  southwestern  United  States.  The 
only  Utah  records  I  have  are  from  Zion 
National  Park  and  Leeds  Canyon  (both  in 
Washington  Co.)  where  the  nymphs  in- 
habit permanent  desert  streams.  I  have 
no  emergence  data  for  the  state.  All  spec- 
imens examined  were  collected  during 
the  month  of  July. 


.■  *' 

N   ,IV;. 

FIG,  9 

,    . 

7    \,  ;^- 

•  Argia  lugens 

<3 

J^-;-- 

*  Argia  moesta 

^v\y^-v 

L,.V-A^^r.;,.;-;-.^?^ 

.A 

f 

--J:','-fl^4^__:__S^----- 

'•  •"' 

;;----r- r'-y 

384 


GREAT   BASIN    NATURALIST 


Vol.  35,  No.  4 


Argia  moesta  (Hagen,  1861) 
Fig.  9 

Although  common  throughout  North 
America  wdth  the  exception  of  the  north- 
western states  and  western  Canada,  this 
species  has  been  taken  in  Utah  only  from 
the  Colorado  River,  where  the  nymphs  can 
be  found  on  rubble  and  in  debris  near  the 
bank.  Emergence  begins  in  late  May  and 
continues  through  most  of  June.  Walker 
(1953)  reports  that  females  oviposit  either 
in  tandem  or  unattended  by  the  male 
and  that  they  may  completely  submerge 
to  deposit  their  eggs.  Unlike  most  western 
species  of  Argia,  which  prefer  to  light  on 
bare  ground,  all  individuals  of  A.  moesta 
observed  in  Utah  rested  on  willows  along 
the  river's  edge. 

Argia  nahauna  Calvert,  1901 
Fig.  8 

This  species  is  known  only  from  the 
southwestern  United  States,  entering  Utah 
only  in  Washington  Co.  It  is  found  at 
small  streams  with  moderate  currents 
and  occasionally  in  permanent  desert 
springs.  At  most  localities  it  was  found 
in  association  with  A.  sedula.  The  earliest 
Utah  record  I  have  is  5  June  and  the  latest 
is  9  September. 

Argia  sedula  (Hagen,  1861) 
Fig.  7 
The  reported  range  of  A.  sedula  in- 
cludes most  of  the  southern  portion  of  the 
United  States,  northern  Mexico,  the  mid- 
western  states,  and  southern  Ontario.  In 
Utah  it  was  found  only  in  Washington 
Co.  at  altitudes  between  2,800  and  4,400 
feet.  The  nymphs  prefer  small  streams 
with  gentle  current  and  a  rich  growth  of 
vegetation.  Emergence  begins  in  mid -May 
and  continues  through  mid- June. 

Argia  vivida  Hagen,  1865 
Fig.  10 

A.  vivida  is  restricted  to  the  western 
United  States  and  southwestern  Canada. 
It  is  fairly  common  in  Utah  and  can  be 
found  at  most  rivers  and  streams  with 
moderate  currents.  During  this  study  they 
were  taken  at  altitudes  between  2,800 
and  6,500  feet.  Although  they  can  be 
found  in  debris  along  the  stream  banks, 
the  nymphs  prefer  stones  and  rubble  well 


FIG.  10 


•  Argia  vivida 


??-iu:. 


within  the  main  current  of  the  stream. 
Emergence  begins  around  the  first  of  June 
and  continues  through  mid-July.  Ovipo- 
sition  takes  place  in  tandem,  and  eggs  are 
laid  just  below  the  water  surface  in  a 
wide  variety  of  aquatic  plants.  Like  most 
other  Argia  species,  the  males  will  fre- 
quently balance  in  a  vertical  position 
during  oviposition  when  the  immediate 
environment  does  not  provide  a  good  ob- 
ject for  them  to  grasp. 

Amphiagrion  abbreviatum  (Selys,  1876) 
Fig.  11 

This  species  is  known  from  the  western 
United  States  and  southwestern  Canada. 
I  found  this  species  at  a  great  many  lo- 
calities throughout  the  state,  but  never  in 
any  large  numbers.  The  habitats  were 
varied,  ranging  from  moderately  flowing 
streams  with  gravel  bottoms,  to  heavily 
vegetated  ponds  and  springs,  to  large 
lakes.  However,  as  Whitehouse  (1941) 
foiuid,  they  tend  to  prefer  shallow,  sunlit 
marshy  areas  with  little  or  no  current. 
They  were  taken  at  altitudes  between 
4,200  and  7,500  feet.  Emergence  begins 
in  early  May  and  the  flight  period  ex- 
tends through  the  end  of  August.  Ovipo- 
sition usually  takes  place"  in  tandem,  but 
I  have  observed  unattended  females  prob- 
ing algal  mats. 


Dec.  1975 


PROVONSHA:    ZYGOPTERA    OF    UTAH 

FIG.   11 
•  Amphiagrion  abbreviatum 


385 


fU''\ 


m^.^-. 


•  .--       • 


Telebasis  salva  (Hagen,  1861) 
Fig.  12 

T.  salva  ranges  from  the  southwestern 
United  States  to  Venezuela.  I  have  seen 
only  three  specimens  taken  in  Utah.  They 
were  all  collected  at  a  small  spring-fed 
pond  near  St.  George,  Washington  Co.: 
two  in  1941  (Univ.  of  Utah  Coll.)  and 
one  in  1959  (Dixie  College  Coll.).  Re- 
cently the  area  has  been  converted  into 
a  golf  course,  greatly  changing  the  origi- 
nal habitat.  It  is  possible  that  this  species 
no  longer  occurs  in  the  state. 

Coenagrion  resolutum  (Hagen,  1876) 
Fig.  12 

This  species  is  known  from  throughout 
most  of  Canada  and  Alaska,  the  northeast- 
ern states,  and  higher  elevations  in  the 
West.  In  Utah  it  was  found  at  small  ponds 
and  slow-flowing  high  meadow  streams 
above  7,000  feet,  where  they  emerge  dur- 
ing the  month  of  July.  Evidently  all  ovi- 
position  is  in  tandem. 

EnaUagma  anna  Williamson,  1900 
Fig.  13 

E.  anna  is  restricted  to  the  western 
United  States.  This  species  is  fairly  com- 
mon  throughout   the   mountainous    areas 


Coenagrion  resolutum 
Telabasis  salva 


^■^^^ 


FIG,  13 


of  northern  Utah  at  altitudes  between 
4,200  and  7,000  feet.  Nymphal  develop- 
ment takes  place  in  rivers  and  streams 
with  slow  to  moderate  currents.  The 
flight  period  ranges  from  mid-May 
through  the  first  of  September,  and,  like 
most  other  species  of  EnaUagma,  ovipo- 
sition  takes  place  in  tandem. 


386 


GREAT   BASIN   NATURALIST 


Vol.  35,  No.  4 


Enallagma  boreale  Selys,  1875 
Fig.  14 

The  range  of  this  species  includes  most 
of  Canada  and  the  northern  United  States 
with  isolated  populations  at  higher  al- 
titudes in  the  southwest.  Next  to  Ischnura 
perparva  Selys,  it  is  the  most  common 
species  in  Utah,  being  absent  only  from 
the  desert  areas  in  the  southeast.  It  is 
most  frequently  found  at  lakes  and  ponds 
and  rarely  at  slow-flowing  streams.  Col- 
lection sites  ranged  from  4,000  to  9,000 
feet.  It  is  an  early  emerger,  and  the  first 
record  I  have  for  the  state  is  8  May.  None 
were  taken  after  mid-August.  Oviposition 
usually  takes  place  in  tandem. 

Enallagma  carunculatum  Morse,  1895 
Fig.  15 

With  the  exception  of  a  break  along  the 
Rocky  Mountains,  this  species  is  found 
transcontinentally  in  southern  Canada  and 
the  northern  United  States,  extending 
southward  in  the  West  into  northern 
Mexico.  This  is  one  of  the  more  common 
and  widely  adapted  species  in  Utah,  often 
occurring  in  great  numbers.  Although  it 
was  most  frequently  found  at  small  ponds, 
it  was  also  taken  at  streams,  rivers,  and 
some  of  the  larger  lakes  and  reservoirs 
at  altitudes  between  3,000  and  8,000  feet. 


E.  carunculatum  is  one  of  the  few  species 
which  can  develop  in  brackish  water, 
such  as  that  occurring  in  drainage  ditches 
west  of  Salt  Lake  Cit}^  near  the  Great 
Salt  Lake,  where  it  was  found  in  close  as- 
sociation with  E.  clausum  Morse.  This  as- 
sociation has  also  been  reported  for  other 
brackish  waters  such  as  Pyramid  Lake, 
Nevada  (Kennedy  1917).  Emergence  be- 
gins in  late  May,  and  oviposition  takes 
place  in  tandem. 


vr^ 


FIG,  15 


•  Enallagma  carunculatum 


imt  ^ 


Enallagma  civile  (Hagen,  1861) 
Fig.  16 

With  the  exception  of  the  far  northwest, 
E.  civile  has  a  wide  range  over  most  of 
the  United  States  and  extends  as  far  south 
as  the  West  Indies  and  Colombia.  Al- 
though this  is  one  of  the  most  common 
species  in  North  America,  it  was  taken 
at  only  a  few  scattered  localities  in  Utah. 
Locally  abundant  populations  were  taken 
in  the  Sevier  River  drainage  system  in 
Millard  Co.,  but  only  small  numbers 
were  taken  at  most  other  collecting  sites. 
Nymphs  were  taken  in  lakes,  ponds,  and 
streams  and  rivers  with  slow  currents  at 
altitudes  from  2,800  to  6,000  feet.  Like  its 
close  relative  E.  carunculatum,  emergence 
begins  in  late  May  and  continues  through 
most   of  the   summer. 


Dec.  1975 


PROVONSHA:  ZYGOPTERA  OF  UTAH 


387 


FIG,  16 


•  Enallagma  civile 

*  Enallagma  ebrium 


W 


Enallagma  clausum  Morse,  1895 
Fig.  17 
This  species  is  found  in  the  western 
United  States  and  southwestern  Canada 
with  isolated  populations  in  Ontario  and 
Quebec.  In  Utah  it  is  restricted  to  the 
western  half  of  the  state  at  altitudes  be- 
tween 4,200  and  6,300  feet.  The  nymphs 


W] 


•  Enallagma  clausum 


M& 


may  be  found  in  slow  streams  and  lakes 
with  some  wave  action.  Enallagma  clau- 
sum is  tolerant  to  saline  conditions  and 
was  frequently  found  in  association  with 
E.  carunculatum  at  brackish  waters  in  the 
western  desert.  UnHke  most  species  of 
Enallagma,  adults  prefer  to  rest  on  bare 
ground  rather  than  on  vegetation;  when 
they  are  at  rest  the  wings  are  frequently 
held  at  a  slight  angle  away  from  the  body. 
Emergence  begins  in  late  May  and  con- 
tinues through  most  of  the  summer. 

Enallagma  cyathigerum 

(Charpentier,  1840) 

Fig.  18 

This  is  a  Holarctic  species,  being  found 
in  Europe,  Asia,  Canada,  and  most  of  the 
northern  United  States.  It  is  one  of  the 
more  common  species  in  Utah  and  is  ab- 
sent only  from  the  western  deserts.  Al- 
though most  common  at  small  ponds,  it 
was  frequently  taken  at  streams  wdth 
slow  to  moderate  currents.  With  the  ex- 
ception of  Coenagrion  resolutum,  it  was 
the  only  species  taken  above  9,000  feet. 
Emergence  begins  in  mid-June,  and  adults 
can  be  found  through  early  September. 

Enallagma  ebrium  (Hagen,  1861) 
Fig.  16 

The  range  of  this  species  is  reported  as 
Boreal  North  America.  It  has  been  taken 
at  only  two  localities  in  Utah.  Fourteen 
specimens  were  collected  at  the  Weber 
River,  West  Weber,  Weber  Co.,  by  C.  J. 
D.  Brown  (1934),  and  two  adults  were 
collected  during  this  present  study  at 
marshy  areas  along  the  Bear  River  in 
Cache  Co.  Both  of  the  above-mentioned 
collections  were  made  in  July.  Walker 
(1953)  records  the  flight  period  in  Canada 
to  be  from  30  May  through  24  August. 

Enallagma  praevarum  (Hagen,  1861) 
Fig.  13 

This  species  ranges  from  the  southern 
United  States  to  southern  Mexico.  In 
Utah  it  is  restricted  to  desert  streams  and 
springs  at  2,800  to  5,800  feet  elevation  in 
the  unique  southwest  corner  of  the  state. 
Although  very  closely  related  to  E.  anna, 
at  no  place  did  their  ranges  overlap. 
Emergence  begins  in  mid-May  and  con- 
tinues through  the  month  of  June. 


388 


GREAT   BASIN   NATURALIST 

FIG,  18 


•  Enallagma  cyathigerum 


Ischnura  barberi  Currie,  1903 
Fig.  19 

Ischnura  barberi  is  known  only  from 
the  western  United  States.  All  collecting 
sites  in  Utah  were  at  ponds  in  the  west- 
ern portion  of  the  state  at  altitudes  rang- 
ing from  2,800  to  4,730  feet.  This  species 
appears  to  be  rare  in  numbers,  since  no 
more  than  two  specimens  were  taken  at 
any  one  locality.  Oviposition  was  never 
observed.  All  collections  were  made  dur- 
ing the  months  of  June  and  July. 

Ischnura  cervula  Selys,  1876 
Fig.  20 

Southwestern  Canada,  western  United 
States,  and  northern  Mexico  are  included 
in  the  range  of  this  species.  It  is  fairly 
common  and  was  frequently  found  at 
ponds  and  slow  streams  with  a  preference 
for  shallow  marshy  areas.  Samples  were 
from  altitudes  ranging  from  4,200  to  7,000 
feet.  Oviposition  does  not  usually  take 
place  in  tandem.  Emergence  begins  in 
early  May  and  continues  through  most 
of  the  summer. 

Ischnura  damula  Calvert,  1901 
Fig.  21 
This    species    is    known    in    southern 
Canada  from  Manitoba  west  and  the  west- 


Vol.  35,  No.  4 


Ischnura  barberi 
Ischnura  demcrsa 


)> 


ern  United  States.  Except  for  the  warm 
springs  in  Tooele  Co.,  its  distribution  in 
Utah  is  restricted  to  the  southern  portion 
of  the  state.  All  specimens  were  collected 
from  small  ponds  and  springs  with  dense 
stands  of  cattails  along  the  margins.  All 
sites  were  between  an  altitude  of  4,225  and 
5,825  feet.  Females  were  observed  ovipos- 


FIG.  20 


Dec.  1975 


PROVONSHA:    ZYGOPTERA   OF    UTAH 

FIG.  21 

•    ISCHNURA   DAMULA 


389 


FIG.  22 


SCHNURA  DENTICOLLIS 


l^-f 


iting  singularly  and  in  tandem.  The  ear- 
liest record  I  have  for  the  state  is  23 
May.  It  is  probable  that  emergence  be- 
gins in  early  May  and  continues  through 
early  July. 

Ischnura  demorsa  (Hagen,  1861) 
Fig.  19 

Ischnura  demorsa  is  reported  from  the 
western  United  States  and  Mexico.  This 
close  relative  of  /.  perparva  is  rare  in 
Utah,  however.  Only  two  specimens  were 
taten  during  this  study,  both  from  Mc- 
Cracken  Spring  in  San  Juan  Co.  at  an  al- 
titude of  4,900  feet.  I  have  no  emergence 
data  for  this  species. 

Ischnura  denticollis  (Burmeister,  1839) 
Fig.  22 
This  species  is  found  only  in  the  south- 
western United  States  and  Mexico.  In 
Utah  it  is  restricted  to  the  cold  deserts 
in  the  western  half  of  the  state.  It  is 
equally  suited  to  both  ponds  and  streams 
and  is  tolerant  to  a  wide  range  of  altitudes, 
being  found  from  2,700  to  6,300  feet. 
Oviposition  usually  takes  place  in  tan- 
dem. This  species  tends  to  emerge  some- 
what later  than  most  other  Utah  isch- 
nurans.  No  specimens  were  taken  before 
the  first  of  June,  and  15  August  was  the 
last  date  they  were  encountered. 


Ischnura  perparva  Selys,  1876 
Fig.  23 

This  species  is  known  from  the  western 
United  States  and  southwestern  British 
Columbia.  This  is  by  far  the  most  common 
damselfly  in  Utah.  It  was  found  at  87  of 
the    152    localities    sampled    during    this 


FIG.  23 


SCHNURA  PERPARVA 


390 


GREAT  BASIN   NATURALIST 


Vol.  35,  No.  4 


Study  (almost  twice  as  many  as  any  other 
species).  Its  great  abmidance  can  be  at- 
tributed, in  part,  to  its  ability  to  survive 
in  many  diverse  habitats.  It  was  found  in 
most  aquatic  situations  between  4,200  and 
7,500  feet  where  there  was  sufficient  vege- 
tation and  a  high  enough  minimum  tem- 
perature to  support  zygopteran  forms.  Al- 
though oviposition  may  take  place  in  tan- 
dem, it  is  usually  accomplished  unattend- 
ed by  the  male.  Emergence  begins  in 
early  May  and  continues  throughout 
most  of  the  summer. 


Acknowledgments 

The  author  wishes  to  thank  Dr.  George 
F.  Edmunds,  Jr.,  University  of  Utah;  Dr. 
Wilford  J.  Hanson,  Utah  State  University; 
Dr.  Vasco  M.  Tanner,  Brigham  Young 
University;  and  Dr.  Andrew  H.  Barnum, 
Dixie  College,  for  the  loan  of  specimens. 
Gratitude  is  also  extended  to  Mrs.  Leonora 
K.  Gloyd,  Museum  of  Zoology,  University 
of  Michigan,  and  Dr.  Minter  J.  Westfal'l, 
Jr.,  University  of  Florida,  for  verification 
of  identifications  and  advice  given  during 
this  study.  Special  appreciation  is  ex- 
pressed to  Dr.  W.  P.  McCafferty,  Purdue 
University,  for  his  encouragement,  advice, 
and  assistance  both  during  the  study  and 
in  the  preparation  of  the  manuscript. 


References 

BiCK,  G.  H.,  AND  J.  C.  BicK.  1961.  An  adult 
population  of  Lestes  disjunctus  australis 
Walker  (Odonata:  Lestidae).  Southwestern 
Nat.    6(3-4):  111-137. 

.     1970.     Oviposition    in    Archilestes    gran- 

dis  (Rambur)  (Odonata:  Lestidae).  Ent. 
News  81:157-163. 

BiCK,  G.  H.,  AND  L.  E.  HoRNUFF.  1972.  Odo- 
nata collected  in  Wyoming,  South  Dakota, 
and  Nebraska.  Proc.  Entomol.  Soc.  Wash. 
74(l):l-8. 


BiCK,  G.  H.,  AND  D.  SuLZBACK.  1966.  Repro- 
ductive behavior  of  the  damselfly,  Hetaerina 
americana  (Fabricius)  (Odonata:  Calopterygi- 
dae).  Anim.  Behav.   14:156-158. 

Brown,  C.  J.  D.  1934.  A  preliminary  list  of 
Utah  Odonata.  Univ.  Mich.  Occas.  Pap.  Mus. 
Zool.  no.  291,   17  pp. 

Johnson,  C.  1961.  Breeding  behavior  in  Hetae- 
rina americana  (Fabricius)  and  H.  titia 
i(Drury)  (Odonata:  Agriidae).  Canadian  Ent. 
93:260-266. 

.     1972.     The    damselflies    (Zygoptera)    of 

Texas.  Bull.  Florida  State  Mus.,  Biol.  Sci. 
16(2):55-128. 

Kennedy,  C.  H.  1915.  Notes  on  the  life  history 
and  ecology  of  the  dragonflies  (Odonata)  of 
Washington  and  Oregon.  Proc.  U.S.  Natl. 
Mus.  99(2107) :259-345. 

.     1917.     Notes    on    the    life    history    and 

ecology  of  the  dragonflies  (Odonata)  of  cen- 
tral California  and  Nevada.  Proc.  U.S.  Natl. 
Mus.  52(2192) :483-635. 

KoRMONDY,  E.  J.  1957.  Records  of  western  Odo- 
nata with  notes  on  Amphiagrion  abbreviatum 
(Selys).  J.  Kansas  Ent.  Soc.  30(3) :  108-110. 

Larson,  W.  P.  1952.  The  dragonflies  (Anisop- 
tera)  of  Utah.  Unpublished  master's  thesis, 
Dept.  of  Zoology,  Univ.  Utah,  Salt  Lake  City, 
95  pp.,  30  plates. 

Martin,  R.  D.  C.  1939.  Life  history  of  Agrion 
aequabile  and  Agrion  maculatum.  Ann.  Ent. 
Soc.  Amer.  32:601-618. 

MussER,  R.  J.  1962.  Dragonfly  nymphs  of  Utah 
(Odonata:  Anisoptera)  Univ.  Utah  Biol.  Sen, 
vol.  12,  no.  6,  Oct.  1962. 

Paulson,  D.  R.  1970.  A  list  of  the  Odonata 
of  Washington  with  additions  to  and  dele- 
tions from  the  state  list.  Pan-Pacific  Ent. 
46(3):  194-198. 

Provonsha,  a.  v.,  and  W.  P.  McCafferty. 
1973.  Previously  unknown  nymphs  of 
western  Odonata  (Zygoptera:  Calopterygidae, 
Coenagrionidae) .  Proc.  Entomol.  Soc.  Wash- 
ington 75(4):  449-454. 

Smith,  R.  F.,  and  A.  E.  Pritchard.  1956. 
Odonata.  pp.  106-153.  In:  Aquatic  insects  of 
California.  R.  L.  Usinger,  ed.  Univ.  Calif. 
Press,  Berkeley,  and  Los  Angeles. 

Walker,  E.  M.  1953.  The  Odonata  of  Canada 
and  Alaska.  Vol.  1,  Univ.  of  Toronto  Press, 
Toronto  XI    +    292  pp. 

Whitehouse,  F.  C.  1941.  British  Columbia 
dragonflies  (Odonata)  with  notes  on  distribu- 
tion and  habits.  Amer.  Midland  Nat.  26: 
488-557. 


NEW  SYNONYMY  AND  NEW  SPECIES  OF  AMERICAN 
BARK  BEETLES  (COLEOPTERA:  SCOLYTIDAE),  PART  IP 

Stephen  L.  Wood- 

Abstr..\ct. —  New  synoiiyiiij-  is  proposed  as  follows:  Pityophthoius  Eichhoff  (=  GmUhophorus 
Schedl.  Gnathopfithorus  Wood).  Araptus  confinis  (Blandford)  (=  Neopilyophthorus  glabricollis 
Schedl),  A.  eruditus  (Srhedl)  (=  Neodryocoetes  buscki  Blackiiian),  A.  hymenaeae  (Eggers)  (=  Neo- 
dryocoetes  humilis  Blackmail),  A.  schedli  (Blackmail)  (=  Neodryocoetes  lenis  Blackmail),  A.  lenel- 
lus  (Schedl)  (=  Ctenyophthorus  mexicanus  Schedl,  Neodryocoetes  granulatus  Schedl,  Araptus  cuspi- 
dus  Wood),  Coccotrypes  carpophagus  (Hornung)  (=  Coccotrypes  liberiensis  Hopkins,  Coccotrypes 
pimctatulus  Eggers),  C.  daclyliperda  (Fabricius)  (  =-  Coccotrypes  bassiavorus  Hopkins).  C.  robuslus 
Eichhoff  (=  Coccotrypes  cylindricus  Schedl).  Cryptocarenus  heveae  (Hagedorn)  (=  Cryptocarenus 
caraibicus  Eggers).  Hypothenemus  setosus  (Eichhoff)  (^  Stephanoderes  congonus  Hagedorn).  Micro- 
corthylus  minutus  Schedl  (=  Microcorthylus  minutissimus  Schedl),  Pseudopityophthorus  limbatus 
Eggers  (=  Pseudopityophthorus  rnicans  'Wood),  Xyleborus  obliquus  (LeConte)  (=  Xyleborus  gil- 
vipes  Blandford,  A',  brasiliensis  Eggers,  illepidus  Schedl).  Hypothenemus  javanus  Eggers  is  a  valid 
species.  The  genus  Dacnophthorus.  type-species  Gnathophthorus  clematus  Wood,  is  described  as  new 
to  science.  The  following  species  are  described  as  new  to  science:  Araptus  consobrinus,  A.  micaceus, 
Pityophthorus  explicitus,  and  P.  inceptis  (Mexico).  P.  costatus  and  P.  mendosus  (Costa  Rica).  P.  de- 
gener  and  P.  timidulus  (Panama).  P.  amiculus  (Mexico.  Costa  Rica),  and  P.  dissolutus  (Costa  Rica, 
Panama).  Xyleborus  californicus  (California).  X.  incultus.  X.  molestulus  (Panama),  and  tristiculus 
(Brazil). 


On  the  following  pages  seAeral  newly 
discovered  cases  of  synonym}',  one  new 
genus,  and  14  sjDecies  new^  to  science  are 
presented  for  American  Scolytidae.  The 
specific  synonymy  is  presented  alphabeti- 
cally for  convenience  of  reference.  The 
species  new  to  science  represent  the  genera 
Araptus  (2),  Pityophthorus  (8),  and 
Xyleborus  (4).  They  are  from  the  fol- 
lowing countries:  United  States  (1), 
Mexico  (4),  Costa  Rica  (2),  Panama  (4), 
Brazil  (1),  Mexico  and  Costa  Rica  (1), 
Costa  Rica  and  Panama  ( 1 ) . 


New  Synonymy 

Pityophthorus   Eichhoff 

Pityophthorus  Eichhoff.  1864,  Berliner  Ent.  Zeit. 
8:39  (Type-species:  Bostrichus  lichtensteini 
Ratzeburg,  subsequent  designation  by  Hop- 
kins.   1914,   Proc.   U.S.   Nat.    Mus.   48:127) 

Gnathophorus  Schedl,  1935  (nee  Kirby,  1837). 
Rev.  de  Ent.  5:342  ( Type-species :  Gnatho- 
phorus sparsipilosus  Schedl.  monobasic).  New 
synonymy 

Gnathophthorus  Wood,  1962,  Great  Basin  Nat. 
22:76  (Replacement  name  for  Gnatho- 
phorus) .   New  synonymy 

The  holotype  of  Gnathophorus  sparsi- 
pilosus Schedl  was  examined  and  com- 
pared directly  to  a  series  of  this  species 
in  my  collection.  It  fits  well  within  the 
limits  of  the  genus  Pityophthorus  anatom- 


^Part   of   this   research   \-\ 
-Department    of    Zoology 


.   sponsored    bj-   the   National    Scienc 
Brigham    Yoiing    University,    Prov 


ically  as  well  as  biologically.  For  this  rea- 
son the  names  Gnathophorus  and  its  re- 
placement, Gnathophthorus,  must  be 
placed  in  synonymy  as  indicated  above. 
The  five  species  subsequently  assigned  to 
this  genus  must  be  transferred  to  another 
genus  that  is  described  below. 

Araptus  confinis   (Blandford) 

Pityophthorus  confinis  Blandford,  1904,  Biol. 
Centr.  Amer.  Coleopt.  4(6):241  (Lectotype, 
male;  Jalapa.  Veracruz.  Me.xico;  British  Mus. 
Nat.   Hist.,   present   designation) 

Neopilyophthorus  glabricollis  Schedl,  1938,  Archiv 
Naturgesch.  7(2):  181  (Holotype.  male;  Teo- 
pisca,  Chiapas,  Mexico;  Schedl  Coll.).  New 
synonymy 

The  first  syntype  in  the  type  series  of 
Pityophthorus  confinis  Blandford  is  here 
designated  as  the  lectotype  of  that  species. 
This  lectotype  was  compared  directly  to 
my  males  from  Guatemala  City,  Guate- 
mala, and  was  found  to  be  identical  in  all 
respects.  My  series  was  later  compared 
directly  to  the  male  holotype  of  Neopity- 
ophthorus  glabricollis  Schedl.  Except  for 
the  loss  of  declivital  and  frontal  setae,  an 
apparent  result  of  abrasion  on  the  Schedl 
type,  these  specimens  are  also  identical. 
As  indicated  above,  the  junior  name  must 
be  placed  in  synonymy.  The  lectotype  of 
confinis  is  labeled  "Type"  and  has  been 
considered  as  the  type  for  many  years. 

e    t'oundation. 

o,    Utah    84602.    Scolytidae    contribution    No.    Gl 


391 


392 


GREAT   BASIN   NATURALIST 


Vol.  35,  No.  4 


Araptus  eruditus  (Schedl) 

Neopityophthorus  eruditus  Schedl.  1938,  Archiv 
Naturgesch..  n.  f.,  7(2):  182  (Holotype,  fe- 
male;   Mexico;    Schedl  Coll.) 

Neodrrocoetes  buscki  Blackman,  1942,  Proc.  U.S. 
Nat.  Mus.  92:192  (Holotype.  female;  Cabima, 
Panama;    U.S.    Nat.    Mus.).    New   synonymy 

The  female  holotype  of  Neopityoph- 
thorus eruditus  Schedl  and  the  female 
holotype  of  Neodryocoetes  buscki  Black- 
man  were  compared  direct!}-  to  my  series. 
All  represent  the  same  species.  The  last 
visible  abdominal  sternum  is  armed  by 
a  subtuberculate  callus  which  aids  in  the 
identification    of   this    species. 

Araptus  hymenaeae  (Eggers) 

Neodryocoetes  hymenaeae  Eggers.  1933.  Mem. 
Trav.  Lab.  d'Ent.  Mus.  Nat!  d'Hist.  Nat.. 
Paris  1(1)  :9  (Holotype,  female;  Gourdon- 
ville,   French   Guiana;    Paris   Mus.) 

Neodryocoetes  humilis  Blackman,  1942,  Proc.  U.S. 
Nat.  Mus.  92:188  (Holotype,  female;  Bonito, 
Pernambuco.  Brazil;  U.S.  Nat.  Mus.)  New 
synonymy 

The  \ariable  frons  of  this  species  has 
led  to  the  description  of  several  synonyms. 
Following  a  study  of  the  holotype  and  six 
cotypes  of  Neodryocoetes  hymenaeae  Eg- 
gers, the  female  holotype  of  Neodryocoetes 
humilis  Blackman,  and  76  other  speci- 
mens, it  was  concluded  that  only  one  spe- 
cies was  represented  and  that  Blackman's 
name  must  be  placed  in  synonymy. 

Araptus  schedli  (Blackman) 

Neodryocoetes  schedli  Blackman,   1942,  Proc.  U.S. 

Nat.  Mus.  92:195   (Holotype,  male;  Tampico. 

Mexico;    U.S.    Nat.    Mus.) 
Neodryocoetes   lenis   Blackman,    1942,    Proc.    U.S. 

Nat.  Mus.  92:198   (Holotype.  male;  Cordoba. 

Veracruz.    Mexico;    U.S.    Nat.    Mus.).    New 

synonymy 

The  male  holotypes  of  Neodryocoetes 
schedli  Blackman  and  Neodryocoetes  lenis 
Blackman  were  compared  directly  to  one 
another  and  to  several  specimens  inter- 
cepted at  New  York  in  seeds  of  Mexican 
origin.  I  am  unable  to  detect  any  differ- 
ences among  these  specimens  that  might 
justify  the  continued  recognition  of  two 
names. 


Araptus  tenellus  (Schedl) 

Neodryocoetes  tenellus  Schedl,  1951,  Dusenia  2: 
109  (Holotype.  male;  Chiapas,  Mexico; 
Schedl    Coll.) 


Ctenyophthorus  mexicanus  Schedl.  1963.  Ent.  Arb. 
Mus.  Frey  14:162  (Holotype,  female;  Tram- 
paluz,  Escarcaga.  Campeche,  Mexico;  Schedl 
Coll.).  New  synonymy 

Neodryocoetes  granulatus  Schedl,  1964,  Reichen- 
bachia  3:311  (Replacement  name  for  Cteny- 
ophthorus mexicanus  Schedl).  New  synon- 
ymy 

Araptus  cuspidus  Wood,  1974,  Brigham  Young 
Univ.  Sci.  Bull..  Biol.  Ser.  19  (1):46  (Holo- 
type, female;  8  km  E  San  Bias,  Nayarit.  Mex- 
ico; Wood  Coll.).  New  synonymy 

The  holotype  of  Neodryocoetes  tenellus 
Schedl  and  the  allotype  of  Araptus  cus- 
pidus Wood  were  compared  directly  to 
one  another.  Except  for  minor  differences 
in  the  convexity  of  the  frons,  they  are 
identical.  The  peculiar  abdominal  stern- 
um 5  was  not  mentioned  by  Schedl.  The 
Schedl  holotype  of  Ctenyophthorus  mexi- 
canus is  a  female,  not  a  male  as  stated  in 
the  original  description.  It  was  compared 
directly  to  the  holotype  of  cuspidus  and 
was  found  to  be  identical.  The  two  junior 
names  and  the  replacement  name,  Neo- 
dryocoetes granulatus  Schedl,  must  be 
placed  in  synonymy  as  indicated  above. 

Coccotrypes  carpophagus  (Hornung) 

Bostrichus  carpophagus  Hornung,  1842,  Stettiner 
Ent.  Zeit.  3:116  (Syntypes;  intercepted  in 
Germany  from  "Betelniissen"  of  "Ostindien" 
origin) 

Coccotrypes  liberiensis  Hopkins,  1915,  U.S.  Dept. 
Agric.  Kept.  99:47  (Holotype,  female;  Mount 
Coffee,  Liberia;  U.S.  Nat.  Mus.).  New  sy- 
nonymy 

Coccotrypes  punctatulus  Eggers,  1951.  Ent.  Blatt. 
45-46:151  (Holotype,  female;  Insel  St. 
Thomas,  Virgin  Islands;  deposited  in  Eggers 
Coll.,  apparently  on  loan  to  Schedl).  New 
synonymy 

The  holotypes  of  Coccotrypes  liberiensis 
Hopkins  and  C.  punctatulus  Eggers  and 
the  syntypes  of  Bostrichus  carpophagus 
Hornung  were  all  examined  and  com- 
pared directly  to  my  series  of  this  species. 
Only  one  species  is  represented  by  the 
three  names. 


Coccotrypes  dactyliperda   (Fabricius) 

Bostrichus  dactyliperda  Fabricius,  1801,  Systema 
Elcutheratorium  2:387  (Syntypes;  in  date 
pits  intercepted  in  Europe;  Copenhagen  and 
Berlin    museums) 

Coccotrypes  bassiavorus  Hopkins.  1915,  U.S.  Dept. 
Agric.  Rept.  99:47  (Holotype,  female;  Wash- 
ington, D.C.;  M.S.  Nat.  Mus.).  New  synon- 
ymy 

Several     female    specimens    of    Cocco- 
trypes dactyliperda  (Fabricius)  that  were 


Dec.  1975 


WOOD:    AMERICAN    SCOLYTIDAE 


393 


compared  by  Eggers  to  the  Fabricius  syn- 
types  were  compared  directly  to  the  fe- 
male holotype  of  C.  hassiavorus  Hopkins. 
Since  they  are  identical  in  all  respects, 
Hopkins's  name  must  be  placed  in  synon- 
>'my. 

Coccotrypcs  robustus  Eichhoff 

Coccotrrpes   robuslus   Eiclihoff.    1878.    Mem.    Soc. 

Roy.   Sci.   Li6ge    (2)8:513    (Syntypes?;   Cuba; 

presumably-  lost  witJi  Hamburg  Mus.) 
Coccotrrpes    cylindricus    Schedl,     1949,    Tijdschr. 

Ent.      91:116      (Holotype,      female;      Crucos. 

Cuba;   Schedl  Coll.).  New  synonymy 

A  female  of  Coccotrypes  robustus  Eich- 
hoff that  was  compared  by  Hopkins  to  a 
syntype  and  the  holotype  of  C.  cylindri- 
cus Schedl  were  compared  to  my  series  of 
this  species.  Since  all  represent  the  same 
distinctive  species,  Schedl's  name  must 
be  placed  in  synonymy. 


Cryptocarenus  heveae  (Hagedom) 

Stephanoderes  heveae  Hagedorn,   1912,  Rev.  Zool. 

Afr.    1:338    (Syntypes,  female;   Eala,   Congo; 

Tervuren  Mus.) 
Cryptocarenus    caraibicus    Eggers.    1937,    Rev.    de 

lEnt.     7:82     (Holotype,    female;    Guadeloupe; 

U.S.    Nat.   Mus.).   New  synony?ny 

The  four  female  syntypes  of  Stephan- 
oderes heveae  Hagedorn  and  the  female 
holotype  of  Cryptocarenus  caraibicus  Eg- 
gers were  compared  to  my  material.  All 
of  these  specimens  represent  the  same 
species. 

Schedl  (1975,  Ent.  Blatt.  71:43)  treated 
C.  lepidus  Wood  as  a  synonym  of  C.  car- 
aibicus. These  species  are  easily  confused 
but  quite  distinct. 


Hypothenemus  setosus  (Eichhoff) 

Hypoborus  (?)  setosus  Eichhoff,  1867,  Berliner 
Ent.  Zeitschr.  11:391  (Syntypes,  Guadeloupe; 
one  syntype  in  U.S.  Nat.  Mus.,  others  lost 
with   Hamburg   Mus.) 

Stephanoderes  congonus  Hagedorn,  1912,  Rev. 
Zool.  Afr.  1:337  (Lectotype,  female;  Eala, 
Congo;  Tervuren  Mus.,  present  designation). 
New  synonymy 

Stephanoderes  congonus  Hagedorn  was 
based  on  a  syntypic  series.  Four  of  those 
syntypes  are  in  the  Musee  Royal  de  I'Afri- 
que  Central,  Tervuren,  labeled  as  one 
"Holotypus"  and  three  "Paratypus." 
These  type  designations,  however,  have 
not  beeen  mentioned  in  print  and  are  con- 


sidered invalid.  The  "Holotypus"  is  se- 
\erely  damaged  but  recognizable.  All  four 
specimens  bear  identical  data  labels: 
"Musee  du  Congo  Beige;  Eala;  Rakusu 
(H??  1140  M,  1911);  D.  Hevea  brasilien- 
sis."  Because  the  first  specimen  is  severely 
damaged,  I  designate  the  third  specimen 
as  the  lectotype. 

The  above  lectotype  was  compared  to 
several  of  my  series  of  setosus  (Eichhoff) 
that  previously  had  been  compared  di- 
rectly to  the  only  known  syntype  of 
setosus.  Only  one  species  is  represented  by 
this  material.  It  is  a  species  distinct  from 
javanus  (Eggers),  as  noted  below.  Other 
synonyms  of  setosus  include  Stephano- 
deres obscurus  Eichhoff  (nee  Ferrari)  and 
S.  depressus  Eichhoff. 

Hypothenemus  javanus  Eggers, 

new  status 

Stephanoderes  javanus  Eggers.  1908.  Ent.  Blatt. 
4:215  (Lectotype.  female;  Java;  U.S.  Nat. 
Mus..  designated  by  Anderson  and  Ander- 
son,  1971,  Smithsonian  Contrib.  Zool.  94:16) 

This  species  was  placed  in  synonymy 
under  setosus  (Eichhoff)  by  Schedl  (1962, 
Ent.  Blatt.  58:204).  Using  the  lectotype 
of  javanus  as  a  point  of  reference,  I 
(Wood,  1972,  Great  Basin  Nat.  32:51) 
added  several  additional  synonyms.  A  re- 
examination of  the  types  of  javanus^  seto- 
sus, and  all  synon}Tns,  and  172  other 
specimens  assigned  to  setosus,  clearly  in- 
dicates the  existence  of  two  easily  dis- 
tinguished species.  Hypothenemus  javanus 
( =  Stephanoderes  obesus  Hopkins,  S.  phil- 
ippinensis  Id^opkins,  S.  bananensis  Eggers, 
S.  kalshoveni  Schedl,  5.  subagnatus  Eg- 
gers) has  a  stouter  body  form  (2.2  times 
as  long  as  wide),  only  12-18  coarse  asper- 
ities on  the  anterior  slope  of  the  pronotum, 
2-4  denticles  on  the  anterior  margin  of  the 
pronotum,  and  several  minor  differences 
in  details  of  sculpture.  It  occurs  in  Indo- 
nesia, Philippines,  Florida,  Cuba,  Jalisco 
(Mexico),  Congo,  Ghana,  and  the  Cam- 
erouns.  Hypothenemus  setosus  has  the 
body  2.4  times  as  long  as  wide,  more  than 
25  pronotal  asperities,  6-8  denticles  on  the 
anterior  margin  of  the  pronotum,  and 
other  differences  in  details  of  surface 
sculpture.  It  occurs  from  Chiapas  (Mex- 
ico) to  Brazil,  in  the  Congo,  and  in  the 
Camerouns.  Therefore,  javanus  must  be 
restored  as  a  valid  name  to  designate  the 
above  species. 


394 


GREAT   BASIN   NATURALIST 


Vol.  35,  No.  4 


Microcorthylus  minutus  Schedl 

Microcorthylus  minutus  Schedl,  1950,  Dusenia  1: 
160  (Syntypes,  females;  Nova  Teutonia, 
Santa  Catarina.  Brazil;  Schedl  and  Plaumann 
colls.) 

Microcorthylus  minutissimus  Schedl,  1952,  Dus- 
enia 3:361  (Syntypes;  Jamaica;  Schedl  Coll.). 
New  synonymy 

Female  syntypes  of  minutus  Schedl  and 
minutissimus  Schedl  were  compared  di- 
rectly to  one  another  and  to  36  other  spec- 
imens from  Veracruz,  Honduras,  Costa 
Rica,  Panama,  and  Brazil.  Only  one  small, 
distinctive  species  is  represented  by  this 
material. 

Pseudopityophthorus  limbatus  Eggers 

Pseudopityophthorus  limbatus  Eggers,  1930,  Ent. 
Blatt.  26:169  (Holotype.  female;  "R.  d.  M.," 
Mexico;  deposited  in  Eggers  collection,  evi- 
dently  now   on   loan   to   Schedl) 

Pseudopityophthorus  micans  Wood,  1967,  Great 
Basin  Nat.  27:44  (Holotype.  male;  96  km 
W.  Durango,  Durango,  Mexico;  Wood  Coll.). 
New  synonymy 

The  female  holotype  of  Pseudopityoph- 
thorus limbatus  Eggers  and  the  female  al- 
lotype of  P.  micans  Wood  were  compared 
directly  to  one  another  and  to  85  other 
specimens  of  this  species  and  were  found 
to  represent  only  one  species. 

Xyleborus  obliquus  (LeConte) 

Pityophthorus  obliquus  LeConte,  1878,  in  Schwarz. 

Proc.    Amer.   Philos.    Soc.    17:432    (Holotype. 

female;     Enterprise,     Florida;     Mus.     Comp. 

Zool.) 
Xyleborus  gilvipes   Blandford,    1898,    Biol.    Centr. 

Amer.,  Coleopt.  4(6):205    (Holotype,  female; 

Zapote,     Guatemala;      British      Mus.),      Neu^ 

synonymy 
Xyleborus   brasiliensis   Eggers,    1928,    Arch.    Inst. 

Biol.,    Sao    Paulo    1:96     (Lectotype,    female; 

Blumenau,  Santa  Catarina.  Brazil;  U.S.  Nat. 

Mus.).  New  synonymy 
Xyleborus  illepidus  Schedl.   1941.  Rev.  Zool.   Bot. 

Afr.     34:402      (Holotype.     female;      Deutsch 

Ostafrika;   Schedl   Coll.).   New  synonymy 

This  species  has  been  reported  from 
areas  disturbed  by  human  activities  in  a 
pattern  that  suggests  introduction.  Its  ori- 
gin has  not  been  established  except  that 
allied  species  are  Neotropical.  Its  occur- 
rence in  Africa  appears  to  be  recent. 

In  establishing  the  above  synonymy, 
the  holotypes  of  obliquus  (LeConte),  gil- 
vipes Blandford,  and  illepidus  Schedl,  the 
lectotype  of  brasiliensis  Eggers,  and  78 
U.S.,  46  Neotropical  (Mexico,  Guatemala, 
Colombia,  Brazil),  and  14  African   (Con- 


go) specimens  were  examined  and  com- 
pared directly.  There  is  a  certain  amount 
of  variability  within  this  material,  but 
the  variation  between  series  is  no  greater 
than  it  is  within  a  series. 

In  addition  to  the  above,  the  synonymy 
of  this  species  with  Under ae  (Hopkins) 
and  mexicanus  Eggers  has  already  been 
established. 

Taxa  New  to  Science 
Dacnophthorus.  n.  gen. 

This  genus  is  distinguished  from  Pity- 
ophthorus Eichhoff  by  the  very  slender 
body  form,  b}^  the  very  large,  coarsely 
faceted  eyes,  by  the  large  antennal  club, 
with  two  partly  septate  sutures,  by  the 
pronotal  summit  being  anterior  to  the 
middle  of  the  pronotum  and  lacking  a 
transverse  impression  posterior  to  it,  by 
the  distinctive  elytral  declivity,  and  by  the 
very  different  habits. 

Frons  dimorphic,  male  convex  above, 
impressed  below,  female  uniformly  shal- 
lowly  concave  to  lower  third,  then  weakly 
convex,  upper  area  variously  sculptured 
and  ornamented  by  hair;  eye  very  large, 
coarsely  faceted,  emarginate.  Antennal 
scape  slender,  elongate;  funicle  5-segmen- 
ted,  some  specimens  apparently  4-segmen- 
ted;  club  large,  much  longer  than  scape, 
sutures  1  and  2  partly  septate,  3  clearly 
indicated  by  setae  and  rather  remote  from 
apex.  Pronotum  elongate,  summit  anterior 
to  middle,  without  a  transverse  impression 
behind  simimit.  Scutellum  rather  large, 
flat.  Elytra  striate;  declivity  rather  short, 
steep.  Legs  as  in  Pityophthorus. 

Type-species. —  Gnathophthorus  de- 
mat  us  Wood. 

Note. —  The  five  species  previously 
I)laced  by  me  in  (rnathophthorus  must  be 
transferred  to  this  genus. 

Araptus  consobrinus^  n.  sp. 

This  species  is  distinguished  from  atten- 
uatus  Wood  by  the  evenly  convex  elytral 
declivity  (attenuatus  has  striae  1  impres- 
sed and  interstriae  1  weakly  elevated) 
i\ui\,  on  the  female  frons,  by  the  coarser, 
more  abimdant,  longer,  yellowish  ves- 
titnre. 

Mai,i:.—  Length  LI-  mm  (])aratypes 
1.4-1.7  mm),  2.7  times  as  long  as  wide; 
( olor  \ery  dark  brown. 


Dec.  1975 


WOOD:     AMERICAN    SCOLYTIDAE 


395 


Frons  similar  to  attenuatus  except  more 
nearly  convex,  punctures  not  as  coarse, 
no  indications  of  aciculation,  median  cal- 
lus at  upper  level  of  eyes  smaller,  less 
strongly  elevated. 

Pronotum  as  in  attenuatus  except  an- 
terior margin  armed  by  about  eight  ser- 
rations. 

Elytra  as  in  attenuatus  except  surface 
less  brightly  shining,  strial  punctures 
slightl}"  larger.  Declivity  more  evenly 
convex,  punctures  much  smaller;  striae 
1  not  impressed,  interstriae  1  not  elevated. 

Female.^ —  Similar  to  male  except  frons 
much  as  in  female  attenuatus  except 
frontal  hair  much  more  abundant,  coarser, 
slightly  longer,  yellowish  in  color. 

Type  Locality. —  Six  km  or  4  miles  N 
Tepic,  Nayarit,  Mexico. 

Type  Material. —  The  male  holotype, 
female  allotype,  and  20  paratypes  were 
taken  at  the  type  locality  on  13-VII-1965, 
1000  m,  No.  241,  from  a  shrub;  seven 
paratypes  are  from  33  km  or  21  miles  N 
Juchitlan,  Jalisco,  Mexico,  3-VII-1965, 
1300  m,  No.  177,  Ficus  twigs  3  cm  in 
diameter,  all  by  me. 

The  holotype,  allotype,  and  paratypes 
are  in  my  collection. 

Araptus  micaceus,  n.  sp. 

This  species  is  distinguished  from  obso- 
letus  (Blandford)  by  the  strongly  im- 
pressed frons,  with  a  pair  of  epistomal 
calluses  at  the  anterior  articulations  of 
the  mandibles,  by  the  smooth  surface  be- 
tween punctures  on  the  area  above  the 
eyes,  and  by  the  coarser  strial  punctures 
on  the  declivity. 

Male. —  Length  1.4  mm  (paratypes 
1.3-1.4  mm),  2.6  times  as  long  as  wide; 
color  light  brown. 

Frons  as  in  obsoletus  except  much  more 
strongly  impressed,  punctures  in  im- 
pressed area  smaller,  surface  on  area 
above  eyes  reticulate;  epistomal  calluses  at 
anterior  articulation  of  mandibles  much 
larger. 

Pronotum  as  in  obsoletus  except  ser- 
rations on  anterior  margin  much  smaller 
(margin  subcostate)  and  reticulation  on 
posterior  areas  more  strongly  impressed. 

Elytra  as  in  obsoletus  except  strial  punc- 
tures   much    larger,    interstriae    one    and 


one-half  times  as  wide  as  striae  on  disc, 
about  twice  as  wide  on  declivity;  im- 
pressed  points   obsolete  on  declivity. 

Female. —  Similar  to  male  except  frons 
planoconvex  on  median  two-thirds  from 
level  of  antennal  insertion  to  slightly 
above  eyes,  finely,  closely  punctured  ex- 
cept impunctate  on  median  line  to  upper 
level  of  eyes,  punctured  area  bearing  fine, 
rather  long,  moderately  abundant  hair; 
hair  shorter,  less  abundant  and  covering 
a  smaller  area  than  in  female  obsoletus. 

Type  Locality. —  Los  Corchos,  Nayarit, 
Mexico. 

Type  Material. —  The  male  holotype, 
female  allotype,  and  seven  paratypes  were 
taken  at  the  type  locality  on  lO-Vn-1965, 
7  m  elevation.  No.  222,  from  a  recently 
cut  vine,  by  me. 

The  holotype,  allotype,  and  paratypes 
are  in  my  collection. 

Pityophthorus  costatus.  n.  sp. 

This  abberant  species  has  a  variable 
number  of  segments  in  the  antennal 
funicle;  it  is  also  distinguished  by  the  long, 
costiform  pronotal  asperities,  by  the  stout 
body  form,  b}'  the  frons,  and  by  other 
characters. 

Female. —  Length  1.1  mm  (paratypes, 
1.1-1.2  mm),  2.3  times  as  long  as  wide; 
color  dark  reddish  brown. 

Frons  rather  strongly  convex,  an  abrupt, 
moderately  deep  impression  immediately 
above  epistomal  margin,  epistomal  margin 
distinctly  elevated  except  for  small  median 
notch;  surface  smooth  shining,  sparsely 
punctured,  punctures  rather  coarse,  area 
above  eyes  somewhat  reticulate;  vestiture 
very  sparse,  short.  Antennal  funicle 
variable,  3-5-segmented;  club  ovate,  small, 
sutures  almost  straight,  1  and  2  septate 
only  at  margins. 

Pronotum  1.0  times  as  long  as  wide; 
widest  at  base,  weakly  arcuate  on  basal 
third  rather  strongly  converging  toward 
narrowly  rounded  anterior  margin;  an- 
terior margin  continuously  costate;  sum- 
mit just  behind  middle,  rather  indefinite; 
asperities  long,  subcostate,  confused,  con- 
tinued to  basal  fourth  in  median  area; 
posterior  areas  strongly  reticulate,  punc- 
tures small,  not  close.  Glabrous. 

Elytra  1.5  times  as  long  as  wide,  1.6 
times  as  long  as  pronotum;  sides  almost 


396 


GREAT   BASIN    NATURALIST 


Vol.35,  No.  4 


Straight  and  parallel  on  basal  two-thirds, 
rather  broadly  rounded  behind;  striae  not 
impressed,  punctures  small,  deej),  spaced 
by  one  to  two  diameters  of  a  puncture; 
interstriae  smooth,  shining,  two  to  three 
times  as  wide  as  striae,  punctures  minute, 
confused,  rather  close.  Declivity  steep, 
convex;  striae  not  impressed,  punctures 
greatly  reduced  to  obsolete;  interstriae  as 
on  disc  except  punctures  reduced  to  obso- 
lete. Vestiture  confined  to  declivity,  of 
sparse,  short,  stout  interstrial  setae. 

Male. —  Similar  to  female  except  epi- 
stomal  impression  usually  more  strongly 
impressed  in  lateral  areas,  with  a  weak 
median  elevation  dividing  this  impression. 

Type  Locality. —  Tapanti,  Cartago, 
Costa  Rica. 

Type  Material. —  The  female  holo- 
type,  male  allotype,  and  12  paratypes 
were  taken  at  the  type  locality  on  2-VII- 
1963,  1300  m.  No.  Ha,  from  a  liana,  by 
me.  Two  paratypes  are  from  13  km  SE 
Cartago,  Cartago,  Costa  Rica,  3-VII-1963, 
1800,  from  the  same  species  of  liana,  by 
me. 

The  holotype,  allotype,  and  paratypes 
are  in  my  collection. 

Pityophthorus  inceptis,  n.  sp. 

This  species  is  distinguished  by  the 
simple  male  frons  and  declivity,  by  the 
female  frontal  vestiture  and  by  the  coarse 
pronotal  and  elytral  punctures.  It  is  not 
closely   allied   to   other   known   species. 

Female. —  Length  1.5  mm  (male  para- 
types 1.5-1.6  mm),  2.7  times  as  long  as 
wide;    color    very    dark    brown. 

Frons  basically  convex  except  flattened 
on  median  half  from  epistoma  to  slightly 
above  eyes;  surface  shining,  punctures 
rather  coarse,  moderately  close;  vestiture 
rather  sparse  and  short  except  on  margins 
of  upper  half  of  flattened  area  forming 
a  dense  fringe  of  long  yellow  hair,  longest 
setae  equal  in  length  to  three-fourths  dis- 
tance from  their  bases  to  epistomal  margin. 
Antennal  club  broadly  obovate,  1.2  times 
as  long  as  wide,  suture  1  almost  straight, 
2  moderately   ])rocurved. 

Pronotum  1 .07  times  as  long  as  wide; 
widest  on  basal  third,  moderately  arcuate 
from  base  to  rather  broadly  rounded  an- 
terior margin;  anterior  margin  armed  by 
about  six  to  eight  low,  basally  contiguous 


teeth;  summit  at  middle;  asperities  rather 
coarse,  confused;  posterior  areas  subrug- 
ose-reticulate,  punctures  coarse,  deep, 
mostly  spaced  by  distances  equal  to  one- 
half  diameter  of  a  puncture,  median  line 
impunctate.  Vestiture  of  sparse  semi- 
recumbent  short  hair  in  lateral  and  as- 
perate areas. 

Elytra  1.7  times  as  long  as  wide,  1.7 
times  as  long  as  pronotum;  sides  almost 
straight  and  parallel  on  basal  three- 
fourths,  somewhat  narrowly  rounded  be- 
hind; striae  not  impressed,  punctures 
rather  coarse,  deep,  occasional  punctures 
not  in  row,  spaced  by  less  than  diameter 
of  a  puncture;  interstriae  almost  smooth, 
shining,  impunctate  except  for  an  oc- 
casional puncture  near  declivity.  Declivity 
steep,  convex;  striae  not  impressed,  punc- 
tures reduced,  about  one-third  as  large  as 
on  disc,  distinctly  impressed;  interstriae 
as  on  disc  except  1  and  3  each  with  a  row^ 
of  fine  punctures.  Vestiture  confined  to 
sides  and  declivity,  that  on  sides  of  minute 
strial  hair,  that  on  declivity  of  fine,  rather 
short  interstrial  setae  on  odd-numbered 
interstriae. 

Male. —  Similar  to  female  except  frons 
more  uniformly  convex,  without  brush  of 
long  hair;  serrations  on  anterior  margin 
of  pronotum  slightly  larger. 

Type  Locality. —  Six  km  or  four  miles 
W  Quiroga,  Michoacan,  Mexico. 

Type  Material. —  The  female  holo- 
type, male  allotype,  and  one  male  para- 
type  were  taken  at  the  type  locality  on 
17-VI-1965,  2200  m.  No.  72,  from  a  shrub- 
by herbaceous  ]:)lant,  by  me. 

The  holotype,  allotype,  and  paratype 
are  in  my  collection. 

Pityophthorus  timididus.  n.  sp. 

This  species  is  distinguished  from  men- 
dosus  Wood  by  the  larger  size,  by  the 
coarser  pronotal  punctures,  and  by  slight 
differences  on  the  elytral  declivity.  Both 
species  are  allied  to  mandihularis  Schedl. 

Male. —  Length  2.0  mm  (paratypes 
1.8-2.0  mm),  2.5  times  as  long  as  wide; 
color  very  dark   brown. 

Frons  convex  above  eyes,  upper  half  of 
area  below  upper  level  of  eyes  rather 
abruptly,  strongly,  transversely  impressed 
almost  from  eye  to  eye,  a  smaller  trans- 


Dec.  1975 


WOOD:     AMERICAN    SCOLYTIDAE 


397 


verse  impression  in  lateral  areas  immed- 
iately above  epistoma;  surface  shining, 
coarsely,  rather  closely  punctured;  ves- 
titure  sparse  exce])t  on  epistoma.  Antennal 
club  oval,  1.3  times  as  long  as  wide,  su- 
tures 1  and  2  moderately  arcuate,  2  at 
middle  of  club. 

Pronotum.  1.1  times  as  long  as  wide; 
sides  on  basal  half  almost  straight,  sub- 
parallel,  rather  broadly  rounded  in  front; 
anterior  margin  armed  by  about  12  low 
serrations;  summit  at  middle;  asperities 
on  anterior  slope  rather  coarse,  close,  con- 
fused; posterior  areas  smooth,  shining, 
with  moderately  abundant  minute  im- 
pressed points,  punctures  rather  coarse, 
deep,  moderately  close,  irregularly  spaced 
by  about  one  to  two  diameters  of  a  punc- 
ture. Glabrous  except  a  few  setae  on 
margins. 

Elytra  1.6  times  as  long  as  wide,  1.6 
times  as  long  as  pronotum;  sides  almost 
straight  and  parallel  on  basal  two-thirds, 
rather  broadly  rounded  behind;  striae  1 
weakly,  others  not  impressed,  punctures 
rather  small,  deep,  spaced  by  diameter  of 
one  puncture;  interstriae  smooth,  shining, 
a  few  small  punctures  on  1  near  declivity, 
others  impunctate.  Declivity  steep,  con- 
vex, shallowly  bisulcate;  interstriae  1  dis- 
tinctly elevated,  armed  by  a  row  of  about 
seven  rounded  tubercles,  2  moderately 
impressed,  slightly  wider  than  1,  smooth, 
shining,  devoid  of  punctures,  3  convex,  as 
high  as  1,  armed  as  on  1,  lateral  areas 
with  punctures  somewhat  confused.  Ves- 
titure  confined  to  declivity,  consisting  of 
rather  short,  moderately  coarse,  sparse, 
interstrial  setae  on  odd-numbered  inter- 
striae. 

Female. —  Similar  to  male  except  frons 
below  upper  level  of  eyes  shallowly, 
broadly,  transversely  impressed,  surface 
regular,  not  granulate,  punctures  moder- 
ately fine,  vestiture  sparse,  declivital  sul- 
cus much  less  strongly  impressed,  gran- 
ules on  interstriae  1   and  3  minute. 

Type  Locality. —  Volcan  Chiriqui, 
Chiriqui,  Panama. 

Type  Material. —  The  male  holotype, 
female  allotype,  and  12  paratypes  were 
taken  at  the  type  locality  on  11-1-1964, 
1800  m,  No.  407,  from  a  sapling  4  cm  in 
diameter,  by  me. 

The  holotype,  allotype,  and  paratypes 
are  in  my  collection. 


Pityophthorus  mcndosiis,  n.  s]). 

This  species  is  distinguished  from 
limidulus  Wood  by  the  smaller  size,  by 
the  smaller  pronotal  punctures,  by  the 
shorter,  stouter  elytra  1  bristles,  and  by  the 
( omplete  absence  of  tubercles  on  female 
declivital  interstriae   1    and   3. 

Male. —  Length  1.7  mm  (paratypes 
1.5-1.7  mm),  2.8  times  as  long  as  wide; 
color  very  dark  brown. 

Frons  as  in  timidulus  except  callus  at 
level  of  antennal  insertion  usually  more 
strongly  develojied  and  with  a  small  cusp 
at  dorsomedian  extremity.  Pronotal  punc- 
tures averaging  slightly  smaller  than  in 
timidulus.  Elytra  as  in  timidulus  except 
declivital  setae  slightly  shorter  and  dis- 
tinctly stouter. 

Female. —  As  in  female  timidulus  ex- 
cept frons  less  distinctly  impressed,  decliv- 
ital granules  absent,  and  declivital  setae 
shorter  and  stouter. 

Type  Locality. —  San  Isidro  del  Gen- 
eral, San  Jose,  Costa  Rica. 

Type  Material. —  The  male  holotype, 
female  allotype,  and  18  j)ara types  were 
taken  at  the  type  locality  on  5-Xn-63, 
1000  m.  No.  282,  from  "Fosforo"  leaf 
petioles,  by  me. 

The  holotype,  allotype,  and  paratypes 
are  in  my  collection. 

Pityophthorus  degener,  n.  sp. 

This  species  is  distinguished  from  the 
allied  timidulus  Wood  and  mendosus 
Wood  by  the  much  more  weakly  im- 
pressed male  frons,  by  the  strongly  im- 
pressed elytral  declivity,  and  by  the  finer 
pronotal  punctures. 

Male. —  Length  2.0  mm  (paratypes  1.7- 
2.0  mm),  2.8  times  as  long  as  wide;  color 
very  dark  brown. 

Frons  convex,  but  with  abrupt,  irreg- 
ular, shallow,  transverse  impressions  just 
below  upper  level  of  eyes  and  just  above 
epistoma;  surface  subshining,  coarsely, 
closely  punctured;  almost  glabrous  ex- 
cept at  epistomal  margin.  Pronotum  and 
elytral  disc  as  in  timidulus  and  mendosus 
except  pronotal  punctures  much  smaller, 
slightly  closer;  elytral  declivity  strongly 
bisulcate,  interstriae  1  almost  as  high  as 
wide,  3  higher  than  1,  each  armed  by  a 


398 


GREAT   BASIN    NATURALIST 


Vol.  35,  No.  4 


row  of  moderately  coarse  tubercles.  Ely- 
tral  vestiture  as  in  timidulus. 

Female. —  Similar  to  male  except  me- 
dian half  of  frons  flattened  below  upper 
level,  surface  smooth,  with  punctures 
rather  fine,  close,  deep,  with  rather  abun- 
dant, fine,  moderately  long  hair;  declivital 
sulcus  half  as  deep,  interstriae  1  and  3 
unarmed. 

Type  Locality. —  Volcan  Chiriqui, 
Chiriqui,  Panama. 

Type  Material. —  The  male  holotype, 
female  allotype,  and  seven  paratypes 
were  taken  at  the  type  locality  on  11-T 
1964,  1800  m.  No.  384,  from  a  tree  limb 
8  cm  in  diameter,  by  me. 

The  holotype,  allotype,  and  paratypes 
are  in  my  collection. 

Pityophthorus  amiculus,  n.  sp. 

This  species  is  distinguished  from  deg- 
encr  Wood  by  the  absence  of  sexual  di- 
morphism, and  by  the  different  frons, 
pronotum,  and  other  characters. 

Male. — Length  1.8  mm  (paratypes  1.7- 
1.9  mm),  2.7  times  as  long  as  wide;  color 
reddish  brown. 

Frons  convex  above  eyes,  with  median 
line  smooth,  impunctate,  a  median  callus 
at  upper  level  of  eyes,  area  below  upper 
level  of  eyes  broadly,  shallowly,  trans- 
versely impressed  to  epistoma;  surface 
almost  smooth,  punctures  moderately 
coarse,  close,  deep;  almost  glabrous  ex- 
cept along  epistoma. 

Pronotum  1.2  times  as  long  as  wide; 
outline  as  in  degener;  asperate  area  con- 
tinued slightly  into  lateral  portion  of  pos- 
terior half  as  weak  rugae;  posterior  areas 
smooth,  shining,  with  rather  numerous 
impressed  points,  punctures  rather  fine, 
moderatel}"  close,  median  line  impunc- 
tate. Sparse  setae  confined  to  asperate 
area. 

Elytra  1.5  times  as  long  as  wide,  1.3 
times  as  long  as  pronotum;  outline  as  in 
timidulus  Wood;  striae  not  impressed, 
punctures  rather  small,  deep,  close,  rows 
occasionally  slightly  confused;  interstriae 
smooth,  shining,  slightly  irregular,  with 
a  few  impressed  lines,  impressed  points 
moderately  abundant,  1  with  obscure  sub- 
granulate  punctures  almost  to  base,  others 
with  an  occasional  similar  puncture.  De- 


clivity steep,  strongly  bisulcate,  sulcus 
commencing  slighth'  behind  middle  of  ely- 
tral  length;  striae  1  and  2  with  punctures 
reduced  in  size  but  clearly  impressed; 
interstriae  1  almost  as  high  as  wide,  3 
higher  than  1,  each  armed  by  a  row  of 
about  eight  subacute  tubercles  of  moderate 
size,  2  wider  than  1,  its  surface  ascending 
laterally,  unarmed.  Vestiture  consisting  of 
interstrial  bristles  on  declivity,  continued 
to  middle  of  disc  on  odd-numbered  inter- 
striae; longest  bristles  rather  slender, 
equal  in  length  to  twice  width  of  an  in- 
terstriae. 

Female. —  Similar  to  male  in  all  re- 
spects. 

Type  Locality. —  Guapiles,  Limon, 
Costa  Rica. 

Type  M.a.terial. — The  male  holotype, 
female  allotype,  and  12  paratypes  were 
taken  at  the  type  locality  on  22-Vin- 
1966,  100  m.  No.  121,  from  a  liana,  by  me. 
Eighteen  paratypes  are  from  Coatzocoal- 
cos,  Veracruz,  Mexico,  26-VL1967,  30  m. 
No.  103,  liana,  by  me. 

The  holotype,  allotype,  and  paratypes 
are  in  my  collection. 

Pityophthorus  dissolutus.  n.  sp. 

This  species  is  distinguished  from  ex- 
plicitus  Wood  by  the  finer  frontal  punc- 
tures, by  the  finer  granules  on  the  pro- 
notal  disc,  and  by  the  shallower  declivital 
sulcus  that  is  armed  by  finer  granules. 

Male. — Length  1.5  mm  (paratypes 
1.4-1.6  mm),  2.7  times  as  long  as  wide; 
color  dark  brown. 

Frons  convex,  a  fine  median  tubercle 
on  epistomal  process;  surface  strongly  re- 
ticulate, punctures  rather  fine,  deep, 
spaced  by  diameter  of  a  puncture  or  more; 
vestiture  fine,  sparse,  inconspicuous. 

Pronotum  1.1  times  as  long  as  wide; 
widest  on  basal  half,  sides  feebly  arcuate, 
subjiarallel.  rather  narrowly  rounded  in 
front;  anterior  margin  armed  by  about 
eight  moderately  coarse  serrations;  sum- 
mit at  middle,  indefinite;  asperities  rather 
fine,  confused;  posterior  areas  strongly  re- 
ticulate, j)unctures  rather  fine,  moderately 
close.  Vestiture  confined  to  marginal  and 
asperate  areas. 

Elytra  1.6  times  as  long  as  wide,  1.5 
times  as  long  as  pronotum;   sides  almost 


Dec.  1975 


WOOD:    AMERICAN    SCOLYTIDAE 


399 


straight  and  parallel  on  basal  two-thirds, 
rather  broadly  rounded  behind;  striae  1 
weakly,  others  not  impressed,  punctures 
fine,  distinct,  decreasing  in  size  posterior- 
ly; interstriae  almost  smooth,  with  some 
indistinctly  impressed  lines,  about  three 
times  as  wide  as  striae  on  basal  fourth, 
six  times  as  wide  near  declivity.  Declivity 
rather  steep,  shallowly  bisulcate;  striae  1 
deejily  impressed,  jninctures  small,  indis- 
tinct, surface  ascending  gradually  to 
lateral  convexity,  striae  2  obscure;  inter- 
striae 1  almost  as  high  as  wide,  almost 
smooth,  with  a  row  of  fine  tubercles,  2  and 
lateral  areas  shining,  rather  densely  cov- 
ered by  impressed  points,  3  slightly  higher 
than  1,  similarly  armed.  Vestiture  of 
minute  strial  hair,  and,  on  posterior  half, 
interstrial  bristles  on  odd-numbered  inter- 
striae; bristles  sparse,  rather  fine,  short. 

Female.^ —  Similar  to  male  except  epi- 
stomal  tubercle  evidently  absent,  and  de- 
clivital  impressed  points  reduced  or  ab- 
sent. 

Type  Locality. —  Thirteen  km  SE  Car- 
tago,  Cartago,  Costa  Rica. 

Type  Material. — The  male  holotype, 
female  allotype,  and  27  paratypes  were 
taken  on  24-IX-1963,  1800  m.  No.  201, 
from  a  liana  1  cm  in  diameter.  Two  para- 
types are  from  Tapanti,  Cartago,  Costa 
Rica,  24-X-63,  1300  m.  No.  244,  liana; 
and  six  paratypes  are  from  Volcan  Chir- 
iqui,  Chiriqui,  Panama,  11-1-1964,  No. 
394,  in  a  sapling;  all  were  taken  by  me. 

The  holotype,  allotype,  and  paratypes 
are  in  my  collection. 

Pityophthorus  explicitus,  n.  sp. 

This  species  is  distinguished  from 
dissolutus  Wood  by  the  coarser  frontal 
punctures,  by  the  larger  granules  on  the 
pronotal  disc,  and  by  the  deeper  declivital 
sulcus  that  is  armed  by  coarser  granules. 

Male. —  Length  1.5  mm  (paratypes 
1.5-1.7  mm),  2.7  times  as  long  as  wide; 
color  very  dark  brown. 

Frons  as  in  dissolutus  except  punctures 
distinctly  larger,  spaced  by  less  than  di- 
ameter of  a  puncture.  Pronotum  as  in 
dissolutus  except  granule  on  lateral  mar- 
gins of  discal  punctures  distinctly  larger. 
Elytra  as  in  dissolutus  except  declivital 
sulcus  deeper,  wider,  interstriae  2  not 
ascending  laterally  on  lower  half. 


Female. —  Female  similar  to  male  in 
all  respects. 

Type  Locality. —  Nine  km  NE  Teziut- 
lan,  Puebla,  Mexico. 

Type  Material. —  The  male  holotype, 
female  allotype,  and  14  paratypes  were 
taken  at  the  type  locality  on  2-Vn-1967, 
1600  m.  No.  143,  from  a  liana,  by  me. 

The  holotype,  allotype,  and  paratypes 
are  in  my  collection. 

Xyleborus  calif ornicus,  n.  sp. 

This  species  almost  certainly  was  intro- 
duced into  California  from  another  area, 
})ossibly  from  South  America  or  south- 
eastern Asia.  Among  North  American 
species  it  might  be  confused  with  pubes- 
cens  Zimmermann,  but  it  is  distinguished 
by  the  smaller  size,  by  the  more  abun- 
dant pubescence,  and  by  the  reticulate- 
granulate  interiors  of  the  strial  punctures 
on  the  declivity. 

Female. — Length  2.0  mm  (paratypes 
2.0-2.2  mmj,  2.9  times  as  long  as  wide; 
color   yellowish   brown. 

Frons  rather  strongly  convex;  surface 
strongly  reticulate,  a  few  small  granules 
from  epistoma  to  upper  level  of  eyes.  Ves- 
titure of  fine,  sparse  hair. 

Pronotum  1.2  times  as  long  as  wide; 
sides  almost  straight  and  parallel  on  basal 
two-thirds,  rather  broadly  rounded  in 
front;  anterior  margin  unarmed;  summit 
in  front  of  middle;  anterior  slope  steep, 
rather  coarsely  asperate;  posterior  areas 
strongly  reticulate,  punctures  small,  shal- 
low, rather  close.  Vestiture  of  fine,  short, 
rather  abundant  hair. 

Elytra  1.7  times  as  long  as  wide,  1.4 
times  as  long  as  pronotum;  sides  almost 
straight  and  parallel  on  basal  two-thirds, 
broadly  rounded  behind;  disc  occupying 
basal  three-fourths;  striae  not  impressed, 
punctures  small,  shallow,  distinct,  in 
rows,  spaced  by  diameter  of  a  puncture; 
interstriae  three  to  four  times  as  wide  as 
striae,  almost  smooth,  shining,  punctures 
fine,  in  definite  rows  in  some  specimens, 
distinctly  confused  on  basal  half  in  others. 
Declivity  steep,  convex,  general  contours 
as  in  pubescens;  strial  punctures  large, 
shallow,  distinct,  their  interior  surfaces 
reticulate-granulate;  interstriae  only 
slightly  wider  than  striae,  their  punctures 
mostly   replaced   by  minute   granules   on 


400 


GREAT   BASIN    NATURALIST 


Vol.  35,  No.  4 


all  interstriae,  a  few  larger  granules  on  1, 
3,  and  lateral  areas;  posterolateral  mar- 
gin rounded,  with  an  indefinite  row  of 
scattered  granules.  Vestiture  of  rather 
abundant,  short,  fine  hair,  distinctly 
longer  on  margins  of  declivity. 

Type  Locality. —  Stanford  University, 
Palo  Alto,  California. 

Type  Material. — The  female  holotype 
and  five  female  paratopes  were  taken  at 
the  type  locality  on  15 -25 Til- 1944,  by  C. 
D.  Duncan.  One  paratype  is  from  Knight's 
Landing,  Yolo  Co.,  California,  10-M949, 
at  light,  by  J.  R.  Fowler. 

The  holotype  is  in  the  California  Acad- 
emy of  Sciences;  three  para  types  are  in 
the  Canadian  National  Collection,  two  are 
in  my  collection,  and  one  is  in  the  U.S. 
National  Museum. 

Specimens  of  this  species  were  sent  to 
Bright,  Browne,  and  Schedl  in  an  effort 
to  associate  it  with  a  named  species  from 
another  area,  but  without  success.  All 
agree  that  its  anatomical  characters,  its 
current  distribution,  and  the  limited  period 
it  has  been  known  in  California  strongly 
suggest  that  it  was  introduced  from 
another  area. 

Xyleborus  incultus.  n.  sp. 

This  species  is  distinguished  from  pristis 
Wood  by  the  larger  size,  by  the  more  dis- 
tinctly impressed  strial  punctures,  and  by 
the  declivital  sculpture  as  described  be- 
low. 

Female. —  Length  2.3  mm,  2.6  times 
as  long  as  wide;  color  dark  brown. 

Frons  and  pronotum  essentially  as  in 
micarius  Wood  except  anterior  margin  of 
pronotum  weakly  produced  at  median  line 
and  armed  by  two  larger  serrations,  punc- 
tures on  posterior  areas  larger,  at  least 
twice  as  large  as  in  micarius. 

Elytra  1.5  times  as  long  as  wide,  1.3 
times  as  long  as  j)ronotum;  sides  almost 
straight  and  y)arallel  on  basal  60  percent, 
narrowly  roundofl  behind,  a  slight  emargi- 
iiation  at  suture;  striae  not  impressed, 
{junctures  small,  distinctly  impressed, 
spaced  within  a  row  by  one  to  two  diam- 
eters of  a  puncture;  interstriae  almost 
smooth,  punctures  fine,  less  than  half  as 
large  as  those  of  striae,  moderately  con- 
fused on  basal  third  of  disc,  uni.seriate  be- 
hind. Declivity  rather  steep,  convex;  sur- 


face minutely  granular  on  lower  two- 
thirds,  dull;  interstriae  1  moderately  ele- 
vated to  near  apex,  armed  by  15  or  more 
small,  pointed,  confused  denticles,  2  rather 
strongly  impressed,  armed  on  upper  third 
by  4-6  similar,  uniseriate  denticles,  3 
slightly  elevated,  armed  as  in  1  except 
denticles  almost  uniseriate,  4-9  each  with 
a  row  of  fine  denticles,  3  and  9  join  and 
continue  submarginally  to  apex  as  a  mod- 
erately strong  elevation,  its  crest  armed 
b}'  a  row  of  about  four  denticles.  Vestiture 
largely  confined  to  sides  and  declivity;  of 
fine,  short  strial  and  slightly  longer  in- 
terstrial  hair,  longest  setae  about  equal 
in  length  to  width  of  an  interstriae. 

Type  Locality. —  Fort  Clayton,  Canal 
Zone,  Panama. 

Type  Material. —  The  female  holotype 
was  taken  at  the  type  locality  on  22-Xn- 
1963,  from  a  Cecropia  branch,  by  me. 

The  holotype  is  in  my  collection. 

Xyleborus  molestulus.  n.  sp. 

This  species  is  distinguished  from  cri- 
nitulus  Wood  by  the  interstrial  granules 
on  the  disc,  by  the  uniseriate  rows  of  ely- 
tral  setae,  and  by  other  characters. 

Female. —  Length  2.0  mm  (paratypes 
2.0-2.2  mm),  2.2  times  as  long  as  wide; 
color  dark  brown. 

Frons  as  in  crinilulus  except  less  strong- 
ly reticulate,  more  sparsely  punctured. 
Pronotum  about  as  in  crinitulus  except 
anterior  margin  armed  by  eight  serrations, 
]:)osterior  areas  very  finely,  shallowly 
punctured. 

Elytra  1.2  times  as  long  as  wide,  1.3 
times  as  long  as  pronotum;  sides  almost 
straight  and  parallel  on  basal  two-thirds, 
broadly  rounded  behind;  striae  not  im- 
pressed, ])unctures  rather  small,  shallow, 
distinct,  spaced  by  one  to  two  diameters 
of  a  puncture;  interstriae  almost  smooth, 
three  times  as  wide  as  striae,  each  with  a 
uniseriate  row  of  fine  tubercles  from  base 
to  a])ex.  Declivity  occupying  posterior 
half  of  elytral  length,  moderately  steep, 
broadly  convex;  sculpture  as  on  disc  ex- 
cept striae  1  feebly  impressed;  postero- 
lateral margin  acutely,  subcrenulately 
elevated  from  apex  to  interstriae  7.  Ves- 
titure of  interstrial  rows  .  of  long,  erect 
hair  from  base  to  apex,  alternate  setae  in 
each   row   twice   as   long  as   distance   be- 


Dec.  1975 


WOOD:    AMERICAN    SCOLYTIDAE 


401 


tween    rows    and    alternating    with    setae 
half  that  length  on  both  disc  and  declivity. 

Type  Locality. —  Barro  Colorado  Is- 
land,  Canal   Zone,    Panama. 

Type  Material. —  The  holotype  and 
six  para  types  were  taken  at  the  type  lo- 
cality on  27-XII-1963,  No.  348,  from  a 
tree  branch.  Four  paratypes  are  from  Fort 
Clayton,  Canal  Zone,  Panama,  22-XII- 
1963,  No.  363,  from  a  tree  branch;  three 
paratypes  are  from  Madden  Forest,  Canal 
Zone,  Panama,  2-1-1964,  70  m.  No.  365, 
from  a  tree  seedling;  and  one  paratype  is 
from  13  km  (8  miles)  S  El  Hato  del  Vol- 
can,  Chiriqiii,  Panama,  7-1-64,  tree  seed- 
Hng.  All  were  taken  by  me. 

The  holotype  and  paratyjies  are  in  my 
collection. 

Xyleborus  tristiculus.  n.  sp. 

This  species  is  distinguished  from  rno- 
lestulus  Wood  b}"  the  more  slender  body, 
by  the  larger  pronotal  punctures,  and  by 
the  differences  on  the  elytra  described 
below. 

Female. —  Length  2.0  mm  (paratypes 
1.9-2.1  mm),  2.4  times  as  long  as  wide; 
color  brown. 

Frons  about  as  in  molestiilus  except 
imiformly,  more  strongly  reticulate. 

Pronotum  1.02  times  as  long  as  wide; 
as  in  rnolestulus  except  median  part  of 
anterior  margin  weakly  produced,  four 
median    serrations    distinctly   larger,    and 


punctures  on  posterior  areas  distinctly 
larger,  spaced  by  at  least  two  to  six  diam- 
eters of  a  puncture;  vestiture  finer,  long- 
er, more  abundant. 

Elytra  1.3  times  as  long  as  wide,  1.3 
times  as  long  as  pronotum;  similar  to 
rnolestulus  in  outline  except  more  nar- 
rowly rounded  behind;  about  as  in  rno- 
lestulus except  discal  interstriae  not  as 
smooth,  interstrial  punctures  very  feebly 
if  at  all  granulate,  declivital  striae  slightly 
smaller,  interstriae  not  as  smooth,  inter- 
strial tubercles  more  closely  spaced,  av- 
eraging smaller.  Vestiture  consisting  of 
rather  long,  fine,  semirecumbent  strial 
hair  on  disc  and  declivity,  and  rows  of 
erect  interstrial  setae  on  declivity  and 
j:)Osterior  half  of  disc,  each  seta  coarser 
and  distinctly  longer  than  those  of  striae, 
distance  between  rows  and  between  setae 
within  a  row  about  two-thirds  length  of 
a  seta. 

Type  Locality.—  Brazil,  12°  49'  S  51° 
46'  W  (not  12°  31'  S  as  given  on  label). 

Type  Material. —  The  female  holotype 
and  four  female  paratypes  were  taken  at 
the  tvpe  locality  8-XL1968,  No.  C-47,  22- 
XL1968,  No.  D-35,  24-IX-1968,  Nos.  170, 
172,  2-Xn-1968,  No.  E-18,  all  by  R.  A. 
Beaver. 

The  holotype  is  in  the  British  Museum 
(Natural  History),  one  paratype  is  in 
Museo  de  Zoologia,  Universidade  de  Sao 
Paulo,  and  three  paratypes  are  in  my  col- 
lection. 


CORRELATES  OF  BURROW  LOCATION  IN  BEECHEY 
GROUND  SQUIRRELS 

Donald    H.    Owings^    and    Mark    Borchert' 

Abstract. —  Partial  correlation  analysis  indicates  that  Beechey  ground  squirrels  show  a  strong  pref- 
erence for  digging  burrows  under  and  around  large  objects,  ma>-  show  a  weaker  tendency  to  locate 
tJieir  burrows  under  the  cover  of  tree  canopies,  and  avoid  digging  burrows  in  areas  with  both  tree- 
canopy  and  ground  cover  (stones,  logs).  These  relationships  hold  for  large  but  not.  small  burrow  sys- 
tems. The  need  for  unobstructed  visual  surveillance  and  an  autumn  food  supply  are  proposed  to  be 
detenninants  of  tliese  preferences. 


Beeche}'  ground  squirrels  {Spermoph- 
ilus  beecheyi)  appear  to  show  preferences 
for  areas  with  certain  characteristics  for 
the  location  of  their  burrows.  At  least 
three  factors  have  been  suggested  to  af- 
fect the  choice  of  burrow  sites:  (1)  bur- 
rows are  often  constructed  under  large 
objects  such  as  stones  or  logs  (Lins- 
dale  1946:  9);  (2)  good  drainage  is  said 
to  be  iinportant,  which  is  best  provided 
by  sloping  terrain  (Tomich  1962);  (3) 
burrows  often  seem  to  be  concentrated 
under  the  cover  of  tree  canopies  (Fitch 
1948).  One  purpose  of  the  research  re- 
ported here  was  to  quantitatively  assess 
the  relationship  between  burrow  location 
and  these  three  independent  variables — 
ground  cover  (stones,  logs),  slope  of  ter- 
rain, and  tree  canopy  cover.  In  addition, 
we  felt  that  the  effect  of  ground  and  tree 
cover  might  interact,  or  at  least  be  ad- 
diti^'e,  in  areas  in  which  these  factors 
physically  overlapped.  We  therefore  ad- 
ded a  fourth  independent  variable  (com- 
mon cover)  to  the  analysis  which  consis- 
ted of  a  measure  of  the  amount  of  area 
having  ground  and  tree  cover  in  common. 

It  is  probably  true,  though,  that  bur- 
row systems  of  different  sizes  may  dif- 
fer in  their  relationship  with  these  var- 
iables. Small  systems,  for  instance,  may 
be  established  by  young  squirrels  who 
have  moved  into  less  than  optimal  habitat 
during  dispersal,  or  by  adults  for  refuge 
from  predators  when  feeding  in  the  open 
(Fitch  1948;  Carl  1971).  A  second  pur- 
pose of  this  research  was  to  divide  the 
burrow  systems  into  size  categories  for 
separate  analysis. 

Study  Area 

This  study  w^as  done  in  the  Department 
of   Zoology   Experimental   Wildlife   Area 

^Departxiient    of    Psychology,    University    of    California.    Davi 


on  the  campus  of  the  University  of  Cali- 
fornia at  Davis  (elevation  about  15.85  m; 
38°  32':  N,  121°  47':  W).  The  study 
plot  was  located  in  the  82-m  wide  origi- 
nal bed  of  Putah  Creek  which  is  now 
permanently  dry.  This  area  contained 
numerous  trees  (principally  black  wal- 
nut, Juglans  hindsi.  and  valley  oak,  Quer- 
cus  lobata),  grasses  (principally  ripgut 
brome,  Brornus  rigidus^  with  some  Italian 
ryegrass,  Lolium  multiflorum,  and  wild 
oats,  Avena  fatua),  and  thistle  (Centau- 
rea  solstitialis  and  Silybum  marianum)^ 
as  well  as  logs,  and  included  most  of  both 
sloping  sides  of  this  cross-section  of  the 
bed.  A  substantial  population  of  squirrels 
inhabited  this  area:  44  were  trapped  and 
marked  in  0.60  ha  in  the  spring  of  1973. 
This  area  was  being  mapped  in  prepara- 
tion for  behavioral   studies. 

Methods 

Our  procedure  was  to  lay  out  a  grid  of 
9.14  X  9.14  m  squares  and  to  map  on 
graph  paper  the  location  of  all  burrow 
entrances,  the  location  and  size  (to  scale) 
of  ground  cover,  and  the  outer  limits  of 
tree  canopies  for  each  of  50  of  these 
squares.  We  derived  measures  of  ground 
cover  area,  tree  canopy  area,  common 
area,  and  nimiber  of  burrow  entrances 
from  these  maps.  Slope  was  measured  over 
uniform  sections  of  the  area;  new  mea- 
sures were  taken  wherever  significant 
changes  in  slope  occurred. 

We  assumed  that  size  of  burrow  sys- 
tem was  positively  correlated  with  num- 
bers of  entrances.  We  used  a  portable 
blower  (Steco  Model  DS-5)  and  non- 
toxic smoke  bombs  (Superior  0.5  min) 
to  assess  the  number  of  entrances  in  a 
system   by   blowing   smoke   into   one   en- 

s.     California     95010, 


402 


Dec.  1975 


OWINGS.   BORCHERT:    GROUND   SQUIRRELS 


403 


trance   and   designating   as    connected    to 
that  entrance  all  burrows  emitting  smoke. 

Results  and  Discussion 

In  order  to  minimize  the  problem  of 
correlated  independent  variables,  we  cal- 
culated partial  correlation  coefficients. 
This  statistic  provides  a  measure  of  the 
correlation  of  burrows  with  each  inde- 
pendent variable,  while  holding  the  ef- 
fects of  the  remaining  independent  var- 
iables constant.  The  results  of  this  analy- 
sis can  be  found  in  Table  1.  The  first 
(uppermost)  row  includes  burrow  sys- 
tems of  all  sizes  and  suggests  that  Beech- 
ey  ground  squirrels  (1)  show  a  strong 
preference  for  digging  burrows  under  and 
around  large  objects,  (2)  may  show  a 
weaker  tendency  to  locate  their  burrows 
under  the  cover  of  tree  canopies,  but  (3) 
avoid  digging  burrows  in  areas  with  both 
tree-canopy  and  ground  cover. 

In  order  to  test  the  hypothesis  that 
the  larger  systems  were  the  ones  most 
consistently  associated  with  our  indepen- 
dent variables,  we  successively  separated 
increasing  sizes  of  small  systems  and 
analyzed  them  separately  from  the  larger 
systems.  The  results  of  these  analyses  are 
presented  in  the  remaining  rows  of  Table 
1 .  It  is  clear  that  there  is  little  effect  upon 
the  relationship  between  burrows  and 
ground  cover  of  deleting  from  the  analy- 
sis burrow  systems  with  1  to  4  entrances. 
However,  deletion  of  5-entrance  systems 
from  the  large-system  analysis  caused   a 


large  drop  in  the  correlation  coefficient. 
Addition  of  5-entrance  systems  to  the 
small-system  analysis  results  in  the  first 
significant  correlation  with  ground  cover. 
We  conclude  that  small  burrow  systems 
tend  not  to  be  associated  with  ground 
cover,  whereas  larger  systems  (5  or  more 
entrances)  do. 

The  negative  correlation  with  common 
cover  and  the  positive  correlation  with 
tree  cover  followed  the  same  pattern  when 
system  size  was  manipulated.  As  a  result, 
we  conclude  that  it  is  principally  the 
larger  systems  which  tend  not  to  be  as- 
sociated with  areas  having  overlapping 
ground  and  tree  cover,  but  which  are  lo- 
cated under  or  near  tree  cover. 

Our  behavioral  observations  indicate 
that  squirrel  concentration  was  correlated 
with  burrow  concentration,  and  G.  L. 
Hunt  (unpubl.  data)  has  quantitatively 
docmnented  this  in  another  location  (r= 
0.80  -  0.85  between  numbers  of  squirrels 
and  burrow  entrances).  We  assume  then 
that  our  data  indicate  microhabitat  se- 
lection by  these  ground  squirrels. 

We  feel  that  the  strongest  determinant 
of  this  propensity  to  live  in  the  vicinity 
of  ground  cover  is  the  visibility  it  pro- 
vides. Hunt  (unpubl.  data)  has  gathered 
considerable  support  for  the  hypothesis 
that  Beechey  ground  squirrels  in  a  much 
more  open  area  tend  to  concentrate  in 
locations  in  which  their  ground  level 
view  (probably  of  approaching  predators) 
is  least  obstructed  by  locating  themselves 


Table  1  Partial  correlation  coefficients  between  the  numbers  of  burrow  entrances  from  burrow 
system  of  various  sizes  (dependent  variable)  and  the  independent  variables  (1)  slope.  (2)  ground  cov- 
er, (3)  tree  cover,  and  (4)  common  cover.  Size  of  burrow  system  is  expressed  as  the  number  of  en- 
trances  to   the   system. 


Burrow   system 

sizes 

Number   ( 

Df   entrances 

Partial 

correlation 

coefficients   of 

entrances   with 

included  in  anal 

lysis 

included 

in  analysis 

Slope 

Ground  cover    Tree  cover 

Common  cover 

1-39 

261 

-0.067 

0.620* 

0.231 

-0.382* 

1 

23 

0.010 

0.012 

0.325* 

-0.220 

2-39 

238 

-0.067 

0.613* 

0.187 

-0.354* 

1-2 

39 

0.106 

-0.116 

0.051 

0.069 

3-39 

222 

-0.089 

0.636* 

0.225 

-0.398* 

1-3 

54 

-0.009 

-0.089 

0.054 

0.013 

4-39 

207 

-0.064 

0.629* 

0.216 

-0.383* 

1-4 

74 

0.109 

0.053 

0.029 

0.043 

5-39 

187 

-0.100 

0.591* 

0.211 

-0.378* 

1-5 

92 

0.079 

0.407* 

0.134 

-0.250* 

6-39 

169 

-0.089 

0.484* 

0.164 

-0.271* 

1-6 

107 

0.025 

0.369* 

0.122 

-0.238* 

7-39 

154 

-0.071 

0.486* 

0.164 

-0.266* 

1-7 

127 

0.027 

0.316* 

0.246* 

-0.233 

8-39 

134 

-0.075 

0,503* 

0.104 

-0.265* 

•■With  <lf  =  -W   on  / 

of  It  0.23G  1 

is   sifinifirnnllv 

(liffcrcnl    f.nni  0 

.n    nt    p    - 

=    n.05. 

404 


GREAT   BASIN    NATURALIST 


Vol.  35,  No.  4 


appropriate!}'  on  slopes  and  in  areas  of 
low  vegetation.  Linsdale  (1946:  63)  has 
argued  that  the  disappearance  of  squirrels 
from  the  Hastings  Reservation  at  the 
termination  of  regular  grazing  was  caused 
by  the  visual  obstruction  of  the  taller 
grass.  In  our  area  the  grass  regularly 
reaches  heights  of  a  meter,  but  the 
squirrels  appear  to  compensate  for  this 
by  using  the  numerous  j^romontories 
(ground  cover)  for  visual  surveillance 
when  disturbances  occur,  as  well  as  when 
simply  lying,  sitting,  or  grooming.  A  sim- 
ilar factor  could  account  for  the  avoid- 
ance of  common  cover,  since  a  consider- 
able amount  of  the  ground  cover  beneath 
tree  canopies  was  made  up  of  small, 
highly  branched  logs,  sticks,  and  twigs. 
Some  of  the  ground  cover  was  beneath 
canopies  which  almost  reached  or  did 
reach  ground  level.  Such  cover  is  often 
neither  readily  mounted  nor  useful  for 
visual  surveillance  since  the  canopy  would 
often  obstruct  the  view.  The  same  factor 
might  also  explain  our  failure  to  find  a 
relationship  with  slope.  It  is  possible  that 
the  preferences  of  squirrels  in  flat  land  for 
embankments  is  related  more  to  enhanced 
visibility  than  to  drainage  (Hunt,  unpubl. 
data). 


Visibility  seems  an  unlikely  determi- 
nant of  the  preference  for  tree-canopy 
cover.  We  concur  with  Fitch's  (1948) 
suggestion  that  this  preference  is  related 
to  the  food  a^'ailable  in  the  trees,  i.e., 
acorns  in  his  case,  and  acorns  and  wal- 
nuts in  ours.  These  are  the  primary  food 
sources  in  the  fall  when  the  grass,  grass 
seeds,  and  forbs  eaten  in  the  spring  are 
no   longer   available    (Schitoskey    1973). 


References 

Carl,  E.  A.      1971.     Population  control  in  Arctic 

ground  squirrels.     Ecology  52:   395-413. 
Fitch,    H.    S.     1948.     Ecology    of    the   California 

ground    squirrel     on    grazing    lands.     Amer. 

Midi.   Nat.   39:    513-596. 
Hunt,    G.    L.      1974,     Habitat    selection    in    the 

'California      ground      squirrel.        Unpublished 

manuscript.     Department     of     Ecology     and 

Evolutionary    Biology,    Univ.    Calif..     Irvine. 
Linsdale,    J.    M.      1946.     The    California    ground 

squirrel.     Univ.     Calif.     Press.     Berkeley    and 

'Los   Angeles. 
Schitoskey,  F.      1973.     Energy  requirements  and 

diet  of  the  California  ground  squirrel.     Ph.D. 

Dissertation,   Ecology  Graduate  Group.   Univ. 

Calif.,    Davis. 
ToMicH,   P.   Q.      1962.     The   annual   cycle   of  the 

California     ground     squirrel.        Univ.     Calif. 

Publ.   Zool.   65:    213-282. 


ARACHNIDS  AS  ECOLOGICAL  INDICATORS 

Dorald  M.  AHred^ 

Abstract.--  In  tliree  ecological  studies  in  the  southern  parts  of  Nevada,  Idaho,  and  Utah,  spiders  of 
138  species,  scorpions  of  10,  solpugids  of  29,  and  phalangids  of  2  species  were  obtained  by  pitfall 
capture.  Significant  differences  in  species  occurrence  were  noted  between  tlie  three  geographic  and 
ecologically  distinct  areas.  Within  each  geographic  area,  arachnids  differed  in  species  occurrence  and 
abundance  in  different  plant  communities.  Only  three  s[)ecies  iif  spiders  and  four  species  of  scorpions 
were   common   to   all    three  geographic   areas. 


This  paper  summarizes  species  distri- 
bution of  arachnids  captured  in  can  pit 
traps  in  the  major  plant  communities  of 
three  geographic  areas — southern  Nevada, 
southern  Idaho,  and  southern  Utah — 
during  the  periods  of  1959  to  1966,  1966 
and  1967,  and  1971  to  1973,  respectively. 
Details  of  the  equipmetit  and  techniques 
used  and  specific  results  of  the  studies  in 
each  area  are  discussed  in  published  ar- 
ticles listed  in  the  references  section  of  this 
paper.  The  comparative  results  of  these 
three  studies  are  summarized  in  Tables 
1  to  4. 

Summary  Discussion 

Only  three  spiders — Calilena  restricta. 
Euryopis  scriptipes.  and  Haplodrassus 
eunis — were  common  to  all  three  areas. 
Fifteen  other  species  were  common  to 
Nevada  and  Utah,  seven  to  Nevada  and 
Idaho,  and  six  to  Idaho  and  Utah.  Paru- 
roctonus  boreus  was  the  only  scorpion 
found  in  all  three  study  areas.  Four  other 


species  were  common  to  Nevada  and 
Utah,  one  to  Nevada  and  Idaho,  and  one 
to  Idaho  and  Utah.  None  of  the  29  spe- 
cies of  solpugids  was  found  in  all  three 
areas.  Two  species  were  common  to 
Nevada  and  Utah,  and  two  to  Nevada 
and   Idaho. 

Apparently  the  habitat  ]:)references  of 
most  species  of  these  arachnids  are  so 
unique  that  even  at  similar  latitudes  and 
in  similar  vegetative  types  the  environ- 
mental factors  are  sufficiently  different 
to  influence  species  occurrence.  Such  dif- 
ferences in  geographic  distribution  may 
be  expected  more  with  groups  such  as 
scorpions  and  solpugids  whose  capability 
for  migration  is  extremely  limited,  where- 
as spiders  through  their  ballooning  adap- 
tations would  be  expected  to  be  more 
widely  distribtited. 

Within  each  geographic  area,  arachnids 
differ  in  species  and  abundance  in  rela- 
tionship to  the  different  plant  commun- 
ities. Where  similar  predatory  niches  are 


Table   1.   No.   of  collection   attempts   and  specimens  of  arachnids  collected  in  three  geographic  areas. 


Area                   Trap    periods^ 

Spiders 

Scorpions 

Solpugids 

Phalangids 

Nevada                     52,080 
Idaho                        20,490 
Utah                           3,590 

5,600 
500 
829 

1,471 
282 
198 

1,000 
72 
34 

575 
0 
0 

^No.   pit  trap>   x  no.   24-hr  periods 

operated 

Table  2.   Species   diversity 

of 

arachnids   in 

three  geographic  areas. 

No.    species 

collected 

Area 

Spiders 

Scorpions 

Solpugids 

Phalangids 

Total    species    collected 

Nevada 

Idaho 

Utah 

In  both  Nevada  &  Utah 

In  both  Nevada  &  Idaho 

In  both  Idaho  &  Utah 

Common  to  all  areas 

138 

94 

42 

40 

15 

7 

6 

3 

10 
9 
1 
5 
4 
1 
1 
1 

29 
28 
6 
3 
2 
2 
0 
0 

2 
2 
0 
0 
0 
0 
0 
0 

^Deparlnicnt    of    Zoolog}-.     Brjgliani     Youjig    University,     Prove.    Utah    S4602. 


405 


406 


GREAT   BASIN    NATURALIST 


Vol.  35,  No.  4 


Table  4.  Ecological  predominance  of  arachnid  species  in  three  geographic  areas. 


Most  Widespread 


Nevada 

Idaho 

Utah 

Nevada 
Utah 

Nevada 
Idaho 


Calilena  restricta 
Calilena  restricta 
Psilochorus  utahensis 

Vaejovis  confusus 
Paruroctonus  boreus 

Hemerotrecha  californica 
Eremobates  septentrionis 


Spiders 


Scorpions 


Solpugids 


Most  Abundant 


Psilochorus  utahensis 
Schizocosa  avida 
Psilochorus  utahensis 

Vaejovis  confusus 
Paruroctonus  boreus 

Hemerotrecha  californica 
Eremobates  septentrionis 


Table  3.   Arachnid   abundance   in   predominant 
plant   communities. 


Most   Species 


Fewest  Species 


Spiders 

Blackbrush  Juniper 

Sagebrush  Rabbitl)iusli 

Juniper  Blackbiusli 


Nevada 

Idaho 

Utah 


Nevada 
Utah 


Nevadc 
Idaho 


Scorpions 
Mixed  Juniper 

Blackbrush  Sagebrush 

Solpugids 
Hopsage  Blackbrush 

Rabbitbrush  Goosefoot 


demonstrated,  as  between  sjnders,  scor- 
pions and  solpugids,  the  occurrence  of  each 
group  would  be  expected  to  differ  within 
the  same  community.  For  example,  in 
Nevada  the  spiders  were  most  common 
in  the  blackbrush  and  mixed  communities, 
whereas  the  solpugids  were  least  common 
in  the  blackbrush  and  most  common  in 
the  pinyon- juniper  and  hopsage- wolf  berry 
where  the  spiders  and  scorpions  were  not 
abundant.  In  Idaho  the  spiders  were  most 
abundant  in  the  sagebrush  and  least  in  the 
rabbitbrush,  whereas  the  solpugids  were 
most  abundant  in  the  rabbitbrush.  In 
Utah  the  scor])ions  were  most  abundant 
in  the  blackbrush,  whereas  the  sjnders 
were  least  abundant  there. 

The  relationship  of  arachnid  species  to 
vegetative  type  is  not  a  direct  one,  but  an 
interrelationship  between  a  variety  of 
environmental-influencing  factors  such  as 
cover,  food,  moisture,  temperature,  and 
edaphic  conditions.  Specific  data  on  these 
influencing  factors  are  not  available  for 
correlation  with  the  results  summarized 
here.  Such  data  are  necessary  for  com- 
plete ecological  interpretations,  and  such 
correlative    studies    are    desirable    for    a 


better  understanding  of  these  interrela- 
tionships. However,  knowledge  of  species 
diversity  alone  in  different  geographic 
areas  as  well  as  predominant  plant  com- 
munities is  sufficient  to  determine  eco- 
logical indicators  that  may  be  monitored 
relative  to  changes  in  environmental 
quality. 

References 

Allred.  D.  M.      1965.  Note  of  phalangids  at  the 

Nevada  Test  Site.  Great  Basin  Nat.  25(1-2): 
37-8. 

.     1968.     Ticks  of    the     National    Reactor 


Testing  Station.  Brigham  Young  Univ.  Sci. 
'Bull.  Biol.   Ser.    10(1):  1-29. 

.  1969.  Spiders  of  the  National  Reac- 
tor Testing  Station.  Great  Basin  Nat.  29(2): 
105-108. 

.  1973a.  Scorpions  of  the  National  Reac- 
tor Testing  Station  Idaho.  Great  Basin  Nat. 
33(4): 25 1-254. 

.      1973b.     Effects   of   a    nuclear   detonation 

on  arthropods  at  the  Nevada  Test  Site.  Brig- 
ham  Young  Univ.  Sci.  Bull..  Biol.  Ser.  18(4): 
1-20. 

At.lred.  D.  M.,  .\ni)  D  E.  Beck.  1964.  Arthro 
pod  associates  of  plants  at  the  Nevada  Test 
Site.  Brigham  Young  Univ.  Sci.  Bull..  Biol. 
Ser.    5(2):  1-16. 

.      1967.     Spiders  of  the  Nevada  Test   Site. 

Great  Basin  Nat.   27(1):  11-25. 

Allred,  D.  M.,  D  E.  Beck,  and  C.  D.  Jorgen- 
SEN.  1963.  Biotic  communities  of  the  Ne- 
vada Test  Site.  Brigham  Young  Univ.  Sci. 
Bull.,  Biol.  Ser.  2(2):  1-52. 

Allred,  D.  M.,  and  W.  J.  Gertsch.  1976. 
Spiders  and  scorpions  from  northern  Arizona 
and  southern  TTtah.  Great  Basin  Nat.  (in 
press) . 

Allreu.  I).  M..  AND  M.  H.  Muma.  1971.  Sol- 
pugids of  the  National  Reactor  Testing  Sta- 
tion. Idaho.  Great   Basin  Nat.    ^1  (3) :  164-168. 

fli.RTscH.  W.  J..  .\ND  I).  M.  Allred.  1965.  Scor- 
jiions  of  tiic  Nevada  Test  Site.  Brigham 
Young  1  luv.   Sci.   Bull..  Biol.  Ser.   6(4):  1-15. 

Mil  ma,  M.  H.  1963.  Solpugida  of  the  Nevada 
Test  Site.  Brigham  Young  TTniv.  Sci.  Bull.. 
Biol.    Ser.    3(2):  1-15. 


NOTES  ON  THE  GENUS  BOM  BY  LI  US  LINNAEUS  IN  UTAH, 

WITH  KEY  AND  DESCRIIH  IONS  OE  NEW  SPECIES 

(DIPTERA:  BOMBYLIIDAE) 


D.   Elmer 


Lucilc    Maughaii    .lohiisoii' 


Abstract. —  A  key  to  the  Utah  species  of  Bombylius  is  piesented.  Bonibylius  hrxinuiculalux.  nigri- 
ventris.  monianus.  abdominalis.  aeslirus.  and  auriferoides.  new  species,  and  lancifer  kanabensis,  sub- 
species new.  are  described  from  Utah.  Bombylius  lassenensis.  new  name,  is  proposed  to  replace  the 
preoccupied   B.   pallescens  .Johnson   and   Maughan. 


The  first  comprehensi\  e  pajjer  on  Utah 
Bombyliidae  (Maughan  1935)  lists  six 
species  in  the  genus  Bombylius  Linnaeus, 
with  the  identity  of  a  seventh  left  in 
doubt  because  of  the  small  amount  and 
poor  condition  of  the  material  available 
for  study.  Additional  collecting  of  spec- 
imens in  good  condition  showed  this  spe- 
cies might  be  B.  clio  WilHston,  but  the 
specimens  are  not  well  enough  preserved 
to  include  in  our  present  key.  The  alter- 
nate species,  B.  fascialis  Cresson,  has 
been  collected  in  abundance  by  us.  These 
results  were  published  in  a  subsequent 
paper  (Maughan  and  Johnson  1936). 
Still  further  collecting  has  added  much  to 
our  knowledge  of  this  genus  as  it  occurs 
in  the  northern  Great  Basin  area  of  Utah, 
but  its  status  in  the  southern  Great  Basin 
and  in  the  Colorado  River  drainage  sys- 
tem is  still  little  known.  We  feel,  never- 
theless, that  publication  of  our  available 
information  at  this  time  is  worthwhile  as 
it  may  stimulate  future  collectors  to  study 
this  group  of  flies.  In  this  paper  we  dis- 
cuss briefl}'  the  forms  previously  known 
to  occur  in  the  state,  describe  seven  new 
taxa,  and  present  a  key  for  their  iden- 
tification. In  addition,  we  use  this  oppor- 
tunity to  give  a  new  name  to  the  homo- 
nym Bombylius  pallescens  Johnson  and 
Maughan. 

A  large  part  of  the  work  on  this  paper 
was  completed  eight  to  ten  years  ago. 
Unfortunately,  changing  circumstances 
and  the  illness  and  death  of  Elmer  John- 
son slowed  progress  on  its  completion.  We 
had  completed  the  key  and  most  of  the 
descriptions   prior   to  that  time.   In   May 


1975  I  (Lucile)  assumed  the  responsibil- 
ity, in  cooperation  with  Dr.  Vasco  M.  Tan- 
ner of  Brigham  Young  University  (BYU), 
of  completing  this  paper  so  it  could  be 
published.  I  have  completed  the  remaining 
species  descriptions  and  revised  the  paper 
where  it  seemed  advisable. 

The  holotypes,  allotypes,  and  ])aratypes 
dealt  with  in  this  paper  are  in  the  Brig- 
ham  Young  University  entomological  type 
collection. 

The  extensive  collection  of  bee  flies 
which  we  built  u])  over  the  past  40  years 
has  been  transferred  to  Brigham  Young 
University  where  it  has  been  carefully 
curated  by  Dr.  Tanner  and  his  associates 
and  is  available  for  study.  Many  of  the 
genera  represented  in  the  collection  in- 
clude apparently  undescribed  species 
which  will  hopefully  be  the  subject  for 
further  research.  Much  of  our  collection 
over  the  years  and  all  of  the  types  we 
have  named  in  other  genera  were  pre- 
viously deposited  in  the  Brigham  Young 
University  Life  Sciences  Museum. 

Nearly  all  of  the  material  dealt  with 
in  this  paper  is  from  Utah,  with  a  few 
specimens  from  neighboring  states,  large- 
ly collected  by  Mr.  Johnson,  some  by  both 
of  us,  and  a  few  by  other  collectors. 

Sincere  thanks  are  expressed  to  those 
who  have  aided  and  given  encouragement 
toward  the  completion  of  this  paper:  to 
our  family;  especially  to  Dr.  Tanner  for 
his  advice  and  active  assistance  in  caring 
for  our  collection;  and  to  Mrs.  Reginald  H. 
Painter,  who  has  been  a  source  of 
strength  and  inspiration. 


Key  to  the  Species  of  Bombylius  Discussed  in  this  Paper 

Wings  with  the  anterior  half  brown    or    blackish,    this    colored    area 

with  a  sinuous  and  sharply  defined  posterior  margin  major  Linneaus 


117    I.   Sircct.    Salt   Lake   City,    Utah   84103.    D.    Elmer  Johnson    died    W    Scptcmhcr    1973. 


407 


408  GREAT    BASIN    NATURALIST  Vol.    35,   No.   4 

Wings  with  brown  area  not  sharply     defined,     nearly     hyaline,     or 

spotted    -- 2 

2(1).        Wings  with  five  to  seven  distinct  dark  spots  on  veins  and  crossveins 

in  hyaline  portion 3 

Wings  at  most  faintly  spotted  on  veins   and    crossveins   other  than   the 

r-m  crossvein  and  free  part  of  vein  Cui  4 


3(2).  Wings  with  dark  spot  near  distal  end  of  vein  R.  .,;  heavily  tomentose, 
female  with  golden  tomentum  on   front;   usually   in   the   mountains 

or  foothills  albicapiUus  albicapillus   Loew 

Vein  Ro  ,  without  dark  spot  near  its  end;  sparsely  tomentose,  female 
with  pale  yellowish  tomentum  on  front;  usually  found  in  the 
foothills  and  valley  floors heximaculatus  n.   sp. 

4(2).         Face  prominent,  facial  pile  wholly  black  , 5 

Face  pollinose,  facial  pile  at  least  in  part  pale 11 

5(4).  Pile  of  dorsum  mostly  pale  yellow  to  golden;  legs  except  bases  of  fe- 
mora and  spots  on  knees  yellow;  female  with  large  trapezoidal 
area  on  front  shining;  both  sexes  with  median  line  of  pale  tomen- 
tum on  abdominal  dorsum;  halteres  pale  brown  to  yellow  

fascialis  Cresson 

Pile  of  dorsum  not  as  above;  usually  with  a  more  or  less  prominent 
line  of  brown  or  black  hairs  extending  from  base  of  wings  forward 
onto  occiput  --- 6 

6(5).        Males - .- 7 

Females    - -. 9 

7(6).  Pile  of  venter  black,  that  of  dorsum  pale  gray;  no  tufts  of  black  pile 
on  sides  of  abdomen  visible  from    above;    relatively    large    species 

(9-12  mm);   frequents  the   foothills nigriventris  n.  sp. 

Pile  of  at  least  first  three  ventral  abdominal  segments  pale;  tufts  of 

black  pile  on  sides  of  abdomen  visible  from  above 8 

8(7).  Pale  pile  of  dorsum  dark  yellowish  gray,  always  several  on  posterior 
callosities  black;  line  of  dark  hair  from  wing  onto  occiput  usually 
broad  and  prominent;  prominent  tufts  of  black  pile  on  hind  angles 
of  second  and  third  terga;  band  of  black  ]Hle  on  front  of  two  and 
often  on  three  also;  frequents  mountains  .-  .  lancifer  lancifer  Osten  Sacken 
Pale  pile  of  dorsum  nearly  white,  on  posterior  callosities  may  be  oc- 
casional black  hairs;  line  of  dark  hairs  from  base  of  wing  onto  occi- 
put thin  and  inconspicuous;  tufts  on  sides  of  terga  prominent  on 
anterior  angles  of  third  segment  only,  on  fourth  visible  from  ven- 
ter only - lancifer  kanahensis  n.  ssp. 

9(6).  Relatively  large  species  (9-12  mm);  lower  frontal  patch  almost  ob- 
solete, the  s])arse  scales  mostly  pale  brown;  ]iatches  of  dense 
white  or  nearly  white  tomentum  on  second  and  following  segments 
form  a  i)rominent  median  line  on  abdominal  dorsum;  tomentum 
of  remainder  of  dorsum  hairlike   and   relati\(4y    sparse;   frequents 

foothills    --  nigriventris  n.  sp. 

Smaller  species  (7-10  mm);  lower  frontal  patch  ]:»rominent,  of  black 
hairlike  scales;  abdomen  with  crossbands  of  more  or  less  dense  pale 
scales  on  posterior  half  of  second  and  following  segments,  median 
line  of  scales  evident  but  not  prominent    10 

10(9).  Pale  hairs  on  dorsum  mostly  dark  grayish  yellow,  many  black  hairs 
on  mesonotal  disc  and  posterior  callosities;  some  of  pale  tomentum 
of   abdominal   dorsum   noticeably     pale     brown;     dorsum     appears 


Dec.  1975  johnson.  johnson:   utah  bomhyliidae  409 

banded  black  and  pale  due  to  sparse    dark    tonientum    in    bands    of 

black  pile  on  anterior  part  of  terga  .   lancifer  laucifcr  Osten  Sacken 

Pale  hairs  of  dorsum  nearly  white,  few  if  an>"  black  hairs  on  nieso- 
notal  disc  and  posterior  callosities;  tomentuni  of  ])Osterior  half  of 
second  and  following  abdominal  segments  whitish  gray,  that  of  an- 
terior half  of  second  and  third  segments  tinged  with  tan,  covering 
segments  lancifer  kanabensis,  n.  ssp. 

11(4).      Lower  frontal  patch  white  to  grayish     12 

Lower  frontal  jjatch  yellow  to  golden     14 

12(11).  Facial  pile  wholly  white;  base  of  wings  brownish;  ])re-  and  postalar 
and  scutellar  bristles  prominent,  brown;  female  with  many  hea\  y 

bristles     on     upper     mesopleura  nionUinus  n.  sj). 

At  least  some  black  hairs  below  antennae  and  along  facial  orbits;  base 

of  wings  blackish 13 

13(12).  Most  of  pile  of  face  black,  only  that  on  oral  margin  yellowish;  entire 
frontal  triangle  of  male  covered  with  dense  appressed  white  scales; 
on  the  female  these  scales  are  found  only  between  antennae  and 
eyes;  dark  of  wings  fills  anteroproximal  one-third  or  more  of  wing; 
dark  spots  present  on  r-m  crossvein  and  free  jiart  of  vein  Cui  .... 

metopium  Osten   Sacken 

Most  of  pile  yellow  to  white,  only  a  narrow  band  of  hairs  below  an- 
tennae and  on  facial  orbits  black;  middle  of  frontal  triangle  of  male 
devoid  of  dense  appressed  tomentum;  only  anteroproximal  fifth  or 
less  of  wing  dark,  no  brown  clouds  on  veins  or  cross  veins;  prom- 
inent crossbands  of  black  pile  on  abdominal  dorsum 

abdominalis  n.   sp. 

14(11).    At  most  bases  of  femora  dark,  rest  of  legs  reddish  tan  15 

Legs  black 16 

15(14).  Pile  at  base  of  wings  golden;  dark  anterobasal  part  of  wing  with  dis- 
tinctly reddish  brown  cast;  male  genitalia  red,  slightly  darker  than 

femora;  late  summer  species   (August)   aestivus  n.  sp. 

Pile  at  base  of  wings  grayish  yellow  to  white,  male  genitalia  brown 

to  black 17 

16(14).  Abundant  erect  pile  of  front,  first  two  antennal  segments  and  wide 
band  below  lower  frontal  patch  black  and  very  long  in  both  sexes, 
female  with  mixed  pale  and  black  on  front;  pale  gray  species,  fe- 
male with  nearly  completely  hyaline  wings  lassenensis  new  name 

No  erect  pile  on  front  in  male;  pile  of  face,  appressed  hairlike  tomen- 
tum on  front  of  both  sexes,  and  erect  pile  of  female  all  same  pale 
color;  black  pile  on  antennae  and  narrow  band  below  lower  frontal 
patch    .  . aurifer  pendens  Cole 

17(15).  Halteres  yellow;  third  antennal  segment  of  female  widest  at  or  be- 
yond middle,  with  sides  nearly  parallel  for  three-fourths  of  its 
length,  then  tapering  abruptly  to  end;  pile  of  front  and  ocellar  tu- 
bercle  of   female   wholly   yellow;  dark  color  in  wing  of  male  does 

not  extend  beyond  tip  of  vein  Ri aurifer  aurifer  Osten  Sacken 

Halteres  various  shades  of  brown;  third  antennal  segment  of  female 
widest  before  the  middle,  not  evenly  tapering  toward  end;  pile  of 
front  and  ocellar  tubercle  of  female  mixed  black  and  yellow;  shin- 
ing wliite  scales  form  complete  band  above  antennae  between 
eyes;  dark  color  in  wing  of  male  more  or  less  fills  all  of  cell  Ri 
auriferoides  n.   sp. 


410 


GREAT   BASIN   NATURALIST 


Vol.  35,  No.  4 


Bombylius  heximaculatus,  ii.  sp. 

Wings  with  five  or  six  distinct  black 
spots  in  the  hyaline  area  or  bordering  the 
dark  color.  Legs  mostly  red.  9-12  mm  in 
length. 

Female. —  Black.  Head  gray  pollinose, 
occiput  densely  so;  lower  frontal  patch 
black,  very  sparse;  tomentum  of  front 
sparse,  golden;  pile  of  antennae,  ocellar 
tubercle,  front,  and  face  black,  a  few 
pale  hairs  around  the  oral  opening;  pile 
of  occiput  white  or  nearly  so,  a  patch  of 
brown  to  black  near  lower  hind  angle  of 
eye;  segments  1  and  2  of  antennae  about 
same  width,  2  one-third  as  long  as  1 ; 
segment  3  somewhat  wider,  about  one- 
fourth  longer  than  first  two  combined, 
widest  at  about  first  third,  sides  of  apical 
third  about  parallel;  proboscis  about  five- 
sixths  as  long  as  head  and  body  combined. 

Mesonotal  disc  velvety  black;  pile  long, 
dense  on  anterior  one-third  and  sides; 
sparse,  gray  mixed  with  a  little  black  on 
center  and  scutellum;  with  some  sparse 
decumbent  grayish  yellow  tomentum; 
pleura  densely  gray  pollinose,  lavender 
glints  in  some  lights;  a  dense  patch  of 
mixed  black,  brown,  and  gray  pile  on 
upper  mesopleura,  just  before  wings,  con- 
tinuous with  dark  hairs  of  lower  occiput; 
most  of  rest  of  pleura  naked  or  nearly  so, 
the  sparse  pile  mostly  pale,  a  few  black 
hairs    intermixed. 

Wings  with  anteroproximal  one-third 
smoky,  remainder  gray  hyaline;  dark 
spots  on  veins  at  r-m  crossvein,  base  of 
R,,  at  junction  of  Mj  +  R,-„  at  free  part  of 
M2,  at  cubital  end  of  median  crossvein, 
and  at  free  part  of  Cui.  Basicostal  pile 
black,  a  few  pale  hairs  on  sides  of  costa. 
Halteres  brown. 

Legs  dark  red,  hind  femora  yellowish 
except  knees,  others  brownish  red,  darker 
basally,  and  tarsi  dark;  pile  of  fore  coxae 
pale  yellowish  gray,  of  middle  coxae 
mixed  black  and  pale,  and  on  hind  ones 
only  a  few  black  in  the  gray;  of  rest  of 
legs  black;  bristles  black;  tomentum  pale, 
like  the  pile. 

Abdomen  faintly  gray  pollinose;  pile 
dense  on  sides,  lateral  fringe  mostly 
almost  white  on  sides  of  first,  anterior 
half  of  second,  fifth,  and  following  seg- 
ments, considerable  black  pile  mixed  in 
on  sides  of  posterior  half  of  second  and 
on  third  and  fourth;  pile  of  dorsum  most- 


ly short  jjale  yellowish  gray,  with  many 
long  black  hairs  on  hind  margins  of  sec- 
ond and  following  segments.  Most  of  pale 
pile  of  venter  black,  a  few  pale  hairs  on 
first  segTnent;  sparse  tomentum  pale 
yellow. 

Type. —  Female  holotype:  Alpine,  Utah 
Co.,  Utah,  20  May  1955'  (L.  M.  Johnson). 
Paratypes:  4  $    same  data. 

Several  males  were  collected  but  be- 
came damaged  so  no  description  will  be 
attempted  at  this  time.  The  males  are 
paler,  with  less  black  and  pale  pile  on  ab- 
dominal dorsum  and  more  black  pile  on 
sides.  Pile  of  venter  is  yellowish  gray. 
Females  are  more  tomentose.  There  is 
some  variation  in  both  sexes  in  the 
amount  of  dark  pile  at  hind  angles  of  eye 
and  on  mesopleura  between  the  wing  and 
head. 

Bombylius  nigriventris.  n.  sp. 

Black.  Anteroproximal  third  of  wings 
blackish,  gradually  evanescent  behind. 
Most  of  pile  pale  yellowish  gray,  that  of 
abdominal  venter  mostly  black.  Length 
6-12  mm  exclusive  of  proboscis.  Proboscis 
long,  three-fourths  as  long  as  to  fully  as 
long  as  body  and  head. 

Male. —  Head  black,  face  except  upper 
and  outer  margin  brown.  Front  and  dark 
margin  of  face  faintly  grayish  pollinose. 
Occiput  and  genae  yellowish  gray  polli- 
nose. Pile  of  face,  front,  antennae,  a  row 
along  u])per  orbits  on  occiput  and  ocellar 
tubercle  black.  That  of  rest  of  head  pale, 
faintly  yellowish  gray,  a  patch  of  bro\vn 
hair  on  lower  occiput.  Lower  frontal 
patch  of  short,  black  hairlike  scales.  An- 
tennae cylindrical,  segment  1  a  little  more 
than  twice  as  long  as  broad,  segment  2 
same  width,  about  as  long  as  broad,  seg- 
ment 3  somewhat  narrower,  slightly 
more  than  twice  as  long  as  first  two  com- 
bined, widest  at  its  proximal  one-fourth, 
sides  of  distal  two-thirds  almost  parallel; 
first  segment  long  pilose,  segment  2  very 
short  fine  pilose,  segment  3  with  a  few 
short  fine  hairs  on  its  u])per  side  on  distal 
half.  Palpi  black,  black  haired. 

Pile  long  and  dense  on  anterior  third, 
rolativel}-  short  and  sparse  on  remainder 
of  mesonotum,  very  pale  yellowish  gray, 
some  dark  hairs  scattered  among  pale  in 
center  of  mesonotal  disc.  Pile  of  pleura 
dense    above,    more    sparse    below,    lower 


Dec.  1975 


JOHNSON.   JOHNSON:    UTAH    HOMBYLHDAE 


411 


pteropleura  bare,  mostly  nearly  white, 
that  on  sterno-  and  hypopleura  slighth 
yellowish.  A  band  of  mixed  brown  and 
pale  hairs  extending  from  the  base  of 
wing  forward  to  join  with  patch  of  brown 
hairs  on  lower  occi])ut.  Pile  of  scutellum 
same  color,  occasional  darker  hairs  along 
hind  margin. 

Dark  of  wings  fills  alula,  basal  third 
of  axillary,  two-thirds  of  anal  cell,  base 
of  fourth  posterior  to  base  of  discal  past 
r-m  cross  vein  to  tip  of  subcostal  cell.  Re- 
mainder of  wing  gray  hyaline,  veins  dark 
brown.  Basicostal  pile  mostly  black,  longer 
and  pale  at  base,  a  few  pale  hairlike  scales 
beneath.  Halteres  brown. 

Coxae  and  base  of  femora  dark  brown, 
remainder  of  legs  reddish  brown;  bristles 
black;  pile  of  coxae  pale,  many  dark  hairs 
intermixed  on  middle  pair,  that  on  femora 
mixed  pale  and  dark  brown;  tomentum  of 
legs  pale  yellowish  gray,  a  line  of  brown 
scales  on  facies  of  femora. 

Abdomen  blue  black;  pile  of  dorsum 
long,  dense,  pale  faintly  yellowish  gray, 
lateral  fringe  on  second  and  third  and  a 
few  hairs  on  hind  margins  of  second  and 
following  segments  black,  somewhat  more 
numerous  on  fifth  and  sixth  terga.  Pile 
of  first  two  sternites  pale,  of  rest  of  venter 
and  on  genitalia  black,  a  very  few  pale 
yellowish    hairlike    scales    intermixed. 

Female.^ —  Less  pilose  and  more  tomen- 
tose  than  male.  Vertex,  front,  and  dark 
upper  and  outer  margin  of  face  brown 
pollinose.  Antennal  segment  1  about  two 
and  one-half  times  as  long  as  wide;  seg- 
ment 2  same  width,  as  long  as  wide;  seg- 
ment 3  widest  at  proximal  third,  thence 
tapering  evenly  to  short  onion-shaped 
style.  Tomentum  of  front  yellow,  a  few 
brownish  scales  near  orbit  in  lower  fron- 
tal patch;  that  of  occiput  nearly  white. 
Pile  of  head  as  in  male,  except  that  there 
are  a  few  pale  hairs  on  front  along  orbits. 

Mesonotum  and  scutellum  with  moder- 
ately dense,  pale  grayish  yellow  curly 
appressed  hairlike  tomentum.  The  band 
of  darker  pile  extending  from  base  of 
wing  to  occiput  is  yellowish  brown,  in- 
termixed with  a  few  black  hairs.  Wings 
paler  than  in  male,  basicosta  has  more 
pale  tomentum. 

Pile  of  abdomen  much  coarser  than 
in  male,  pale  hairs  mostly  more  yellow; 
many  more  black  hairs,  that  of  fifth  and 
sixth  terga   mostly  black;   black  hairs  of 


lateral  fringe  of  second  and  third  terga 
confined  to  hind  angles;  lateral  fringe  of 
fourth  and  fifth  segments  white. 

Dense  median  patches  of  faintly  yellow 
to  white  tomentum  on  second  and  follow- 
ing terga  form  a  prominent  line  down 
middle  of  abdominal  dorsum.  Pale  hairlike 
tomentum  on  hind  margins  of  first  two 
and  in  median  spots  on  following  ster- 
nites; rest  of  sparse  tomentum  of  venter 
black. 

Types. —  Male  holotype:  Alpine,  Utah 
Co.,  Utah  23-V-1955  (D.  E.  Johnson), 
female  allotype,  topotypical,  20-V-1955 
(D.  E.  Johnson),  paratypes  13cf  9,  topo- 
typical, ll-V-1954,  16-V-1954,  18-V-1954, 
22-V-1955,  19-VI-1955,  (D.  E.  and  L.  M. 
Johnson);  2cf  1  5  Dry  Canyon,  Salt  Lake 
Co.,  Utah;  Id  1  9  Fort  Douglas,  Salt 
Lake  Co.,  Utah,  lO-V-1933,  6-V-1940  (W. 
Anderson);  2^  Oak  Creek  Canyon,  Mil- 
lard Co.,  Utah,  24-V-1957  (D.  E.  John- 
son); 1  cT  Mt.  Carmel  Junction,  Kane  Co., 
Utah,  21-V-1957  (D.  E.  Johnson). 

In  addition  to  the  variation  in  size, 
there  is  some  variation  in  the  amount  of 
black  pile  on  the  mesonotal  disc  and  hind 
margins  of  the  abdominal  terga  in  the 
males,  and  in  the  amount  and  color  of 
tomentum  on  the  abdominal  dorsum  of 
the  females.  In  both  sexes  there  is  some 
variation  in  the  amount  and  color  of  the 
dark  pile  which  forms  the  conspicuous 
dark  line  from  the  base  of  the  wrings  for- 
ward onto  the  lower  occiput. 

This  species  has  the  same  type  of  face 
and  wings  as  lancifet\  but  can  readil}^ 
be  separated  from  that  species  by  its 
larger  size,  by  the  absence  in  both  sexes 
of  the  crossbands  of  black  pile  found  on 
the  hind  margins  of  the  second  and  third 
terga  of  lancifer,  by  the  presence  of  most- 
ly clark  vestiture  on  the  abdominal  venter, 
where  lancifer  is  mostly  pale,  and  by  the 
presence  in  the  female  of  the  prominent 
median  line  of  white  tomentum  on  the 
abdominal  dorsum.  Moreover,  lancifer  is 
a  species  of  the  mountains,  while  nigri- 
ventris  appears  to  be  confined  to  the  foot- 
hills and  valley  edges,  and  is  on  the  wing 
about  a  month  earlier  in  the  season. 

While  hovering,  the  females  of  this 
species  produce  a  sound  somewhat  lower 
in  pitch  than  the  hum  of  a  honeybee, 
and  of  a  different  quality,  being  more  of 
a  buzz  than  a  hum.  The  hovering  males 
produce  a  hum  considerably  higher  than 


412 


GREAT   BASIN   NATURALIST 


Vol.  35,  No.  4 


that  of  the  bee.  This  high-pitched  hum 
was  audible  to  us  for  a  distance  of  forty 
or  fifty  feet,  and  was  frequently  used  as 
a  means  of  locating  specimens.  Frequent- 
ly males  were  found  hovering  or  darting 
about  some  7  to  12  or  so  feet  above  a  given 
small  area.  We  found  our  first  female, 
as  well  as  several  subsequent  ones,  resting 
on  the  ground  beneath  such  a  hovering 
male.  We  did,  indeed,  use  the  males  as 
guides  to  the  females,  collecting  first  the 
female  and  then  the  male.  Another  spe- 
cies of  Bombylius  heximaculatus  n.  sp., 
was  present  in  the  same  habitat  as  nigri- 
ventris,  and  was  found  in  far  greater 
numbers.  There  was  sufficient  difference 
in  the  sounds  produced  by  the  two  species 
to  readily  recognize  and  locate  nigriven- 
tris  specimens  b}-^  the  lower  pitch. 

The  types  were  taken  about  one  mile 
west  of  the  village  of  Alpine,  at  an  eleva- 
tion of  5,340  feet.  The  site  is  a  narrow 
unplowed  strip  alongside  a  road  through 
agricultural  lands.  The  whole  area  is  foot- 
hills of  the  Wasatch  Mountains  above  the 
highest  level  of  Pleistocene  Lake  Bonne- 
ville. With  the  exception  of  Mt.  Carmel 
.function,  the  other  collection  sites  are 
also  foothill  areas  of  the  Wasatch,  at  ele- 
vations between  5,300  and  6,000  feet.  The 
elevation  at  Mt.  Carmel  Junction  is  listed 
as  5,241  feet. 

Although  other  species  were  available, 
the  only  flowers  at  which  we  saw  these 
beeflies  feeding  were  those  of  Phlox  longi- 
folia.  Wherever  we  have  found  nigriven- 
tris  flying  we  have  also  found  Phlox 
blooming. 

Bomhylius  lancifer  kanahensis.  n.  ssp. 

Like  lancifer  O.S.  except  it  has  far  few- 
er black  hairs  on  thorax  and  abdomen. 
Pale  pile  of  dorsum  glistens  in  reflected 
light,  making  the  fly  appear  to  shine. 
Length  8-10  mm. 

Male. —  Head  black,  face  shining 
brown,  face  and  front  from  some  angles 
lightly  brownish  gray  pollinose,  occiput 
and  chin  gray  i)ollinose;  pile  of  ocellar 
tubercle,  front,  face,  antennae,  and  a  few 
hairs  and  scales  in  a  patch  on  occiput  near 
lower  corner  of  eye  brown,  with  a  few- 
black  intermixed;  lower  frontal  patch 
black,  sparse  hairlikc  tomentum  of  front 
])ale  grayish  yellow;  pile  of  chin  white, 
of  occiput  grayish  yellow   (same  color  as 


that  of  adjoining  thorax).  Antennae 
slender;  segment  1  about  three  times  as 
long  as  wide;  segment  2  same  width  as 
first,  about  as  long  as  wide;  segment  3 
not  quite  as  wide  as  first  two,  its  sides 
nearly  parallel,  not  quite  twice  as  long 
as  first  two  combined,  its  style  evenly  long 
conical.  Proboscis  black,  about  four-fifths 
as  long  as  head  and  body  combined. 

Thorax  black,  bluish  reflections  in  some 
lights,  pleura  thinly  gray  pollinose  over 
shining  surface;  pile  fairly  long  and 
dense,  very  pale  yellowish  gray,  nearly 
white  on  pleura,  a  patch  of  dark  brown 
with  a  few  black  hairs  before  the  wing 
extending  onto  occiput;  macrochaetae  not 
strongly  developed,  nearly  same  color  as 
pile.  Two  black  hairs  on  left  posterior 
callus,  none  on  others. 

Wings  brownish  hyaline,  anteroproxi- 
mal  third  brownish,  color  evanescent  be- 
hind; veins  brown;  basicostal  pile  black, 
a  few  white  hairs  near  base  shading  to 
brown  along  heavy  vein;  pile  of  alulae 
long,  very  pale  yellowish.  Halteres  brown. 
Legs  yellow,  knees,  tarsi,  and  bases  of 
femora  brown,  first  two  pairs  broadly 
so;  pile  of  coxae  nearly  white,  of  remain- 
der of  legs  mostly  dark  brown  to  black; 
tomentum  nearly  white;   bristles   black. 

Abdomen  blue  black;  pile  of  dorsum 
long  and  dense,  faintly  yellowish  (paler 
than  thorax),  tufts  on  hind  angles  of 
second,  on  hind  angles  of  third  when 
viewed  from  below,  and  a  few  hairs  on 
hind  margins  of  second  and  succeeding 
.segments  black;  tomentum  nearly  white, 
a  sj^arse  jiatch  in  center  of  second  and 
fairly  dense  crossbands  on  ])osterior  third 
of  tiiird  to  fifth  segments,  covering  last 
segments.  Pile  of  venter  mostly  white,  a 
few  black  hairs  on  hind  margins  of  last 
three  segments;  sparse  tomentum  whitish. 
Genitalia  dull  black,  shading  to  brownish 
at  tips. 

Female. —  Much  like  the  male.  Less 
pilose,  but  much  more  tomentose  than 
male.  Front  brown  pollinose;  pile  black; 
tomentum  golden,  a  few  of  the  pale 
scales  mixed  into  the  lower  frontal  patch, 
especially  on  inner  end.  Face  with  a  few 
scattered  yellow  hairlike  scales  scattered 
among  pile.  Sparse  tomentum  of  occiput 
pale  yellowish,  a  few  black  hairs  mixed 
with  pale  buff  pile  near  vertex,  and  a  few 
black  and  dark  buff  hairs  near  lower 
inner   corner   of   eye.    Onl}^   a   few   black 


Dec.  1975 


JOHNSON,   JOHNSON:     UTAH    BOMBYLnDAE 


413 


hairs  in  dark  band  before  the  wing.  Pos- 
terior two-thirds  of  mesonotuni  and  scutel- 
lum  with  sparse  pale  yellow  curly  tomen- 
tum.  Wings  only  slightly  paler  than  in 
male.  Dark  ])art  of  femora  less  extensive 
than  in  male. 

Bristly  black  hairs  in  tufts  on  hind 
angles  of  second  and  third  terga  similar 
to  male,  relatively  more  bristly  black  hairs 
on  hind  margins  of  second  and  succeeding 
terga.  Second  sparsely  except  in  middle, 
and  succeeding  terga  densely  tomentose 
on  posterior  half  of  segments  and  in 
centers,  forming  a  distinct  nearly  white 
median  line;  anterior  half  of  segment  1 
almost  bare  of  tomentum,  front  of  3  and 
4  with  sparse  tomentum.  Most  of  pile  on 
first  three  ventral  segments  white,  on 
succeeding  segments  mostly  black;  tomen- 
tum pale  yellow  to  white. 

Types. —  Male  holotype  and  female 
allotype:  Six  miles  north  of  Kanab,  Kane 
Co.,  Utah,  5-VI-1965  (D.  E.  &  L.  M. 
Johnson).  Paratypes:  21  cJ",  12  9  same 
data. 

The  type  locality,  six  road  miles  north 
of  Kanab,  on  U.S.  Highway  89,  is  in  the 
red  sandstone  hills  of  the  Vermillion 
Cliffs  area.  The  flies  were  on  a  red  sandy 
sidehill  among  scattered  Juniperus  osteo- 
carpa  and  Artemesia  tridentata.  Asso- 
ciated with  this  species  were  males  of 
Lordotus  apicula  Coq.  The  males  were 
hovering  and  the  females  feeding  be- 
tween 8:30  and  10:00  a.m. 

This  series  of  flies  is  quite  uniform. 
There  is,  however,  some  variation  in  the 
amount  of  black  pile  on  the  thorax  and 
abdomen.  But  even  the  darkest  specimens 
have  very  few  black  hairs.  This  subspe- 
cies is  readily  distinguishable  from  the 
nominal  subspecies  by  this  scarcity  of 
black  pile,  and  by  the  paler  color  of  the 
pale  pile,  which  in  this  subspecies  is 
nearly  white.  On  females  there  may  be  a 
few  brown  to  black  hairs  on  occipital  or- 
bits near  the  vertex;  tomentum  of  front 
may  be  pale  yellow,  on  the  occiput  and 
the  thorax  in  front  of  the  wing  there  may 
be  no  dark  pile;  there  may  be  no  brown 
on  the  femora. 

Bombylius  major  Linnaeus 

This  is  the  earliest  bee  fly  on  the  wing 
in  the  spring  in  our  area..  It  appears  as 
early  as  mid-March  on  the  edges  of  the 


valley  floors  and  in  the  foothills.  We  have 
not  done  any  early  collecting  in  the 
southern  part  of  the  state  and  have  not 
seen  this  species  in  the  collections  of 
others  from  there.  We  have  seen  it  in  col- 
lections from  Arizona. 

Bornhjiius  nu'topiutri  Osten  Sacken 

This  small  species  appears  shortly  after 
major  around  the  edges  of  the  valleys  of 
the  northern  part  of  the  state.  As  the  seas- 
on progresses,  metopium^  like  se\eral 
others  of  its  congeners,  moves  up  into  the 
canyons  and  onto  the  foothills.  We  have 
found  it  common  in  only  a  few  places,  but 
it  is  so  inconspicuous  that  we  may  easily 
have  overlooked  it  at  other  sites  where  we 
were  not  specifically  seeking  it.  The  shin- 
ing white  tomentum  on  the  male  com- 
pletely obscures  the  front;  on  the  female 
it  is  found  only  at  the  sides  of  the  anten- 
nae. The  specimens  have  a  gray  appear- 
ance. 

Bombylius  lancifer  lancifer  Osten  Sacken 

We  have  collected  this  species  in  the 
mountains  abo^e  7,000  feet  elevation 
from  May  into  July.  We  have  found 
them  in  the  canyons  and  on  northern 
slopes  of  practically  every  mountain 
range  in  which  we  ha^e  sought  them  in 
the  northern  part  of  Utah.  We  do  not 
know  how  far  south  they  may  be  found. 
Specimens  have  been  examined  from 
Utah,  Oregon,  and  California. 

Bombylius  montanus.  n.  sp. 

Facial  pile  and  lower  frontal  patch 
wholly  white;  macrochaetae  well  devel- 
oped. Length  9-11   mm. 

Male. —  Head  dark  brown,  densely 
gray  pollinose,  face  and  oral  margin  yel- 
lowish; pile  white,  a  line  of  dark  brown 
to  black  hairs  on  each  side  of  bare  pol- 
linose area  on  frontal  triangle  between 
antennae  and  vertex,  some  on  ocellar  tu- 
bercle, and  upper  and  outer  side  of  first 
antennal  segment;  tomentum  of  occiput 
and  lower  frontal  patch  at  sides  above 
antennae  subshining  white,  a  few  brown 
decumbent  hairlike  scales  at  junction  of 
inner  orbits  on  upper  front.  Antennal 
segment  1  about  twice  as  long  as  wide; 
segment  2  about  as  wide  as  first,  as  long 
as  wide;  segment  3  about  as  wide  as  first 


414 


GREAT   BASIN    NATURALIST 


Vol.  35,  No.  4 


two,  about  twice  as  long  as  first  two  com- 
bined, widest  at  about  the  middle,  tapering 
evenly  thence  to  blunt  tip;  style  evenly 
conical.  Proboscis  brown,  about  two- 
thirds  as  long  as  body  and  head;  palpi 
brown,  pile  yellow. 

Thorax  brown,  dorsum  faintly  brown- 
ish, pleura  densely  gray  pollinose;  pile 
very  dense,  pale  gray  and  reddish  brown 
on  anterior  one-third  of  mesonotum, 
sparse  on  posterior  two-thirds  except  along 
sides;  posterior  callosities  have  dense  tufts 
of  pile,  reddish  at  base,  gray  at  ends, 
bordered  next  to  mesonotum  with  shorter 
almost  black  pile.  Sparse  pile  of  mesono- 
tum has  pale  gray  pile  with  hairlike 
gray  tomentum  beneath  it  in  two  indis- 
tinct longitudinal  stripes,  with  a  few  black 
hairs  intermixed  in  the  darker  contrasting 
brownish  pile  on  the  rest  of  the  disc.  Pile 
and  tomentum  of  scutellum  like  that  of 
posterior  mesonotum;  most  of  macrochae- 
tae  brown  to  black,  well  developed,  three 
or  four  on  notopleura,  a  number  on  pos- 
terior callosities  and  along  hind  margin 
of  scutellum.  Pleura  with  pile  moderately 
dense,  yellowish  gray  on  upper  half,  less 
dense  and  white  on  lower  half. 

Anteroproximal  half  of  wing  brown, 
somewhat  darker  along  veins,  the  color 
evanescent  behind,  becoming  gray  hya- 
line; veins  brown,  basicostal  pile  black,  a 
few  white  hairs  at  extreme  base  of  wings; 
pile  of  alulae  dense,  long,  white.  Halteres 
brown.  Legs  yellow,  knees  and  tarsi 
brown;  pile  and  tomentum  pale  yellowish 
to  white,  bristles  black. 

Abdomen  grayish  brown,  pollinosity 
almost  same  color;  pile  of  first  three 
terga  pale  brownish  yellow  on  sides, 
nearly  white  in  middle;  dark  brown  to 
black  bristly  hairs  form  single  rows  on 
hind  margins  on  outer  third  of  first  and 
across  second  and  third  segments  and 
form  prominent  tufts  on  sides  of  second 
segment,  with  small  tufts  on  sides  of  third; 
pile  of  rest  of  abdominal  dorsum  white, 
a  few  black  hairs  on  hind  margin  of  sixth 
segment;  tomentum  white,  sparse  on 
second  and  third,  moderately  dense  on 
succeeding  segments,  more  concentrated 
on  posterior  half  and  middle  of  second, 
third,  and  fourth;  pile  of  venter  mostly 
white,  a  few  black  hairs  on  terminal  three 
segments;  sparse  tomentum  pale  yellow. 

Genitalia  dark  brown  and  reddish,  pos- 
terior  angles   of   ninth   sternite   produced 


into  sharply  pointed  spinelike  projections; 
pile  yellow. 

Female. —  Much  like  male  but  less 
pilose  and  much  more  tomentose.  Wings 
almost  completely  hyaline.  Pile  of  occi- 
put yellowish,  that  of  front  sparse,  mixed 
black  and  white.  Front  and  face  densely 
pale  gray  pollinose.  Dark  pile  of  thorax 
paler  than  in  male.  Halteres  paler  than 
in  male.  Tomentum  on  abdomen  nearly 
white,  appearing  pale  brownish  on  mid- 
dles of  second,  third,  and  fourth  terga  be- 
cause less  dense,  rest  of  tomentum  dense; 
pile  mostly  white,  that  on  hind  margins 
of  segments  two  to  four  black  and  bristly, 
brown  to  pale  on  fifth  and  following  seg- 
ments, brown  tufts  on  sides  of  two  and 
three.  Pile  of  first  three  sterna  mostly 
white,  rest  mostly  black  and  brown,  bris- 
tly on  hind  margins  of  segments. 

Types. —  Male  holotvpe:  Aspen  Grove, 
Utah  Co.,  Utah,  19-VIU1953  (D.  E.  John- 
son) .  Female  allotype:  Pebble  Basin,  Ban- 
nock Co.,  Idaho,  24-VII-1948  (D.  E. 
Johnson).  Paratypes:  1  cT  same  data  as 
holotype;  1  d"  Strawberry  Valley,  Wa- 
satch Co.,  Utah,  1()-VII-1969  (on  flax  and 
cinquefoil). 

In  general  appearance  this  species  re- 
sembles eboreus  Painter  except  that  the 
dark  of  the  wings  is  distinctly  brown 
rather  than  blackish.  However,  the  strong 
development  of  the  macrochaetae  sets  this 
species  apart  from  all  of  its  congeners  in 
our  area.  We  have  seen  specimens  from 
Pasadena,  California,  which  may  be  this 
species,  but  more  material  will  be  neces- 
sary to  be  certain.  Apparent  variation 
shows  pile  of  posterior  callosities  all  pale 
gray.  Slight  rubbing  destroys  distinctive 
lines  on  mesonotum.  Macrochaetae  may  be 
pale. 

The  holotvpe  and  paratype  specimens 
were  collected  on  the  Mt.  Tim])anogos 
trail  about  a  half  mile  from  Aspen  Grove. 
There  were  three  males  and  one  female 
alternately  hovering  and  darting  about 
the  site,  but  only  two  of  the  males  could 
be  ca[)tured. 

Bomhylius  ahdotninalis.  n.  sp. 

Black,  pale  pile  of  dorsum  mostly  yel- 
lowish gray.  Length  9-12  mm  exclusive 
of  proboscis. 

Male. —  Head  black,  densely  gray  pol- 
linose,   oral    margin     vellow    shading    to 


Dec.  1975 


JOHNSON.   JOHNSON:     UTAH    BOMBYLHDAE 


415 


black  below  antennae.  Appressed  shining 
white  tomentuni  above  antennae  leaves 
center  pollinose  strip  bare  except  for  a 
ver}'  few  long  white  scales.  Black  erect 
pile  on  bare  area  next  to  white  tomentum. 
Several  rows  of  black  pile  below  antennae 
and  on  facial  orbits,  dense  yellowish  white 
pile  on  lower  face  and  oral  margin.  Tuft 
of  black  pile  on  ocellar  tubercle,  tomen- 
tum of  occiput  white,  pile  yellowish  gray 
abo^•e,  white  when  light  reflects  off  it. 
Antennae  with  first  two  segments  quite 
straight-sided,  slightly  wider  than  third; 
first  more  than  twice  as  long  as  second, 
third  about  one  and  one-half  times  the 
length  of  first  tw^o  combined.  Black  pile 
on  first  two  segments.  Proboscis  about 
three-fourths  the  length  of  body  and  head 
combined,  black,  as  are  palpi.  The  latter 
about  one  and  one-half  times  as  long  as 
the  width  of  the  base  of  the  proboscis, 
pile  black. 

Thorax  and  abdomen  black  with  pale 
pile  from  third  tergite  forward  3'ellowish 
gray,  shining  almost  white  in  some  lights, 
that  of  fourth  and  following  tergites 
white.  Dorsum  of  thorax  wdth  pile  dense 
at  sides  and  front  third  of  mesonotum  and 
on  scutellum;  shorter  on  rear  two-thirds, 
sparse,  with  a  few  black  hairs  intermixed, 
and  with  scattered  decumbent  fine  golden 
tomentum.  Macrochetae  yellowish.  Sparse 
black  pile  on  scutellum  and  post  allar  cal- 
losities. 

Wings  hyaline  except  anterobasal 
fourth  cloudy  reddish  brown,  evanescent 
behind.  Basicostal  pile  white,  with  black 
on  front  edge.  Pile  of  alulae  white,  dense 
and  long.  Halteres  brownish  with  part  of 
knobs  paler. 

Legs  have  black  femora,  tarsi  dark  yel- 
lowish brown.  Vestiture  white,  except 
spines  are  black. 

Dorsum  of  abdomen  with  pale  pile 
bushy,  black  pile  in  prominent  band  on 
rear  of  second  tergite,  narrower  band  on 
third  and  some  black  pile  on  rear  of  other 
segments.  All  pile  slightly  shorter  on 
sides  of  second  and  third  tergites.  All  to- 
mentum of  abdomen  is  white;  scant  on 
second  and  third  tergites,  more  dense  in 
center  of  each  segment,  giving  the  effect 
of  a  stripe,  wider  in  the  white  pile,  very 
narrow  in  the  black  pile.  Fourth  and  fol- 
lowing segments  with  dense  tomentum. 
Venter  of  second  and  third  segments  wjth 
long  white  pile   and  dense   long  hairlike 


tomentum,  following  segments  with  black 
and  white  pile  scant,  and  with  short  white 
tomentum  also  not  dense,  but  forming  a 
stripe  at  center  of  last  three  segments. 
Genitalia  very  dark,  yet  not  black  in  color. 

Female. —  Much  like  the  male.  Tufts 
of  black  pile  on  sides  of  wide  upper  front 
and  a  few  fine  golden  scales  on  front. 
Mesonotum  with  disc  back  of  long  pile 
covered  with  decumbent  fine  golden  to- 
mentum, more  dense  on  scutellum,  where 
it  is  arranged  with  the  ends  pointing  to- 
ward center.  Abdomnial  dorsum  much 
more  white  tomentose,  longitudinal  stripe 
more  than  twice  as  wide  as  on  male  and 
more  continuous,  with  diamond-shaped 
effect  on  second  and  third  segments.  Much 
less  white  pile  than  male  on  dorsum,  sides 
about  same.  Venter  less  pilose,  heavily 
tomentose  on  second  and  third  sternites, 
following  ones  with  short  tomentum 
forming  a  stripe  on  fourth  to  sixth,  cover- 
ing seventh.  Type  with  front  femora  dark, 
other  femora  partly  so,  and  tarsi  paler 
than  male. 

Types. —  Male  holotype  and  female  al- 
lotype: Stansbur}'  Mountain,  Tooele  Co., 
Utah,  23-V-1969  (D.  E.  Johnson).  Para- 
types:  236^,  39,  topotypical,  23-V-1967, 
18-V-1969,  23-V-1969;  8d,  Hickman 
Canyon,  Stansbury  Mountain,  Tooele  Co., 
Utah,  2-VI-1957;  6d,  H  ?  Clover  Creek, 
Tooele  Co.,  Utah,  9-VI-1957;  16cf ,  Alpine, 
Utah  Co.,  Utah,  18-V-1954,  24-V-1969 
(D.  E.  and  L.  M.  Johnson);  Id  Little 
Mt.,  Salt  Lake  Co.,  Utah,  17-VL1968. 
Other  specimens  have  been  collected  from 
Lark,  Salt  Lake  Co.,  Utah,  9-VL1953. 

In  addition  to  variations  in  size,  there 
may  be  little  black  pile  on  the  thorax; 
knobs  of  halteres  are  sometimes  mostly 
dark;  three  segment  stripe  of  white  to- 
mentum on  posterior  sternites  of  male 
faint  or  not  ])resent;  some  females  have 
completely  pale  yellowish  brown  femora. 

The  description  of  this  species  was  in- 
complete at  the  time  of  Mr.  Johnson's 
death.  lie  did  not  record  any  information 
about  the  ecology  of  the  Stansbury 
Mountains,  Hickman  Canyon,  or  Clover 
Creek  areas,  which  are  all  within  a  10- 
mile  distance  on  the  east  side  of  the  Ona- 
qui  Mountains.  However,  the  Alpine  spec- 
imens were  collected  in  the  same  strip  be- 
tw^een  cultivated  fields  as  nigriventris  in 
the  foothills  of  the  Wasatch  Mountains, 


416 


GREAT   BASIN    NATURALIST 


Vol.  35,  No.  4 


and  all  the  other  areas  are  wild  habitat 
at  similar  altitudes  and  with  similar  vege- 
tation. 

Bombylius  aestivus^  n.  sp. 

Similar  to  aurifer  O.S.  but  has  all  red 
femora,  and  pile  of  face,  front,  and  pleura 
pale  tawny  instead  of  nearly  white. 
Length  5-7%  nun. 

Male. —  Dark  brown;  yellowish  gray 
pollinose  on  front,  face,  first  and  second 
antennal  segments,  mesonotum,  pleura, 
first  tergum,  and  venter  of  abdomen. 
Antennae  black,  segment  1  about  twice 
as  long  as  wide;  segment  2  about  same 
width,  almost  round;  segment  3  almost 
twice  as  long  as  other  two  together,  not 
quite  as  wide,  almost  parallel  sided.  Pro- 
boscis black,  about  two  and  one-half  times 
as  long  as  head;  palpi  yellow  at  base, 
black  about  half  of  length,  somewhat  long- 
er than  width  of  proboscis. 

Pile  long  and  shaggy,  moderately  dense 
on  face,  occiput,  thorax,  and  sides  of  ab- 
domen; color  of  pile  shining  pale  tawny, 
somewhat  paler  on  pleura  and  abdominal 
venter,  with  black  pile  found  in  a  row 
below  lower  frontal  patch,  a  few  on  an- 
tennal segments  1  and  2,  and  sparse  tufts 
of  hairs  on  anterior  lateral  angles  of  third 
and  fourth  terga.  Tomentum  hairlike, 
shining,  of  about  the  same  color  as  pile, 
moderately  dense  on  thorax  and  abdomen, 
a  dense  appressed  patch  above  each  an- 
tenna.   Genitalia   pale   red. 

Legs  red,  knees  narrowly  black,  tarsi 
darkening  distally;  pile  and  tomentum 
as  on  remainder  of  body,  bristles  black. 

Wings  gray  hyaline,  brown  on  anterior 
proximal  one-third,  color  evanescent  be- 
hind and  distally;  pile  on  base  of  wings 
and  alulae  pale  tawny,  long  and  shaggy. 
Halteres  brown,  knobs  pale. 

Female. —  Very  much  like  the  male. 
Fewer  black  hairs  on  face  and  on  anterior 
angles  of  abdominal  segments  than  in 
male.  Abdomen  relatively  broader,  more 
densely  tomentose;  front  with  pile  and  to- 
mentum as  in  remainder  of  body. 

Types. —  Holotype  male:  Provo  Can- 
yon, Wasatch  Co.,  Utah,  14-Vin-1966  (D. 
E.  .Johnson).  Allotype  female:  Tryol  Lake, 
Uintah  Mts.,  Summit  Co.,  Utah  (no 
other    information,    but    collected    before 


1931).  Paratypes:  4cf,  1?  topotypical 
with  holotype. 

Some  damaged  specimens  in  the  col- 
lection show  some  variation  in  the  amount 
of  black  pile  on  the  face,  on  antennal  seg- 
ment 2  and  on  the  abdomen.  Some  speci- 
mens, particularly  females,  have  only  a 
very  few  black  hairs,  or  even  none  at  all 
in  these  places. 

This  species  may  be  readily  separated 
from  aurifer  aurifer  O.S.  which  may  occur 
in  the  same  area  by  the  red  femora  and 
by  the  darker  color  of  the  pile  of  the  face, 
occiput,  and  pleura,  which  in  aurifer 
aurifer  are  nearly  white.  Cole's  aurifer 
pendens  has  only  partially  red  femora. 

Whereas  the  other  species  of  Bombylius 
in  this  area  are  flies  of  the  spring  and  very 
early  summer,  this  species  is  apparently 
not  on  the  wing  until  much  later  in  the 
season  and  is  not  in  evidence  until  long 
after  the  others  have  disappeared  from 
even  the  high  country  that  seems  to  be 
home  for  aestivus. 

The  type  locality  is  in  the  Wasatch 
Momitains  about  one  mile  southeast  of 
the  Deer  Creek  Resenoir  in  Provo  Can- 
yon. The  flies  were  collected  on  a  north- 
facing  slope  in  open  areas  between  patches 
of  scrub  oak,  Quercus  gambeli^  and  choke 
cherries,  Prunus  melanocarpa.  They  were 
feeding  on  Helianthus  sp.  and  Aster  sp. 
The  elevation  is  between  7,000  and  8,000 
feet  above  sea  level.  The  allotype  has  only 
a  locality  label,  but  Tryol  Lake  is  a 
similar  ecological  habitat  in  the  Uintah 
Mts.  Other  species  of  bee  flies  common  in 
the  area  at  the  same  time  include  Poccil- 
anthrax  signatipennis  (Cole),  P.  sackenii 
(Coq.),  P.  willistoni  (Coq.),  Sparnopolius 
coloradensis  Grote,  Villa  edititoides  Paint- 
er, Aphoebantus  mormon  Melander,  and 
Lor  dolus  gibbus  Loew. 

Bombylius  lassenensis,  new  name 

At  the  time  Bombylius  pallescens  John- 
son and  Maughan  (1935)  was  described, 
we  were  unaware  that  Hesse  (1938:226) 
had  used  the  name  for  an  African  spe- 
cies. Since  our  species  is  a  junior  homo- 
nym of  Bombylius  pallescens  Hesse,  the 
new  name  Bombylius  lassenensis  is  pro- 
])osed  as  a  replacement.  This  species,  col- 
lected in  Lassen  Co.,  California,  is  in- 
cluded in  the  key  in  this  paper,  even 
fhouerh  it  is  not  from  Utah. 


Dec.  1975 


JOHNSON,   JOHNSON:     UTAH    BOMBYLHDAE 


417 


Bombylius  auriferoides,  n.  sp. 

Black,  pile  mostly  pale  yellowish  gray; 
similar  to  aurifer  0.  S.  but  with  brown 
halteres.  4.5-10  mm  in  length  exclusive 
of  proboscis. 

Male.^ —  Head  black,  gray  pollinose; 
pale  pile  faintly  yellowish  gray,  that  of 
ocellar  tubercle,  antennae,  and  outer  face 
mostly  black;  lower  frontal  patch  covered 
by  dense,  appressed,  shining,  nearly  white 
scales  which  nearly  completely  obscure 
the  front.  Antennal  segment  1  about 
twice  as  long  as  wide;  segment  2  same 
width  as  first,  about  as  long;  segment  3 
about  one  and  one-half  times  as  long  as 
first  two  combined,  about  as  wide  at  its 
widest  part  as  first  two,  widest  at  its  basal 
third;  pile  of  segment  2  mostly  pale,  that 
of  segment  1  black.  Proboscis  about  half 
as  long  as  head  and  body  combined. 

Thorax  faintly  grayish  pollinose  on  the 
mesonotal  disc,  densely  so  on  pleura; 
pile  long  and  dense,  that  on  pleura  nearly 
white;  short,  moderately  sparse  shining 
hairlike  scales  on  mesonotum  and  scutel- 
lum.  Wings  brown  on  anteroproximal 
half,  gray  hyaline  behind,  color  extends 
to  distal  end  of  cell  R,,  evanescent  behind; 
veins  brown,  darker  distally;  pile  of  squa- 
mae and  extreme  base  of  wing  pale,  rest 
of  pile  of  costa  black.  Halteres  brown. 
Legs  black,  dense  tomentum  and  most  of 
pile  pale,  a  few  dark  hairs  beneath  on  first 
and  second  femora;  bristles  black. 

Abdomen  moderately  densely  pilose, 
three  tufts  of  black  hairs  at  anterior  an- 
gles of  third,  fourth,  and  fifth  terga;  to- 
mentum of  abdomen  of  short,  curly,  ap- 
pressed hairlike  scales,  same  color  as  pile. 
Genitalia  brown,  members  yellowish  dis- 
tally. 

Female. —  Much  like  male,  somewhat 
less  pilose  and  more  tomentose.  Upper 
half  of  front  and  vertex  brownish  polli- 
nose, very  sparsely  tomentose;  lower  half 
of  front  gray  pollinose  and  densely  pale 
yellow  tomentose  in  an  uninterrupted 
band  between  eyes;  pile  of  ocellar  tu- 
bercle, front,  and  antennae  with  black 
and  pale  yellow  pile,  pale  pile  being 
mostly  on  posterior  part  of  tubercle, 
along  orbits,  and  on  upper  part  of  an- 
tennae. Wings  somewhat  paler  than  in 
male.  Fewer  black  hairs  at  sides  of  ab- 
domen. 

Types. —    Male    holotype    and    female 


allotype:  Mt.  Timpanogos,  Utah  Co., 
Utah,  elevation  8,600  ft.,  30-VI-1957 
(D.  E.  Johnson).  Paratypes:  Idaho:  Ban- 
nock Co.,  1  cf  Lava  Hot  Springs,  23-VL 
1935.  Utah:  IScT,  same  data  as  types; 
Utah  Co.,  Utah:  37cr,  18?  American 
Fork  Canyon;  IcT,  1  ?  Prove ;  2cf  Aspen 
Grove;  8cf,  2$  Pay  son  Canyon.  Salt 
Lake  Co.,  Utah:  1  J"  Lambs  Canyon; 
4cr,  2  $  Parley's  Canyon;  28 cf,  7  5  L'ttle 
Mountain.  All  sj)ecimens  collected  by  D. 
E.  .Johnson. 

In  addition  to  the  remarkable  range 
in  size,  there  is  considerable  variation  in 
some  other  characters.  The  color  of  the 
halteres  varies  from  rather  pale  to  Aery 
dark  brown,  with  the  females  usually 
being  somewhat  paler  than  males.  The 
color  of  pile  varies  from  nearly  white  to 
distinctly  yellowish,  particularly  on  the 
dorsum  of  some  s])ecimens.  The  smaller 
specimens  are  usually  paler  than  the  larg- 
er. The  very  early  season  flies  are  usually 
paler  in  all  respects  than  later  season 
specimens.  The  amount  of  black  pile  on 
the  sides  of  the  abdomen  varies  from  three 
distinct  tufts  in  some  specimens  to  only 
a  few  inconspicuous  black  hairs  in  others. 
Some  males  have  no  black  hairs  on  either 
the  ocellar  tubercle  or  antennae,  while 
others  have  a  few  on  either  or  both.  There 
is  variation  in  the  number  of  black  hairs 
on  the  antennae,  front,  and  ocellar  tuber- 
cle of  the  females;  but  always  there  are  at 
least  same  black  hairs  at  all  three  of  these 
locations,  and  always  there  are  some  pale 
hairs  along  the  orbits  on  the  lower  front. 
There  is  some  \ariation  in  the  relative 
length  of  the  proboscis.  The  third  anten- 
nal segment  varies  somewhat  in  shape  in 
drying,  but  always,  in  both  sexes,  there 
is  some  constriction  between  the  base  of 
the  segment  and  its  widest  point,  which 
is  nearly  always  at  the  basal  third.  Also, 
this  segment  is  always  narrowest  just  be- 
fore the  tip,  flaring  more  or  less  to  the 
end  where  the  onion-shaped  style  is  at- 
tached. 

This  species  and  aurifer  0.  S.  are  so 
similar  in  general  appearance  that  it  was 
not  until  we  were  studying  a  long  series 
of  specimens  of  what  we  assumed  were 
aurifer,  from  a  single  locality  to  determine 
the  intraspecific  variation,  that  we  re- 
alized we  were  dealing  with  more  than 
one  taxon,  and  that  the  small  differences 
we  had   noted   consistently  separated  the 


418 


GREAT   BASIN    NATURALIST 


Vol.  35,  No.  4 


whole  into  two  distinct  groups.  Inasmuch 
as  the  two  forms  have  sympatric  distri- 
bution they  can  only  be  considered  as 
distinct  species.  The  following  characters 
may  be  used  to  separate  them:  In  aurifer 
the  halteres  are  always  pale  yellow;  in 
auriferoides  they  are  some  shade  of 
brown;  the  pile  of  the  vertex  and  front 
of  the  females  of  aurifer  is  always  wholly 
yellow,  while  in  auriferoides  it  is  always 
mixed  black  and  yellow.  The  sides  of 
the  third  antennal  segment  of  aurifer  fe- 
males are  always  nearly  parallel  for  the 
proximal  half,  and  then  evenly  tapering 
to  the  end;  this  segment  in  auriferoides  is 
always  more  or  less  restricted  near  the 
base  and  before  the  tip,  and  the  widest 
part  is  usually  before  the  middle.  The  dark 
color  in  the  wing  of  the  males  of  aurifer 
does  not  extend  beyond  the  tip  of  vein 
Ri;  in  auriferoides  it  more  or  less  fills 
all    of    cell    Ri-    The    females    are    more 


readily   separated  than  the  males. 

Reexamination  of  the  material  listed  in 
the  earlier  paper  by  Maughan  (1935) 
shows  that  all  specimens  studied  are 
auriferoides  rather  than  aurifer.  The 
specimens  are  in  too  poor  condition  to 
include  as  paratypes.  We  have  found 
auriferoides  to  be  more  numerous  than 
aurifer  in  the  study  area. 

Literature  Cited 

Hesse.  A.  J.  1938.  A  revision  of  the  Boniby- 
liidae  (Diptera)  of  southern  Africa.  Ann.  So. 
Afr.   Mus.   34(1053):  332,   text   figures. 

Maugii.xn,  L.  1935.  A  systematical  and  morph- 
ological study  of  Utah  Bombyliidae  with 
notes  on  species  from  inteniiountain  states. 
.1.  Kansas  Ent.  Soc.  8(  1-2)  :27-80.  4  pis. 

Maughan,  L..  and  D.  E.  Johnson.  1936.  Notes 
on  Utah  Bombvliidae  (Diptera).  Proc.  Utah 
Acad.  Sci.,  Arts.  Lett.  13:197-201. 

.Johnson,  D.  E,  and  L.  Maughan.  1953.  Stu- 
dies in  Great  Basin  Bombvliidae.  Great  Basin 
Nat.    13  (1-2):  17-27. 


BREEDING  RANGE  EXPANSION  OF  THE  S  J  ARLING  IN  UTAH 

Dwight  G.   Sniitli' 

Abstract. —  The  discover^'  and  observation  of  colonies  of  starlings  nesting  in  the  eastern  Great  Basin 
desert  indicates  further  expansion  of  the  breeding  range  of  this  species  in  ITtah.  Data  on  nest  site  se- 
lection, nesting  productivity,  food  habits,  and  relationships   with    other    avian    s])ecies    are    presented. 


The  dispersal  and  establishment  of  star- 
Hngs  (Sturnis  vulgaris)  in  North  Ameri- 
ca has  been  well  documented  (Bent  1950; 
Davis  1960).  In  Utah  information  on  the 
initial  spread  of  this  species  was  sum- 
marized b}-  Behle  (1954).  While  study- 
ing raptor  populations  in  central  Utah 
from  1967  to  1972  I  observed  several 
small  colonies  of  starlings  nesting  in  des- 
ert habitats.  I  believe  these  colonies  show 
the  establishment  of  starlings  as  mem- 
bers of  the  breeding  avifauna  of  the  cen- 
tral Utah  Great  Basin  desert.  As  such, 
they  represent  a  breeding  range  expan- 
sion which  reflects  the  general  adapta- 
bility of  this  species. 

History  of  the  St.^rling  in  Utah 

The  range  expansion  of  starlings  in 
Utah  was  initially  gradual.  Starlings  were 
first  observed  in  the  state  in  February 
1939  near  Salt  Lake  City.  In  the  next 
year  small  winter  flocks  were  observed 
in  other  parts  of  Salt  Lake  Valley  and  a 
lone  individual  was  reported  near  Lehi, 
Utah  County.  In  January  1941  a  flock  of 
200  was  observed  at  Alt.  Carmel,  Kane 
County,  in  south  central  Utah  (Behle 
1958).  From  1941  to  1947  small  winter 
flocks  were  frequently  observed  near 
feedlots  and  ranches  in  Salt  Lake  Valle}^ 
In  1948  their  winter  range  again  expand- 
ed; a  flock  of  1,000  individuals  was  seen 
northward  in  Davis  County;  and  a  single 
individual  was  found  in  Kanab,  Kane 
County.  Their  winter  range  expansion 
continued  in  1950,  when  they  were  re- 
corded for  the  first  time  northward  in 
Box  Elder  County  and  westward  in 
Tooele  County.  During  the  decade  1950- 
1960  winter  flocks  of  starlings  increased 
tremendously.  Bailey  (1966)  reported 
flocks  as  large  as  100,000  and  noted  that 
they  constituted  an  important  agricultural 
pest     of     feedlots     and     orchards     in     16 

^Dcpl.     of    Biology.     Soullicrn    Coniicclicul    Stntc    C;ollegc,    Nc\ 


counties  of  the  state,  from  Washington 
Count}'  in  the  south  tf)  Box  Elder  Coimty 
in  the  north. 

The  first  nest  of  the  starling  in  Utah 
was  found  on  25  May  1949  in  an  old 
woodpecker  or  flicker  hole  on  the  west 
side  of  Salt  Lake  City.  In  the  following 
year  a  nest  was  discovered  in  a  shed  at 
Randolph,  Rich  County  (Behle  1954). 
Within  six  years  starlings  were  nesting 
at  many  localities  in  the  central,  popu- 
lated valleys  of  the  state.  In  1956  a  star- 
ling nest  containing  young  was  found  in 
a  shed  at  Lynn,  in  the  northwestern  cor- 
ner of  the  state  (Behle  1958).  Starlings 
have  since  spread  throughout  most  of 
the  state  and  now  are  a  sizable  com- 
ponent of  the  breeding  bird  populations 
of  the  towns,  settlements,  and  ranches 
of  the  northern  Great  Basin  area  (Hay- 
ward  1967).  During  my  study  of  raptor 
populations  in  Cedar  Valley,  LTtah 
County,  and  Rush  Valley,  Tooele  County, 
I  found  starlings  and  house  sparrows 
(Passer  domesticus)  to  be  the  most  com- 
mon breeding  birds  in  small  towns  such 
as  Fairfield  and  Cedar  Fort.  In  these  set- 
tlements they  typically  nested  in  a  variety 
of  buildings  and  in  holes  in  cottonwoods 
and  willows.  They  were  also  frequently 
observed  nesting  in  wind  breaks  border- 
ing agricultural  fields  and  pastures.  Here 
they  usuall}^  nested  in  holes  in  living  and 
dead  trees,  but  unused  bulky  stick  nests 
of  hawks  and  magpies  [Pica  pica)  were 
also  appropriated.  My  observations  in 
other  settled  areas  of  Tooele,  Juab,  and 
Millard  counties  showed  similar  choices 
of  nesting  site  selection  and  habitats  by 
starlings. 

I  belie^■e  that  my  observations  of  star- 
lings nesting  in  desert  habitats  warrant 
particular  interest  because  this  suggests 
invasion  and  adaptation  to  a  new  habitat 
as  well  as  a  further  range  extension  by 
this  species  in  Utah. 


419 


420 


GREAT   BASIN    NATURALIST 


Vol.  '35,  No.  4 


Nesting  Localities  and  Nest  Sites 

I  observed  nine  breeding  sites  located 
in  three  counties,  all  judged  to  be  ex- 
amples of  a  starling  breeding  range  ex- 
pansion into  habitats  of  the  northern 
Great  Basin  desert.  It  should,  however,  be 
mentioned  that  each  spring  and  summer 
I  frequently  observed  starlings  in  addition- 
al localities,  and  the  breeding  range  ex- 
pansion of  this  species  is  undoubtedly  oc- 
curring over  a  much  wider  area  than  my 
records  indicate.  Four  of  the  nine  sites 
supported  small  but  regular  breeding 
populations  during  the  six-year  study. 
The  other  sites  were  irregularly  active. 

Starlings  displayed  their  well-known 
adaptability  in  choosing  nest  sites  (Kes- 
sel  1957),  and  a  comparison  of  nesting 
habitats  and  nest  site  selection  indicates 
some  degree  of  opportunism.  Although 
Michael  (1971)  considered  his  observa- 
tions of  starlings  nesting  in  rocky  cliffs 
in  Kentucky  and  West  Virginia  to  be 
significant,  I  found  18  of  42  (42.9  per- 
cent) of  the  nests  I  actually  located  to  be 
placed  in  crevices  in  the  sheer  walls  of 
quarries  and  cliffs.  Of  the  remainder,  9 
(21.4  percent)  nests  were  located  in  some 
type  of  abandoned  mining  structure,  8 
(19  percent)  were  found  in  holes  in  juni- 
pers {Juniperus  osteosperma)  ^  and  7 
(16.7  percent)  in  stick  nests  of  hawks, 
owls,  and  ravens  (Corvus  corax).  The 
choice  of  nesting  sites  reflected  the  di- 
versity of  the  nesting  habitat.  Usually 
pairs  of  a  colony  were  found  nesting  in 
a  variety  of  sites  where  available. 

Several  sites  will  be  described  in  some 
detail  to  facilitate  comparison  with  ])os- 
sible   future   breeding  locations. 

Utah  County. —  Five  sites  were  found, 
all  in  the  western  section  of  the  county. 

A  colony  was  located  in  the  vicinity 
of  an  abandoned  mine  and  clay  pit  oper- 
ation at  Five  Mile  Pass,  which  is  about 
six  miles  west  of  Fairfield.  The  habitat 
at  this  locale  is  desert  scrub  intermixed 
with  widely  scattered  junipers.  Four  mine 
structures  are  still  standing  and  there  are 
six  large  quarries  and  clay  pits.  From  1967 
to  1972  a  total  of  17  nests  were  found  at 
this  site  as  follows:  2  in  1967;  4  in  1968: 
5  in  1969;  3  in  1970;  1  in  1971  and  2  in 
1972.  Several  additional  pairs  were  seen 
each  year  after  1968.  and  some  may  have 
been   nesting.   Of  the  nests   found,   seven 


were  placed  in  crevices  in  the  rock  walls 
of  clay  pits,  four  in  holes  in  junipers, 
three  in  the  walls  of  a  mine  shack,  two  in 
mine  bunkers,  and  one  in  the  beam  sup- 
ports   of   a   mine    tunnel. 

A  second  colony  active  during  all  six 
breeding  seasons  was  located  in  the  vi- 
cinity of  the  abandoned  Little  Topliff 
quarry  at  Ten  Mile  Pass.  This  site  was 
approximately  14  miles  southwest  of  Fair- 
field and  5  miles  northwest  of  Allan's 
Ranch.  The  habitat  of  this  site  is  a  mix- 
ture of  grasses  and  desert  scrub.  Fourteen 
nests  were  found  at  this  site:  2  in  1967, 
2  in  1968,  5  in  1969,  3  in  1970,  1  in  1971, 
and  1  in  1972.  Again,  additional  pairs 
were  observed  each  3'ear  after  1968.  Eight 
of  the  nests  were  located  in  creA'ices  in 
the  sheer  rock  face  of  the  quarry,  two  in 
an  unused  golden  eagle  {Aquila  chrysae- 
tos)  nest,  one  in  an  abandoned  prairie 
falcon  {Falco  mexicanus)  nest,  and  three 
in  cracks  in  a  wooden  retaining  wall. 

Another  Ten  Mile  Pass  site  was  located 
in  the  abandoned  Big  Topliff  quarry 
which  is  about  one  mile  east  of  Little 
lopliff.  This  quarry,  one  of  the  largest 
in  the  area,  is  bordered  entirely  by  desert 
scrub  communities.  Two  nests  were 
found  in  1970,  both  constructed  in  an  un- 
used golden  eagle  nest.  Several  individ- 
uals, some  of  which  may  have  been 
paired,  were  observed  in  1971. 

In  1969  a  third  colony  was  found  at 
Ten  Mile  Pass,  about  two  miles  east  of 
Big  Topliff  quarry  and  three  miles  north- 
west of  Allan's  Ranch.  Several  individuals 
and  two  nests  each  were  found  in  1969 
and  1970  in  a  large  limestone  cliff  line 
400  feet  above  the  valley  floor.  The  sur- 
rounding habitat  is  sparse  desert  scrub  de- 
void of  trees. 

Three  of  the  four  nests  were  located  in 
crevices  in  the  cliff  face.  The  fourth  was 
in  an  unused  red-tailed  hawk  {Butco  ja- 
maicensis)  nest  constructed  in  a  large 
crevice. 

A  fifth  colony  was  found  in  the  vicinity 
of  the  abandoned  Tintic  Empire  Mine, 
located  in  the  foothills  of  the  Boulter 
Mountains  approximately  four  miles 
north  of  Eureka  and  two  and  one-half 
miles  southeast  of  Allan's  Ranch.  The 
surrounding  habitat  of  this  colony  was  a 
large  stanrl  of  widely  spaced  junipers. 
Two  nests,  both  located  in  holes  in  juni- 
pers,   were    found    in    1969;    other    pairs 


Dec.  1975 


SMITH:    STARLING    IN    UTAH 


421 


were  present.  Onl}-  one  bird  was  seen 
when  this  site  was  rechecked  in  1970, 
and  none  were  observed  in  1971.  I  was 
unable  to  visit  this  site  during  the  1972 
breeding  season. 

Tooele  County. —  I  found  nests  at 
two  separate  sites  in  this  county  and 
evidence  of  nesting  activity  at  one  ad- 
ditional site. 

In  1968  and  again  in  1969  I  found  a 
starling  nest  in  a  hole  in  a  juniper  near 
the  entrance  to  Black  Rock  Canyon.  This 
nest  site  was  approximately  seven  iniles 
east  of  Vernon.  This  site  was  unusually 
interesting  because  it  was  located  in  a 
juniper  which  also  supported  an  active 
great  horned  owl  {Bubo  virginianus) 
nest  during  the  two  nesting  seasons.  Both 
starling  and  great  horned  owl  nesting 
attempts  were  successful  during  the  two 
nesting  seasons.  No  nest  was  present  in 
1970  or  1971,  but  starlings  were  observed 
in  May  and  June  in  the  same  juniper 
stand,  although  about  one-half  mile  above 
the  original  nesting  site. 

On  15  June  1969  I  observed  several 
starlings  in  a  juniper  stand  in  the  north- 
ern foothills  of  Simpson  Mountains.  This 
site  is  approximately  16  miles  northeast 
of  Simpson  Springs.  One  nest  was  found 
in  a  hole  in  a  juniper,  and,  judging  by 
the  behavior  of  the  other  birds,  additional 
nests    may    have    been    present. 

Juab  County. —  Murphy  et  al.  (1969) 
observed  12  starlings  near  a  golden  eagle 
nest  located  in  sandstone  cliffs  at  Yuba 
Dam  State  Park  and  presumed  them  to 
be  nesting. 

On  15  May  1970  a  pair  of  starlings 
was  observed  approximately  six  miles 
north  of  Trout  Creek  in  the  foothills  of 
the  Deep  Creek  Mountains.  One  carried 
nesting  material  and  was  seen  in  the  vi- 
cinity of  a  small  stand  of  junipers.  We 
were,  however,  unable  to  locate  a  nest. 


Reproductive  Chronology  and  Success 

Starlings  were  not  found  in  the  vi- 
cinity of  the  desert  nesting  sites  during 
the  fall  and  winter  months  (September 
through  February),  although  small  flocks 
were  frequently  observed  in  nearby  set- 
tlements. They  began  appearing  in  the 
future  nest  site  vicinity  in  early  March, 
and  the  majority  were  present  by  late 
March  and  early  April.  During  this  time 
they  were  frequently  seen  ins])ecting  di- 
lapidated mine  buildings  and  holes  in 
junipers  and  often  reacted  to  my  pres- 
ence near  these  sites  by  protesting  vigor- 
ously. 

Adults  carrying  nesting  material  were 
seen  in  late  March  and  throughout  April. 
Most  nests  were  constructed  of  grass,  pri- 
marily wheat  grass  (Agropyron  spica- 
tum),  and  frequently  lined  with  feathers. 
Four  of  the  nests  I  found  were  decorated 
with  juniper  greenery. 

Nests  containing  eggs  were  found  from 
late  April  through  mid- June.  Dates  of  the 
24  nests  with  eggs  which  we  found  are 
as  follows:  27,  30  April;  1,  2,  4(3),  10 
(2),  17(4),  19,  21(2),  30  May;  4,  9(3), 
and  19  June.  I  found  no  evidence  of  at- 
tempts to  raise  a  second  brood  in  July 
and  August. 

The  clutch  size  of  these  nests  averaged 
4.2  -  1.2  eggs  (range,  3-7;  mode,  4).  A 
clutch  size  comparison  with  other  areas 
is  presented  in  Table  1.  Utah  clutch  size 
averaged  significantly  smaller  than  those 
of  New  York  and  Holland  (t  =  3.95,  2.84 
respectively;  P<0.05  for  both)  but  not 
significantly  different  from  northwestern 
England    (t=1.5,   P>0.90). 

Young  were  in  the  nest  from  mid-May 
through  mid- July.  My  earliest  and  latest 
dates  for  nests  with  young  are  14  May 
and  28  July.  The  brood  size  of  17  nests 
was  5.9  ±1.1  young  (range,  2-7;  mode, 
4).  Interestingly,  there  was  no  significant 
difference    among    brood    sizes    of    Utah, 


Table  1.  Clutch  and  brood  size  comparison  of  central  Utah  nests.* 


Location                          No. 

No.  eggs 

No. 

No.  young 

Author 

clutches 

per  clutch 

broods 

per  brood 

Central    Utah                24 

4.5  ±  1.2 

17 

3.9  ±  1.1 

Present  Study 

Ithaca,  New  York       199 

5.5  ±  0.9 

230 

4.3  ±  1.3 

Kessell    (1957) 

Holland                         1592 

5.2  ±  1.0 

1377 

4.4  ±  1.3 

Kluijver    (1933) 

NW  England               105 

4.9  ±  1.1 

913 

4.2  ±  1.1 

Lack  (1948) 

•Data    is    average    ^    one    standard    deviation. 


422 


GREAT   BASIN    NATURALIST 


Vol.  35,  No.  4 


New  York,  Holland,  and  England  nests, 
indicating  perhaps  a  somewhat  higher 
overall  hatching  success  of  Utah  nests. 

Overall  reproductive  success  was  high. 
Of  13  nests  on  which  I  was  able  to  obtain 
complete  information,  12  successful!}' 
hatched  young;  and  of  these,  11  nests 
fledged  young.  Two  nests  were  aban- 
doned; one  containing  three  eggs  and 
one  with  five  young.  Neither  jDair  at- 
tempted to  renest.  Overall  hatching  and 
fledging  success  was  94.2  percent  and 
84.6  percent,  respectively.  Both  percent- 
ages are  slightl}"^  higher  than  reported 
from  previous  studies  in  other  areas. 

Foraging  and  Food  Habits 

Information  on  foraging  and  food  habits 
is  limited  and  w^as  obtained  from  morning 
observation  of  three  nests,  two  in  1969 
and  one  in  1970,  all  located  in  the  Five 
Mile  Pass  nesting  colony.  Adults  were 
observed  from  a  parked  vehicle  with  a 
40X  spotting  scope  attached  to  a  window 
mount.  Only  those  food  items  brought 
to  the  nest  site  which  could  be  identified 
are  included  in  the  results  presented  in 
Table  2. 

Adults  foraged  predominantly  in  the 
sagebrush-wheatgrass  {Artemisia- Agropy- 
ron)  associations  which  were  the  common 
plant  communities  in  the  nesting  site  vi- 
cinities. They  spent  considerably  less  time 
in  the  ground  layer  vegetation  of  pinyon- 
juniper  (Pinus-Juniperus)  communities 
and  among  the  rubble-strewn  floor  of 
quarries. 

Over  86  percent  of  the  arthropod  food 
items  brought  to  the  nest  were  insects. 
Of  these,  Orthoptera  comprised  56  per- 
cent and  Colcoptera  almost  27  percent. 
Araenids  were  the  only  other  animal  food 
which  was  taken  in  significant  quantities. 
In   a   food   habit   study   in   eastern   Texas 


Table   2.     Arthropod   food    of   starlings    in    the 
eastern  Great  Basin. 


Item 

No.  indv. 

%  Freq. 

Acrididae 

31 

47.0 

Tettigoniidae 

6 

9.1 

Carabidap 

11 

16.7 

Tenebrionidae 

3 

k") 

Scarabidac 

3 

4.5 

Cicadellidae 

2 

10 

Formicidae 

1 

1.5 

Araneidae 

9 

13.6 

Totals 

66 

99.9 

based  on  stomach  contents,  Russell  (1967) 
found  Orthoptera  and  Coleoptera,  particu- 
larly Carabidae,  to  comprise  84  percent 
of  the  total  insects  eaten  and  68  percent 
of  the  total  food,  with  other  arthropods 
and  some  plant  material  constituting  the 
remainder  of  the  diet.  I  did  not  identify 
any  utilization  of  plant  material  for  food, 
but  results  are  undoubtedly  biased  be- 
cause the  small  nesting  populations  pre- 
cluded collection  of  adults  and  young  for 
stomach  contents  analysis.  Both  Killpack 
and  Crittenden  (1952)' and  Bailey  (1966) 
noted  the  extensive  use  of  such  plant  ma- 
terials as  grain  and  corn  silage  by  win- 
tering flocks  of  starlings.  Starlings  are 
undoubtedly  opportunistic  in  their  feed- 
ing habits  and  utilize  the  most  available 
food.  This  is  reinforced  by  a  comparison 
of  the  food  habits  of  these  desert  nesting 
starlings  with  the  result  of  Fautin's 
(1946)  investigations  of  the  invertebrate 
populations  of  the  sagebrush  community. 
Analysis  reveals  that,  with  the  exception 
of  Formicidae,  starlings  utilized  the  most 
prevalent  ground  layer  invertebrates  in 
the   sagebrush    community. 

Relations  with  Other  Species 

Starlings  appear  to  be  the  predominant 
avian  species  in  the  vicinity  of  their 
breeding  locales.  Other  birds  observed  in 
the  same  locale  included  the  house  spar- 
row, pinyon  jays  {Gyrnnorhinus  cyano- 
cephala)^  scrub  jays  {Aphelocoma  coe- 
rulescens),  mourning  doves  (Zenaidura 
macroura),  common  nighthawks  (Chor- 
deiles  minor),  and  mountain  bluebirds 
(Sialia  currucoides) .  Only  house  spar- 
rows were  common  nesting  associates,  and 
at  two  sites.  Big  and  Little  Topliff  quar- 
ries, they  outnumbered  the  starlings.  At 
these  and  other  sites  the  two  species  ap- 
peared to  mutuall}'  tolerate  one  another 
and  no  aggressive  interactions  were  ob- 
served. We  did  observe  starlings  inter- 
acting aggressively  with  scrub  jays 
(twice)  and  mourning  doves  (twice) 
which  had  perched  in  the  immediate  vi- 
cinity of  the  nesting  site.  In  each  in- 
stance the  starlings  displaced  the  intrud- 
ing birds.  The  only  direct  evidence  of 
nest  displacement  which  we  obsened 
concerned  a  mountain  bluebird  nest 
which  contained  six  eggs  when  discovered 
on  15  May  1970.  When  this  nest  was  re- 
checked    five    days    later    we    found    an 


Dec.  1975 


SMITH:    STARLING   IN    UTAH 


423 


adult  starling  sitting  on  a  clutch  of  six 
eggs.  No  trace  of  the  mountain  bluebird 
was  found,  and  the  adults  were  not  ob- 
served  again   in   the   nesting   territory. 

In  a  previous  study  of  the  food  habits 
of  raptors  breeding  in  the  eastern  Great 
Basin  Desert,  starlings  were  occasionally 
recorded  as  prey  of  several  hawk  and 
owl  species  (Smith  and  Murphy  1973). 
It  is  possible  that  their  plumage,  size,  and 
aggressive  habits  make  them  conspicuous 
targets  and  hence  more  liable  to  be  preyed 
upon  by  raptors. 

Discussion 

Within  the  relatively  short  time  span 
of  30  years  the  starling  has  become  a  pre- 
dominant component  of  Utah's  avian 
fauna.  Its  successful  invasion  and  estab- 
lishment can  be  roughly  delineated  into 
three  segments,  each  of  approximately 
10  years  duration.  In  the  first  10-year 
period  (1939-1948)  following  their  ap- 
pearance in  the  state,  starlings  were  ob- 
served only  as  individuals  or  small  flocks 
of  winter  visitants.  In  the  next  10  years 
(1949-1958)  these  winter  flocks  increased 
tremendously  in  size  and  greatly  ex])and- 
ed  their  winter  range  to  include  most  of 
the  populated  central  j)ortions  of  the  state. 
The  first  nesting  attempts  were  reported 
early  in  this  period  and  by  the  end  of  the 
decade  small  nesting  populations  were 
found  in  many  widely  separated  towns 
and  settlements  of  the  state.  In  the  third 
10-year  period  (1959-1968)  both  winter 
and  summer  starling  populations  had  in- 
creased in  size  to  the  point  at  which  the 
starling  had  achio\ed  the  status  of  a 
major  pest  species.  During  this  time  the 
starling  became  (with  the  possible  ex- 
ception of  the  house  sparrow  in  some 
areas)  the  most  abundant  bird  throughout 
most  of  the  settled  portions  of  the  state. 
Although  most  common  in  cities  and 
towns,  they  were  also  found  in  widely 
separated  settlements  and  ranches.  In 
winter  they  formed  large  flocks  which, 
through  their  feeding  and  roosting  ac- 
tivities, became  an  economic  nuisance  to 
feedlot  owners  and  fruit  growers  through- 
out the  state. 

At  the  present  time,  so  far  as  is  known, 
starlings  nest  in  all  the  settlements  and 
towns  in  the  state  and  have  recently  ex- 
tended their  breeding  range  into  desert 
habitats.      Although      initiallj^      sporadic. 


their  occupation  of  distinctly  desert  habi- 
tats for  nesting  pur])oses  has  recently  be- 
come more  widespread,  suggesting  that 
they  have  been  able  to  adapt  to  a  new 
habitat  type. 

In  analyzing  the  range  expansion  and 
establishment  of  starlings  in  Utah  I  con- 
sider the  following  to  be  of  significance: 
(1)  mobility,  (2)  suitable  climate,  (3) 
suitable  habitat,  and  (4)  sufficient  popula- 
tion })ressure.  The  four  factors  are,  in  fact, 
a  measure  of  the  starling's  adaptability 
and,  when  considered  together,  explain 
the  success  of  this  species.  Although  not 
specifically  investigated  in  this  study, 
their  rapid  range  expansion  across  North 
America  indicates  that  starlings  have 
adequate  powers  of  mobility  and  wide 
climatic  tolerances.  While  largely  non- 
migratory,  their  behavorial  adaptation  of 
forming  large,  mobile  winter  flocks  which 
break  up  and  disperse  to  fa\orable  nesting 
areas  in  spring  undoubtedly  allows  some 
exploratory  activity  which  may  in  turn 
lead  to  further  range  expansion. 

Water,  but  not  food,  may  pose  the  only 
j)otential  limiting  factor  for  starling  range 
expansion  into  the  Great  Basin  desert. 
Starlings  at  their  desert  breeding  sites 
were  observed  drinking  water  from  ephe- 
meral rain  puddles  and  from  liA'estock 
watering  troughs. 

The  adaptability  of  starlings  to  new 
habitats  is  well  known  (Kessell  1957). 
A  review  of  the  pattern  of  starling  in- 
vasion in  Utah  suggests  that  a  habitat 
with  some  form  of  human  modification 
provides  a  favorable  impetus  for  range  ex- 
pansion. Thus,  widely  separated  towns 
and  settlements  were  occupied  within  20 
years,  but  the  interAening  areas  of  desert 
were  not  invaded  until  after  starling 
populations  were  well  established  in  near- 
Ijy  settlements.  Only  after  starling  popula- 
tions had  occupied  these  locales  did  a 
further  range  expansion  into  the  upper 
Great  Basin  desert  take  place.  It  is  quite 
])ossible  that  the  human  habitats  pro- 
vided breeding  sites,  probabl}'  due  to  the 
well-known  breeding  behavior  adaj)ta- 
tions  of  this  species  with  reference  to 
man,  which  resulted  in  local  population 
increases.  Pressures  resulting  from  these 
local  population  increases  may  have  en- 
couraged starlings  to  examine  the  adja- 
cent habitats  of  the  upper  Great  Basin 
desert. 


424 


GREA'l'    BASIN    NATURALIST 


Vol.  35,  No.  4 


The  short  time  span  ol  their  successful 
utilization  of  this  new  habitat  suggests 
behavorial  adaptations  rather  than  genetic 
changes  within  the  population.  However, 
the  possibility  of  future  evolutionary 
changes  in  populations  occui)ying  such 
habitats  may  warrant  further  study. 

Literature  Cited 

Bailey.  E.  P.  1966.  Abuiidanre  and  activity 
of  starlings  in  winter  in  northern  lUah.  Con- 
dor  68:152-162. 

Behi.e,  W.  H.  1954.  Changing  status  of  tiie 
starling   in    Utah.    Condor    56:49-50. 

.      1958.     The     birds     of     tlie     Raft     River 

Mountains,  northwestern  Utah.  Univ.  Utah 
•Biol.    Ser.    11(6):  1-49. 

Bent,  A.  C.  1950.  Life  histories  of  North 
American  wagtails,  shrikes,  vireos  and  their 
allies.    U.S.    Natl.    Mus.    Bull.    197:182-214. 

D.wis.  D.  E.  1960.  Comments  on  the  migration 
of  starlings  in  eastern  United  States.  Bird- 
Banding   31:216-219. 

F.AUTiN,  R.  W.  1946.  Biotic  communities  of  the 
northern  desert  shrub  biome  in  Western 
Utah.  Ecol.  Monogr.   16:251-310. 


H.\YW.-\RD,  C.  L,  1967.  Birds  of  tlie  Upper  Colo- 
rado Rivei-  Basin.  Brigham  Young  Univ.  Sci. 
Bull..    Biol.    Ser.    9(2):  1-64. 

Kessei.i.,  B.  1957.  A  study  of  the  breeding  biolo- 
gy of  the  European  starling  (Slurnis  vul- 
garis) in  North  America.  American  Midi. 
Nat.    58:257-331. 

Kii.i.PACK,  M.  C,  AND  D.  N.  Crittenden.  1952. 
Starlings  as  winter  residents  in  the  Uinta 
Basin,   Utah.   Condor  54:338-344. 

Kluijver,  I.  H.  N.  1933.  Bijdrage  tot  die  bio- 
logic en  do  ecologie  van  den  spreeuw  (Stur- 
nus  vulgaris  vulgaris  L.)  gedurende  zijn 
voortplantingstiid.  Versl.  Meded.  Planten- 
ziekt    (Wageningen)    69:1-145. 

Lack.  D.  1948.  Natural  selection  and  family 
size   in    the   stalling.    Evolution   2:95-110. 

Michaei,.  E.  D.  1971.  Starlings  nesting  in 
rocky  cliffs.   Bird-Banding  42:123. 

Murphy,  J.  R.,  F.  J.  C.mviinzand,  D.  G.  Smith, 
and  ,I.  B.  Weston.  1969.  Nesting  ecology 
of  raptorial  birds  in  central  Utah.  Brigham 
Young   Univ.    Sci.    Bull.,    Biol.    Ser.    10:1-36. 

RussELi,.  D.  N.  1971.  Food  habits  of  the  star- 
ling   in   eastern   Texas.   Condor    73:369-372. 

Smith,  D.  G.,  and  J.  R.  Murphy.  1973.  Breed- 
ing ecology  of  raptors  in  the  eastern  Great 
Basin  of  Ihdh.  Brigham  Young  Univ.  Sci. 
Bull..    Biol.    Ser.    18(3):  1-76. 


SOME  PARASITES  OF  PADDLEFISII  (POLYODON  SPAT  HULA) 
FROM  THE  YEELOWSTONE  RIVER,  MONTANA 


Lawrence  I^.   Lockai 


nd   R.   Randall   Parsons- 


Abstract. —  One  species  of  copepod  (Ergasilus  elongatus),  one  tiematode  (Dichholhriuiu  liaiuu- 
laturn),  two  nematodes  (Camallanus  oxrcephalus  am\  Contracaecum  sp.).  and  two  cestodos  (Marsi- 
pomctra  hastata  and  M.  pnrva)  were  recovered  from  17  i)addlefisli  [Polyodun  spalhula)  collected 
from  the  Yellowstone  River.  Montana,  on  11  and  18  May  1971  The  male  paddlefish  averaged  60.4 
nematodes  and  157.6  cestodes  per  infected  fish  while  the  females  averaged  12.0  nematodes  and 
415.7  cestodes.  The  higher  intensity  of  cestode  infection  in  female  paddlefish  was  attributed  to 
their  larger  size  and  consequent  greater  intake  of  food  resulting  in  more  exposure  to  the  cestode 
intermediate     hosts     (Cyclops    bicuspidalus) . 

The  gastrointestinal  tract  was  dissected, 
and  the  contents  were  washed  onto  a 
2()()-niesh  screen,  then  transferred  to  an 
illuminated  tray  (Barber  and  Lockard 
1973)  for  examination.  Cestodes  were 
fixed  in  AFA  (alcohol-formalin-acetic 
acid)  and  stained  with  Delafield's  hema- 
toxylin. Nematodes  were  placed  in  a  mix- 
ture of  70  percent  alcohol  and  5  percent 
glycerine  and  later  mounted  in  glycerine. 
The  liver,  heart,  gall  bladder,  spleen,  and 
gills  were  dissected  and  each  was  placed 
in  a  jar  wdth  water  and  agitated  on  a 
mechanical  shaker  for  5-10  minutes. 
The  contents  were  poured  onto  a  200- 
mesh  screen,  washed,  transferred  to  an 
illuminated  tray,  and  examined.  Trema- 
todes  and  copepods  were  placed  in  a  mix- 
ture of  70  percent  alcohol  and  5  per- 
cent glycerine.  The  ovaries,  testes,  and 
gas  bladder  were  examined  grossly  and 
observed  abnormalities  checked  micro- 
scopicall}'. 

Results 


There  have  been  relatively  few  reports 
on  the  parasites  of  paddlefish  (Polyodon 
spat  hula).  Linton  (1898)  found  ta])e- 
worms  in  paddlefish  from  the  Ohio  River, 
Ohio.  Stockard  (1907)  mentions  large 
numbers  of  cestodes  in  paddlefish  from 
the  state  of  Mississippi;  and  Wilson 
(1914)  found  parasitic  copepods  on 
paddlefish  from  the  Mississippi  River  in 
Illinois  and  Iowa.  Simer  (1929  and  1930) 
and  Beaver  and  Simer  (1940)  examined 
171  paddlefish  from  the  Tallahatchie 
River,  Mississippi,  and  reported  on  tre- 
matodes  and  cestodes  from  them.  Bang- 
ham  and  Vernard  (1942)  found  one  pad- 
dlefish from  Reelfoot  Lake,  Tennessee, 
infected  with  trematodes,  nematodes,  and 
cestodes.  Meyer  (1946)  discovered  leech- 
es parasitizing  paddlefish  while  Causey 
(1957)  examined  one  paddlefish  and 
found  parasitic  copepods.  Meyer  (1960) 
studied  cestodes  from  paddlefish  collected 
from  the  Mississippi  River  in  Iowa  and 
the  Missouri  River  in  South  Dakota. 
Hugghins  (1972)  examined  one  paddle- 
fish from  Fort  Randall  Resenoir  on  the 
Missouri  River  in  South  Dakota  and  found 
it  parasitized  with  tapeworms  and  nema- 
todes. Weisel  (1973)  reported  uniden- 
tifted  tapeworms  and  nematodes  from  3 
paddlefish  from  the  Yellowstone  River, 
Montana.  The  present  study  was  under- 
taken in  an  effort  to  gain  information  on 
the  parasites  of  paddlefish  from  Montana. 

Methods 

This  report  was  based  upon  the  necrop- 
sy of  17  paddlefish  collected  from  the 
spawning  run  in  the  Yellowstone  River 
near  Intake,  Montana,  on  11  and  18  May 
1973.  The  fish  were  eviscerated,  and  the 
viscera  and  gills  were  placed  in  plastic 
bags    and    frozen    for    later    examination. 


The  ten  male  paddlefish  had  an  av- 
erage weight  of  10.6  kilograms  and  range 
of  6.8  to  13.6  kilograms,  while  the  seven 
females  had  an  average  weight  of  22.8 
kilograms  and  range  of  20.0  to  27.2  kilo- 
grams. 

Table  1  lists  the  rates  of  infection  with 
copepods,  trematodes,  nematodes,  and 
cestodes  recovered  from  paddlefish  during 
this  study.  The  parasitic  copepod  Ergasilus 
elongatus  was  identified  by  Dr.  Z.  Kabata 
of  the  Fisheries  Research  Board  of  Can- 
ada. The  trematode  Diclybothrium  ham- 
ulaturn  was  identified  by  Dr.  Fred  P. 
Meyer  of  the  Bureau  of  Sport  Fisheries 
and  Wildlife.  Identifications  of  the  nema- 
todes Camallanus  oxycephalus  and  Con- 
tracaecum sp.  w^ere  confirmed  by  Dr.  E.  J. 


J312    S.    Grant    Ave..    Picirc.    SD 
-Department    of    Biology.     Montana 


State     Universily.     Bozcmnn.    Montana 


425 


426 


GREAT   BASIN    NATURALIST 


Vol.  35,  No.  4 


Table  1.  Incidence  of  parasites  of  Pohdou  spathula  collected  from  the  Yellowstone  River.  Montana. 
Percent  infected  Mean   no.    of   parasites    (range) 

Parasite Male       Female  Male  Female 

Ergasilus   elongatus 
Diclybothrium    hamulatum 
Carnal  lanus    oxycephalus 
Contracaecum  sp. 
Nematode  cysts 
Marsipometra   hastala 
M.  parva 


30 

14 

100 

100 

80 

86 

80 

86 

100 

100 

100 

100 

90 

100 

69.4    (12-302)  12.0    (4-25) 

157.6    (34-356)  415.7    (37-1013) 


Hugghins  of  South  Dakota  State  Univer- 
sity and  Dr.  G.  L.  Hoffman  of  the  United 
States  Fish  and  Wildlife  Service,  respec- 
tively. Spherical  cysts  (1-3  mm  in  diam- 
eter) containing  larval  nematodes  were 
present  on  the  surface  of  the  stomach, 
pyloric  caecum,  and  the  intestine  of  all 
fish  examined.  All  17  paddlefish  ex- 
amined were  infected  by  tapeworms 
identified  from  Beaver  and  Simer  (1940) 
and  Meyer  (1960)  as  Marsipometra  has- 
tata    and    M.    parva. 

Discussion 

It  was  felt  that  the  handling  procedures 
greatly  reduced  chances  of  discovering 
parasitic  copepods  as  well  as  monogenetic 
flukes  which  may  have  been  present  on 
the  gills  of  the  fish.  Causey  (1957)  says 
statements  of  incidence  of  infection  for 
copepods  have  little  value.  Thus,  the  in- 
cidence of  infection  of  copepods  found  in 
Table  1  may  not  be  indicative  of  the  num- 
ber of  infected  fish  actually  present.  Like- 
wise, quantitative  analysis  on  the  pres- 
ence of  the  monogenetic  fluke  would  be 
of  little  value. 

Differences  in  intensity  of  nematode 
infections  (Table  1 )  between  male  and 
female  paddlefish  were  noted  but  were 
difficult  to  interpret  because  of  the  pres- 
ence of  immature  Contracaecum  sp.  in 
the  intestine. 

Simer  (1930)  and  Beaver  and  Simer 
(1940)  name  three  species  of  cestodes  in- 
fecting paddlefish:  Marsipometra  hastata, 
M.  parva,  and  M.  confusa.  Meyer  (1960) 
examined  tapeworms  from  these  studies 
and  his  own  material  and  concluded  that 
M.  hastata  and  yi.  confusa  were  synony- 
mous. He  also  found  Cyclops  hicuspidatus 
to  be  the  intermediate  host  of  M.  hastata. 
In  this  study  no  attempt  was  made  to 
distinguish  M.  confusa  from  .M.  hastata. 
Females  had  a  higher  intensity  of  tape- 
worm infection  than  male  paddlefish, 
and  this  may  be  attributable  to  larger  size 
of   the    females.    Females    weighed    more 


than  twice  as  much  as  males  and  thus 
would  have  ingested  more  of  the  inter- 
mediate hosts  of  these  parasites  while 
feeding. 

Acknowledgment 

The  authors  express  their  appreciation 
to  Jeffrey  Bagdanov  for  assisting  with  the 
necropsy  of  the  paddlefish. 

Literature  Cited 

B.^NGH.^M.  R.  V.,  .and  C.  E.  Vernard.  1942. 
Studies  on  parasites  of  Reelfoot  Lake  fish. 
IV.  Distribution  studies  and  checklist  of 
parasites.  J.  Tcnn.  Acad.   Sci.   17(1):    22-38. 

Barber.  D.  L.,  and  L.  L.  Lockard.  1973.  Some 
helminths  from  mink  in  southwestern  Mon- 
tana, with  a  checklist  of  their  internal  para- 
sites. Great  Basin  Nat.  33(1):   51-60. 

Beaver,  P.,  and  P.  H.  Simer.  1940.  A  restudy 
of  the  three  existing  species  of  the  cestode 
genus  Marsipometra  Cooper  (Amphicoty- 
lidae)  from  the  Spoonbill.  Polyodon  spathula 
(Wal.),   Tr.   Am.    Micr.    Soc.    59:    167-182. 

Causey,  D.  1957.  Parasitic  Copepoda  from 
Louisiana  freshwater  fish.  Am.  Midi.  Nat. 
58(2):    378-382. 

HuGGiNs.  E.  I.  1972.  Parasites  of  fishes  in 
South  Dakota.  South  Dakota  Exper.  Sta.  Bull. 
484:    1-73. 

Linton,  E.  1898.  Notes  on  cestode  parasites  of 
fishes.  Proc.  U.S.  Nat.  Museum  20:   423-456. 

Meyer.  F.  P.  1960.  Life  history  of  Marsipo- 
metra hastala  and  the  biology  of  its  host. 
Polyodon  spathula.  Iowa  State  ITniv.  Library, 
Ames,    Iowa    (unpubl.). 

Meyer,  M.  C.  1946.  Further  notes  on  the 
leeches  (Piscicolidae)  living  on  freshwater 
fishes  of  North  America.  Tr.  Am.  Micr.  Soc. 
65(3):    237-249. 

Simer.  P.  H.  1929.  Fish  trematodes  from  the 
lower  Tallahatchie  River.  Am.  Midi.  Nat. 
11(12):   563-588. 

Simer,  P.  H.  1930.  A  preliminary  study  of  the 
cestodes  of  the  spoonbill.  Polyodon  spathula 
(Wal.).  Tr.  111.  State  Acad.  Sci.  22:    139-145. 

SiocKARD.  C.  R.  1907.  Observations  on  the  nat- 
ural history  of  Polyodon  spathula.  Amer. 
Nat.  41:   753-766. 

Weisei..  G.  E.  1973.  Anatomy  and  histology 
of  the  digestive  system  of  the  paddlefish 
{Polyodon  spathula)  J.  Morph.  140(2): 
243-251. 

Wii.soN,  C.  B.  1914.  Copepod  parasites  of  fresh- 
water fislies  and  theii-  economic  relations  to 
mussel  glochidia.  Bull.  F.S.  Bur  Fish. 
34(1914):    331-374. 


REPRODUCTIVE  CYCLE  OF  THE  BELDINC;  GROUND  SQUIRREL 

{SPERMOPHILUS  BELDINGl  BELDINGI)  -. 

SEASONAL  AND  AGE  DIFFERENCES 

Martin  L.   Morton^   and  John   S.   Gallup- 

Abstract. —  The  reproductive  cycle  in  Belding  ground  squirrels  was  studied  in  the  Sierra  Nevada 
Mountains  at  two  locations,   one   at  2,100   ni   elevation,  the  other  at  3,000  m. 

Adults  emerged  from  hibernation  completely  prepared  physiologically  for  reproduction.  Males 
tended  to  emerge  slightly  ahead  of  females  and  yearlings  tended  to  emerge  later  than  adults.  Year- 
ling females  were  fertile  but  produced  smaller  litters  than  adults,  4.48  vs.  6.31.  Yearling  males  were 
infertile.  They  exhibited  a  slight  seasonal  cycle  in  testicular  growth  but  did  not  reach  sexual  maturity. 
Testicular  growth  and  spermatogenesis  were  incipient  in  many  adults  and  in  yearlings  prior  to  hiber- 
nation. 


Hibernation  and  seasonal  breeding  are 
important  survival  strategems  of  rodents 
living  at  high  latitude  or  high  altitude. 
Typically  in  these  environments  there  are 
extreme  seasonal  oscillations  in  ambient 
conditions.  Winters  tend  to  be  long  and 
cold  and  summers  brief  and  sharply  de- 
limited. Dormancy  is  employed  as  a 
means  of  bridging  the  long  gap  of  energy 
shortage  in  winter,  and  breeding  is  co- 
incident with  the  clement  weather  and 
abundant  food  of  surmner.  Both  respon- 
ses require  advance  preparation  and  ac- 
curate timing  to  be  maximally  adaptive. 
The  physiology  of  hibernation  is  cur- 
rently a  viable,  active  field  of  study, 
whereas  seasonal  breeding  has  aroused 
less  interest  and  its  complexities,  especial- 
ly in  wild  populations,  are  poorly  under- 
stood  (Chapman   1972). 

Herein  we  report  on  seasonal  changes 
in  reproductive  functions  of  the  Belding 
ground  squirrel  {Spertnophilus  beldingi 
beldingi),  a  hibernator  that  lives  at  high 
altitude  in  the  Sierra  Nevada  Mountains 
of  California. 

Methods 

This  study  extended  from  1969  to  1973 
and  was  on  S.  b.  beldingi  living  principal- 
ly in  meadows  of  Lee  Vining  Canyon, 
Mono  County,  California.  A  number  of 
squirrels  were  live-trapped  or  shot  at  Big 
Bend  (elevation  ca.  2,100  m),  but  most 
data  are  from  those  living  near  Tioga 
Pass  (elevation  ca.  3,000  m).  At  both 
areas  we  carried  on  an  extensive  mark- 
release  program  throughout  the  time 
squirrels  were  active  above  ground.  This 
program  enabled  us  (1)  to  follow  seasonal 


changes  in  body  weights  and  dimensions 
and  external  appearance  in  individuals 
of  known  age  and  (2)  to  collect  animals 
of  known   age  for   specimens. 

Live-trapping  was  conducted  with 
Tomahawk  wire  mesh  traps  baited  with 
peanut  butter.  A  few  specimens  were  col- 
lected with  a  .22  caliber  rifle.  Animals 
trapped  for  the  first  time  were  toe-clipped 
in  a  standard  pattern  never  involving 
more  than  one  toe  per  foot.  All  animals 
were  examined  externall}^  for  appearance 
of  vulva  and  mammae  or  of  scrotal  pig- 
mentation and  position  of  testes. 

Freshly  excised  reproductive  organs 
were  fixed  in  Bouin's  solution  and  trans- 
ferred to  70  percent  ethanol.  At  the  time 
of  transfer  they  were  debrided,  blotted, 
and  weighed  to  the  nearest  0.1  mg.  Tis- 
sues were  sectioned  at  7  or  1 0/t  and  stained 
with  hematoxylin  and  eosin.  Measure- 
ments of  seminiferous  tubules  were  taken 
with   an   ocular  micrometer. 

Results 

Arousal  schedule.^ —  The  sequence 
and  pace  of  events  in  the  active  season 
were  alike  at  the  two  study  areas,  but 
the  active  season  at  Big  Bend  usually  be- 
gan at  least  six  weeks  in  advance  of  that 
at  Tioga  Pass.  Snowcover  was  not  com- 
parable at  the  two  sites  at  the  beginning 
of  the  season.  At  Big  Bend  most  burrow 
sites  were  clear  of  snow  when  emergence 
occurred.  At  Tioga  Pass  emergence  tended 
to  begin  on  knolls  that  were  the  first 
areas  to  become  snow  free,  but  many  in- 
dividuals tunneled  out  at  sites  covered  by 
snow  up  to  a  depth  of  2  m.  Similar  ef- 


^Biology    Department,    Occidental    College,    Los   Angeles,    California   90041. 
-Alaska    Department    of   Fish    and    Game,    Homer,    Alaska. 


427 


428 


GREAT   BASIN    NATURALIST 


Vol.  35,  No.  4 


fects  of  snowpack  on  emergence  of  S. 
columbianus  were  noted  by  Shaw  (1925). 
There  was  considerable  annual  variation 
in  snow  conditions  and  the  schedule  of 
S.  b.  beldingi  was  affected  accordingly 
(see   beyond). 

Adult  males  were  the  first  animals  seen 
above  ground  at  a  given  location,  but  a 
few  adult  females  and  an  occasional  year- 
ling could  be  found  within  a  few  days 
thereafter. 

Testes. —  At  emergence  adult  males 
had  scrotal  testes  weighing  about  2  to  3 
g,  the  maximum  weight  seen  during  the 
entire  active  season  (Fig.  1).  The  scrotum 
was  darkly  pigmented.  Within  a  month 
after  emergence  testicular  weight  of 
adults  began  to  decrease  noticeably  and 
testes  had  become  inguinal  or  abdominal 
in  position  and  scrotal  pigmentation  was 
decreasing.  Six  weeks  after  emergence  tes- 
ticular weight  had  decreased  to  a  seasonal 
minimimi  that  was  maintained  thereafter 
at  Big  Bend  until  onset  of  hibernation 
(Fig.  1,  upper).  At  Tioga  Pass,  however, 
considerable  increase  in  testicular  weight 
of  adults  occurred  during  the  last  few 
weeks  of  the  season  (Fig.  1,  lower). 


' 

' 

■ 

Big     Bend 

. 

■ 

• 

•.1 

: 

t 

: 

° 

"          o             I 

I 

r 

I 

.> 

.     :»•  *°    8 

• 

^11 ' 


"I    .8  °  + 


Fig.  1.  Paired  testes  weights  of  Spermophilus 
beldingi  beldingi  throughout  tlie  active  season  at 
two  study  areas.  Note  log  scale  on  ordinate. 
Cross  on  lower  right  corner  of  Tioga  Pass  data 
indicates  mean  testicular  weight  of  juveniles  prior 
to  hibernation. 


In  histological  perspective,  testes  of 
adults  showed  intense  spermatogenic  ac- 
tivity from  time  of  emergence  through 
onset  of  weight  collapse.  During  this  time 
seminiferous  tubules  were  of  large  diam- 
eter (150  to  250/i)  with  spermatozoa  fill- 
ing the  liunina.  As  testicular  weight  de- 
creased spermatogenesis  ceased  and  tu- 
bule diameter  decreased.  During  June  at 
Tioga  Pass,  for  example,  diameters  went 
from  about  150  to  60/j..  Beyond  June, 
lumina  in  seminiferous  tubules  were  ab- 
sent. Recrudescing  testes  of  adults  col- 
lected in  late  August  at  Tioga  Pass  had 
a  thickening  germinal  layer  with  num- 
erous primary  spermatocytes  and  a  few 
secondary  spermatocytes  present.  Semi- 
niferous tubules  had  enlarged  slightly  to 
a  diameter  of  80  to  lOO/i. 

Yearling  males  tended  to  emerge  later 
than  adult  males,  were  of  smaller  body 
size  (Morton  and  Parmer,  in  press),  and 
had  considerably  smaller  abdominal  testes 
(Fig.  1).  This  was  most  clearly  ob- 
served at  Tioga  Pass  where  we  had  a 
larger  pool  of  marked  animals  to  collect 
from.  Testes  of  yearlings  were  about  three 
times  heavier  at  emergence  than  when 
they  entered  hibernation  the  previous  fall 
as  juveniles.  A  decrease  in  testicular 
weight  occurred  soon  after  emergence,  fol- 
lowed by  an  increase  toward  the  end  of 
the  season. 

Two  yearlings  collected  at  Big  Bend  on 
18  April  1973  had  paired  testes  weighing 
920  and  940  mg.  Seminal  vesicles  of  these 
animals  were  37.4  and  34.2  mg,  respec- 
tively. Although  well  above  resting  level, 
neither  set  of  glands  approached  those  of 
sexually  active  adults  in  weight  or  cel- 
lular maturation. 

There  was  some  evidence  of  a  cycle  in 
spermatogenic  activity  in  yearling  testes 
in  that  a  few  spermatocytes  were  pro- 
duced early  in  the  season.  Diameter  of 
seminiferous  tubules  remained  small  (be- 
low lOO/i)  throughout  the  season,  and  no 
advanced  stage  of  spermatogenesis  was 
found  in  any  yearling  testis.  As  in  adults 
at  Tioga  Pass,  tubule  diameter  increased 
slightly  in  concert  with  increased  testic- 
ular weight,  and  spermatoc>'te  numbers 
increased    just   prior   to   hibernation. 

Seminal  vesicles. —  Seminal  vesicles 
of  adults  tended  to  increase  in  weight  for 
a  few  days  following  emergence,  re- 
mained   at    maximum    size    for    a    few 


Dec.  1975 


MORTON,  GALLUP:    GROUND  SQUIRREL 


429 


weeks,  then  decreased  to  minimum  size 
for  the  rest  of  the  season  (Fig.  2).  These 
glands  were  only  slightly  enlarged  in 
yearlings  early  in  the  season  and  tended 
to  decrease  in  weight  thereafter.  As  shown 
in  S.  lateralis  by  McKeever  (1964),  sem- 
inal vesicle  growth  is  controlled  by  tes- 
ticular hormone.  The  near-maximum  size 
of  seminal  vesicles  in  recently  emerged 
S.  h.  heldingi  suggests  that  upon  final 
arousal  they  have  fully  secretory  testes. 
In  microscopic  appearance  the  heaviest 
seminal  vesicles  of  adults  had  a  distended 
mucosal  epithelimn  and  lumina  filled 
with  seminal  fluid  in  a  colloidal  state.  As 
involution  occurred  the  colloid  disap- 
peared, the  mucosal  layer  became  shrun- 
ken and  folded,  and  the  lumina  nearly 
disappeared.  This  appearance  was  main- 
tained through  onset  of  dormanc}'.  Sem- 
inal vesicles  of  yearlings  were  without  de- 
tectable cellular  change  throughout  the 
active  season. 

5exual  cycle  of  females. —  Adult  fe- 
males appeared  to  be  sexually  receptive 
almost  immediately  after  emergence  as 
judged  by  their  swollen,  open  vulvae  and 
enlarged,  turgid  uteri.  Copulation  was 
never  observed,  but  additional  evidence 
that  mating  occurred  soon  after  emer- 
gence is  that  a  few  adults  were  already 
lactating  during  the  fifth  week  after  the 
first  active  females  were  seen.  Gestation 
period  in  S.  beldingi  is  thought  to  be  27 
to  31  days   (Turner  1972). 

Yearlings  were  in  estrous  later  in  the 
season  than  adults  because  they  tended  to 
emerge  later  and  because  estrous  appeared 
to  be  delayed  in  smaller  yearlings  until 
additional   body   growth   had   occurred. 

Seasonal  changes  in  ovarian  weight 
were  about  two-fold  and  were  similar  for 
the  two  age  groups.  For  the  first  six  weeks 
after  emergence  ovaries  weighed  20  to  30 
mg.  Weight  then  decreased  to  8  to  15  mg 
for  the  remainder  of  the  season. 

Anovulatory  follicles  and  corpora  lutea 
tended  to  enlarge  during  gestation  and 
reached  maximum  diameters  at  partu- 
rition. These  structures  shrank  in  post- 
partum females.  By  the  end  of  lactation 
follicular  cavities  were  much  reduced  or 
absent  and  corpora  lutea  were  becoming 
indistinct.  No  ovarian  recrudescence  was 
observed  in  yearlings  or  adults  prior  to 
hibernation,    but    follicular    enlargement 


1 

• 

Big  Bend 

•    AcJuM 

. 

, 

o    Yeorlmg 

. 

• 

; 

8 

.. 

•• 

; 

» 

oo 

i 

°           '«         o 

Fig.  2.  Paired  seminal  vesicle  weights  of 
Spermophilus  beldingi  beldingi  throughout  the 
active  season  at  two  study  areas.  Note  log  scale 
on  ordinate. 


began  in  juveniles  during  their  last  weeks 
of  activity  after  older  animals  had  al- 
ready become  dormant. 

Emergence  of  females  at  Tioga  Pass 
occurred  over  a  span  of  several  weeks. 
As  a  result  parturition  dates  were  spread 
out  and  the  percentage  of  females  lac- 
tating at  a  given  time  was  almost  never 
100  percent  (Fig.  3).  As  far  as  we  know, 
all  females  reproduced,  although  some  of 
the  smaller  yearlings  collected  in  late 
June  at  Tioga  Pass  had  unscarred,  thin, 
virgin-like  uteri.  Note  that  testicular 
atrophy  was  well  advanced  in  adult  males 
at  that  time  (Fig.  1,  lower).  It  is  possible 
that  a  few  yearling  females  did  not  bear 
young.  Another  possibility  is  that  implan- 
tation was  delayed  in  those  of  small  body 
size.  There  are  indications  in  other  studies 
that  female  Spermophilus  may  become 
impregnated  well  after  testicular  collapse 
has  begun  in  males  (Wells  1935;  Tomich 
1962;  McKeever  1966). 

If  these  smaller  females  do  have  young, 
they  will  be  born  relatively  late  in  the 
season  (we  have  observed  considerable 
disparity  in  size  of  juveniles  at  the  end 
of    the    season;    Morton,    Maxwell,    and 


430 


GREAT   BASIN   NATURALIST 


Vol.  35,  No.  4 


Fig.  3.     Mean  percentage  of  female  Spermophilus  beldingi  beldingi  thought  to  be  lactating  at  spe- 
cific 5-clay  intervals  at  Tioga  Pass.   Numerals  indicate  sample  sizes. 


Wade  1974),  and  if  these  young  survive, 
a  self-perpetuating  cycle  of  late  repro- 
duction by  small  yearling  females  will 
have  been  established. 

Litter  size. —  Our  specimens  and  rec- 
ords of  frequently  retrapped  females  in- 
dicate that  they  have  one  litter  per  sea- 
son. This  was  true  even  of  the  earlist  fe- 
males to  breed.  We  obtained  measure- 
ments of  litter  size  through  counts  of 
placental  scars  and  implanted  embr3'os 
in  collected  specimens  and  through  litters 
born  in  captivity  (Table  1).  The  differ- 
ence in  means  between  counts  of  scars 
and  of  embryos  was  not  different  ac- 
cording to  a  t  test  (P>  0.05)  but  some 
prenatal  loss  did  occur.  Ten  of  the  228 
embryos  examined  (4.4%)  were  partially 
reabsorbed.  This  is  similar  to  the  re- 
absorption  rate  found  in  S.  lateralis 
(McKeever  1964;  Skryja  and  Clark  1970) 
and  in  S.  richardsonii  (Sheppard  1974). 
The  lowest  estimate  of  litter  size  was  ob- 
tained from  births  in  captivity.  Cannibal- 
ism of  their  young  by  confined  females 


Table  1 .  Measurements  of  litter  size  in  Spermo- 
philus beldingi  beldingi. 


Mean 

S.D. 

N 

Placental  scars 

Adults 
Yearlings 

6.88 
5.00 

1.22 

17 
2 

Implanted  embryos 
Adults 
Yearlings 

6.33 
4.75 

2.01 
1.07 

21 
20 

Young  per  captive 
Adults 
Yearlings 

5.71 
4.00 

1.45 
1.00 

17 
5 

Total,   all   measurements 

Adults 
Yearlings 

6.31 
4.48 

1.67 
1.40 

55 
27 

was   noted   in   a   few   cases   but   all    cases 
may  not  have  been  detected. 

When  all  measurements  were  summed, 
mean  litter  size  of  adults  (6.31)  was  sig- 
nificantly larger  (P<  0.01)  than  that  of 
yearlings  (4.48).  This  is  quite  similar  to 
age  differences  in  litter  size  observed  in 
S.  armatus  (Slade  and  Balph  1974). 


Dec.  1975 


MORTON.   GALLITP:    OHOUNI)   SQl'IKKEL 


431 


Discussion 

In  most  published  reports  there  is  little 
indication  that  yearling  Sperniophilus  of 
either  sex  are  functionally  or  visually- 
separable  from  older  animals  (compare 
Mayer  and  Roche  1954;  McKeever  1963, 
1964;  Carl  1971;  Zimmerman  1972),  al- 
though they  may  constitute  a  considerable 
portion  of  the  breeding  population.  How- 
ever, yearlings  engage  in  unique  behav- 
ioral interactions  with  older  animals  as 
they  are  integrated  into  the  conmnniity 
of  reproducers  ( Michener  and  Michener 
1973)  and  should  be  the  focus  of  more 
study.  Even  in  the  larger-bodied  sciurids 
such  as  Marmota  monax  (Christian,  Stein- 
berger,  and  McKinney  1972)  and  Cy- 
nomys  leucurus  (Bakko  and  Brown  1967) 
some  males  are  sexually  mature  as  year- 
lings. 

The  lack  of  I'eproductive  ca])acity  in 
yearling  male  but  not  in  female  S.  b.  hel- 
dingi  is  an  interesting  contrast  that  indi- 
cates the  operation  of  sex-specific  selec- 
tion factors.  As  pointed  out  by  Conaway 
(1971)  the  nonpregnant  cycle  is  a  rarity 
and  cannot  be  afforded  by  most  natural 
populations.  The  breeding  season  is  de- 
layed slightly  in  yearling  female  S.  b. 
beldingi  by  their  emergence  schedule  and 
the  apparent  necessity  for  some  to  attain 
greater  body  size  before  pregnancy.  Nev- 
ertheless, all  or  nearly  all  do  reproduce. 
In  males,  however,  there  is  greater  total 
body  size  to  be  reached  than  in  females 
(Morton  1975)  and  selection  seems  to 
have  favored  the  strategy  of  diverting 
energy  expenditure  from  reproductive 
activities  toward  growth.  Indeed  from  the 
last  half  of  the  yearling  season  onward 
males  are  significantly  larger  than  fe- 
males (Morton  and  Parmer,  in  press). 
We  have  additional  evidence  that  delayed 
sexual  development  in  male  S.  b.  beldingi 
is  tied  directly  to  body  size.  Two  males 
born  in  captivity  and  kept  under  class- 
room conditions  for  display  became  ex- 
cessively obese  in  their  first  autumn,  went 
through  bouts  of  estivation  for  several 
months,  and  possessed  scrotal  testes  soon 
after  resuming  normothermia  at  about 
nine  months  of  age. 

Growth  rates  of  captive  S.  b.  beldingi 
juveniles  are  comparable  to  those  of  other 
hibernatory  Sperniophilus,  but  there  are 
indications  that  growth  is  much  slower  in 
feral   animals   than   in   captives   even   be- 


fore weaning  (Morton  and  Tung  1971). 
It  may  be  that  it  is  unusually  difficult  for 
S.  b.  beldingi  living  at  high  altitude  to 
achieve  adult  size  and  reproductive  i)o- 
tential  simultaneously.  We  have  sug- 
gested (Morton  and  Tung  1971)  and  now 
shown  clearly  (Morton  and  Parmer,  in 
press)  that  iS.  b.  beldingi  do  not  reach 
maximum  body  size  until  late  in  their 
second  year  of  life  or  beyond.  Likewise, 
Sheppard  (1972)  found  that  yearling  S. 
richardsonii  aged  on  the  bases  of  eye  lens 
weight,  tooth  wear,  and  epiphyseal  clo- 
sure had  smaller  mean  body  weights  than 
older  animals. 

The  habitual  lack  of  a  reproductive 
cycle  among  certain  male  members  of  a 
population  could  not  be  tolerated  unless 
a  mating  system  were  employed  that  as- 
sured impregnation  of  all  receptive  fe- 
males. A  priori  one  might  expect  repro- 
ductively  active  males  in  such  a  system 
to  be  polygamous.  The  details  of  the  mat- 
ing system  of  S.  b.  beldingi  should  be 
elucidated  shortly  by  behavioral  studies 
in  progress  at  Tioga  Pass  by  Paul  Sher- 
man of  the  University  of  Michigan. 

The  social  system  of  a  ground  squirrel 
population  has  recently  been  implicated 
in  sexual  development  of  yearling  males. 
Slade  and  Balph  (1974)  found  that  year- 
ling male  S.  armatus  seldom  had  scrotal 
testes  and  rarely,  if  ever,  bred.  After  the 
population  was  artificially  reduced,  how^- 
CAer,  many  yearling  males  were  sexually 
active.  Slade  and  Balph  associate  this 
precocity  with  low  harassment  as  juve- 
niles, early  arousal  from  hibernation,  and 
decreased  encounters  with  aggressive 
squirrels  following  emergence.  They  did 
not  report  on  body  size  in  these  animals. 

It  is  possible  but  improbable  that  we 
have  been  studying  a  ground  squirrel 
population  with  unique  growth  patterns 
and  mating  system.  It  is  important  to 
recognize  that  accurate  aging  in  many 
studies  has  not  been  possible  due  to  their 
brevity  or  to  lack  of  history  on  recogniz- 
able individuals.  For  example,  in  his 
study  of  S.  b.  oregonus  conducted  in  Las- 
sen County,  California,  at  1,370  to  1,730 
m  elevation,  McKeever  (1963)  refers 
only  to  adults  and  jmeniles.  Body  weights 
shown  for  adult  S.  b.  oregonus  are  5  to  10 
percent  lower  throughout  the  season  than 
those  of  S.  b.  beldingi  (Morton  1975), 
but   juvenile  5.    b.   oregonus  are  at  least 


432 


GREAT    BASIN    NATURALIST 


Vol.  36,  No.  4 


20  percent  heavier  than  S.  b.  hcldingi 
(Morton,  Maxwell  and  Wade  1974)  be- 
fore entering  hibernation.  Mean  maxi- 
mum testicular  weight  occurs  in  both 
subspecies  at  the  beginning  of  the  active 
season,  but  it  is  about  three  times  greater 
in  S.  b.  beldingi  than  in  S.  b.  oregonus. 
This  seems  anomalous  to  us  and,  coupled 
with  bod}'  weight  data  and  McKeever's 
observation  that  only  70  percent  of  adult 
male  S.  b.  oregonus  were  sexually  active, 
suggests  that  at  least  some  j^earling  S.  b. 
oregonus  could  be  distinguished  from 
older  animals  on  the  bases  of  body  size 
or  reproductive  capacity  if  ages  were 
known . 

Finally,  it  should  be  recognized  that 
unusual  ecological  conditions  exist  at 
high  altitude.  Many  factors,  both  terres- 
trial and  extraterrestrial  in  origin,  could 
function  as  inhibitors  of  growth  and  de- 
velopment   in    young    animals. 

Seasonal  breeding. —  Gro\^i;h  of  the 
reproductive  tract  during  the  whole  hi- 
bernation period  is  well  known  among 
Spermop/iilus.  Remarkably,  gonadal  re- 
crudescence, at  least  of  testes,  begins 
even  prior  to  dormancy  in  several  spe- 
cies with  short  active  seasons  such  as  S. 
undulatus  (Mitchell  1959;  Hock  1960), 
5.  b.  oregonus  (McKeever  1963),  S.  later- 
alis (McKeever  1964),  S.  richardsonii 
(Clark  1970),  and  5.  b.  beldingi  (pres- 
ent study).  McKeever  (1963)  noted  that 
S.  b.  oregonus  emerged  with  testes  of  max- 
imum size.  Usually  the  final  stages  of 
spermatogenesis  in  hibernatory  Spermo- 
philus  are  not  reached  until  ten  days  or 
more  after  emergence  even  in  species  at 
high  latitude  (Hock  1960).  The  total  read- 
iness of  5.  b.  beldingi  to  reproduce  at  time 
of  emergence  is  undoubtedly  a  response 
to  conditions  imposed  by  the  short  .sum- 
mers of  high   altitude. 

Seasonal  breeding  is  highly  adaptive 
in  that  young  are  born  at  a  time  favorable 
for  their  survival.  At  high  altitufle  this 
favorable  season  is  comprcss(Hl  temporally 
and  has  rather  sliarply  drawn  boundaries. 
To  cope  successfully  with  these  circum- 
stances the  cycle  of  sexual  maturation  in 
S.  b.  beldingi  is  completed  during  dor- 
mancy.  A  corollary  is  that  reproductive 
preparation  must  ha\('  a  precise  })hase 
relationship  to  average  snowTnelt  patterns 
and  related  euA  ironmental  effects.  There 
is    sensitivity,    however,    on    the    part    of 


newly  emerged  animals  to  ambient  con- 
ditions. At  Tioga  Pass  considerable  an- 
nual variation  in  snowpack,  schedule  of 
snowmelt,  and  emergence  of  vegetation 
were  documented  (Morton,  Maxwell, 
and  Wade  1974;  Morton,  in  press).  The 
schedule  of  S.  b.  beldingi  was  affected  ac- 
cordingly. In  1969,  for  exam])le,  snow- 
pack  was  about  240  percent  above  nor- 
mal, whereas  in  1972  it  was  35  percent 
below  normal.  As  judged  by  subsequent 
emergence  times  of  juveniles  and  their 
growth  curves,  reproduction  occurred 
about  three  weeks  later  in  1969  than  in 
1972  (Morton,  Maxwell,  and  Wade 
1974).  Reproduction  is  not  delayed  in- 
ordinately, however,  even  in  heavy 
snow  years,  because  about  25  percent  of 
prehibernatory  fat  reserves  still  remain 
at  emergence,  ])ro-\iding  a  buffer  to  food 
requirements  during  the  first  weeks  of 
activity   (Morton  1975). 

Acknowledgments. —  Many  Occiden- 
tal College  students  participated  in  por- 
tions of  this  study.  We  are  grateful  to  all 
of  them  but  would  like  to  acknowledge  in 
particular  Phil  Rekey,  Cassie  Cusick, 
Shena  Huang,  Roland  Leong,  Cathy 
Maxwell,  Robert  Parmer,  and  Charlie 
Wade.  Financial  support  was  provided  by 
Occidental  College  and  by  National 
Science  Foundation  Grant  GR  29146X1. 

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LTniv.  Arizona,  149  pp. 

Wells,  L.  J.  1935.  Seasonal  sexual  rhythm 
and  its  experimental  modification  in  the 
male  of  the  thirteen-lined  ground  squirrel 
{Citellus  tridecemlineatus) .  Anat.  Rec.  62: 
409-447. 


A  NEW  COMBINATION  IN  PENSTEMON 
( SCROPHULARI ACEAE) 

Stephen  L.  Clarki 

Abstract. —  Field  and  herbarium  studies  of  Penstenion   cyananthus   Hook.   ssp.   longiflorus   Pennell 
suggest  that  this  taxon  be  elevated  to  species  rank. 


Penstemon  cyananthus  Hook.  ssp.  longi- 
florus Pennell  is  a  tall,  attractive  blue- 
flowered  plant  presently  known  only  from 
Beaver,  Millard,  and  Piute  counties  of 
southern  Utah.  It  was  first  collected  by 
Edward  Palmer  (376,  NY)  near  Beaver 
City,  Utah,  and  was  distributed  as  P. 
glaber  cyananthus,  a  synonym  of  P.  cyan- 
anthus. 

Pennell  recognized  that  Palmer's  col- 
lection differed  from  the  species  as  fol- 
lows: the  calyx  lobes  are  broader  than  in 
the  species,  the  flowers  longer,  the  stems 
and  lower  leaves  are  puberulent,  and  the 
inflorescence  is  strongly  secund.  In  ad- 
dition, the  distributions  are  verj'  differ- 
ent. Penstemon  cyananthus  ssp.  cyanan- 
thus has  never  been  collected  from  south- 
ern Utah,  but  it  is  one  of  the  most  fre- 
quenty  encountered  species  of  Penstemon 
in  the  Wasatch  mountains  of  northern 
Utah,  southeastern  Idaho,  and  southwest 
Wyoming  (Fig.  1). 

On  the  basis  of  these  morphological  and 
geographical  differences,  Pennell  (1920) 
described  and  named  the  subspecies  P. 
cyananthus  ssp.  longiflorus. 

It  is  the  writer's  opinion  that  differ- 
ences of  such  magnitude  warrant  the  ele- 
vation of  this  taxon  to  specific  status, 
hence  the  following  new  combination  is 
proposed: 

Penstemon  longiflorus  (Pennell)  Clark 
comb,  nov.,  based  on  P.  cyananthus  Hook, 
ssp.  longiflorus  Pennell  in  Contr.  from 
the  U.S.  Nat.  Herb.,  vol.  20,  part  9,  p. 
353,  1920. 


Penstemon  longiflorus  can  easily  be 
recognized  by  its  puberulent  lower  stem 
and  leaves,  broad  sepals,  longer  corolla, 
secund  inflorescence,  and  its  blue  tipped 
staminode. 

Both  taxa  have  a  chromosome  number 
of  2n  =  16. 


\                     1 

"T 

(Xrif'* 

AS> 

•     ; 

1^ 

V|,..i.i-i 

1 — 

.... 

—  — ^^ — ^__ 

.._ ■}■/ 

y 

"\ 

...Jr 

Fig.   1.   Distribution  of  P.  cyananthus   (circles) 
and  P.  longiflorus  (squares). 


'Department    of    Botany,     Weber    State    College,     Ogden.     Utah    84043. 


434 


SOME  RELATIONSHIPS  BETWEEN  WATER  FERTILITY  AND 

EGG  PRODUCTION  IN  BROWN  TROUT  {SALMO  TRUTTA) 

FROM  MONTANA  STREAMS 

Lawrence  L.  LockarcP 

Abstract. —  Relationships  between  water  fertility  (as  measured  by  conductivity  and  alkalinity)  of 
17  Montana  streams  and  the  attainment  of  sexual  maturity  and  fecundity  of  their  resident  female 
brown  trout  were  studied.  Fish  from  the  streams  having  conductivity  and  alkalinity  levels  greater  than 
100  micromhos/cm  and  ppm  CaCO,^,  respectively,  were  younger  at  sexual  maturity  than  fish  from 
waters  with  lower  levels.  The  attainment  of  earlier  sexual  maturity  in  fish  from  the  former  streams 
could  not  be  completely  explained  on  the  basis  of  greater  growth  rates.  Fish  from  the  stream 
having  the  highest  levels  of  conductivity  had  the  slowest  growth  rate  but  became  sexually  mature  at 
the  youngest  age.  A  positive  relationship  was  found  between  chemical  fertility  of  streams  and  the 
fecundity-  of  their  fish.  However,  in  the  stream  having  the  highest  levels  of  conductivity,  fish  were 
the  least  fecund.  It  was  concluded  that  the  chemical  fertility  of  these  streams  is  generally  related  to 
the  age  at  sexual  maturity  and  fecundity  of  fish. 


The  size  and  age  at  sexual  maturity 
and  the  fecundity  of  female  fish  appear 
to  be  related  to  features  of  their  environ- 
ment. In  Pennsylvania  brown  trout 
{Salmo  trutta)  from  infertile  waters  had 
a  smaller  proportion  of  mature  fish  per 
age  class  and  smaller  weight  of  eggs  than 
comparable  fish  from  fertile  waters  (Mc- 
Fadden,  Cooper,  and  Anderson  1965). 
Scott  (1962)  and  Bagenal  (1969)  dem- 
onstrated that  rainbow  trout  {Salmo  gaird- 
neri)  and  brown  trout,  respectively, 
brought  a  lower  number  of  eggs  to  ma- 
turity under  reduced  nutritional  levels 
than  fish  on  higher  levels  of  nutrition. 

This  study  is  an  attempt  to  determine 
the  relationships  between  the  conduc- 
tivities and  alkalinities  of  Montana 
streams  and  (1)  the  size  and  age  at  sex- 
ual maturity  and  (2)  the  fecundity  of 
the  brown  trout  in  those  streams.  Field 
collections  were  made  from  8  September 
to  23  October  1972  and  from  1  Septem- 
ber to  19  October  1973. 

Methods 

A  total  of  449  female  brown  trout 
were  collected  by  electrofishing  at  sites 
on  streams  in  the  Clark  Fork  of  the  Co- 
lumbia River  and  in  the  Yellowstone  and 
Missouri  River  drainages  (Fig.  1).  These 
streams  had  a  wide  range  of  physical  and 
chemical  conditions   (Table  1). 

At  least  one  fall,  winter,  and  summer 
measurement  of  conductivity  and  alka- 
linity was  made  at  each  collecting  site. 
The  field  measurements  from  each  stream 


were  averaged  with  the  }'early  conduc- 
tivity and  alkalinity  averages  obtained 
from  Water  Resources  Data  for  Montana 
(U.S.G.S.  1972)  where  available.  Dis- 
charge values  were  obtained  by  aver- 
aging available  yearly  values  from  the 
above  U.S.G.S.  records  with  values 
measured  or  estimated  by  fisheries  biolo- 
gists of  the  Montana  Fish  and  Game  De- 
partment. 

All  fish  were  collected  during  Sep- 
tember and  October  of  1972  and  1973 
(Table  2).  Fish  taken  were  preserved  in 
10  percent  formalin  and  later  washed  in 
water  and  stored  in  40  percent  isopropyl 
alcohol.  Fixation  in  formalin  causes  spe- 
cimens to  shrink  about  3-4  percent  in 
length  and  increase  5-12  percent  in 
weight  (Parker  1963).  After  preservation, 
fish  were  measured,  weighed,  and  scale 
samples  were  removed  for  age  determi- 
nations. Each  fish  was  classified  as  ma- 
ture or  immature  according  to  the  condi- 
tion of  the  eggs  in  its  ovaries.  Mature 
fish  containing  eggs  in  a  gradient  of 
sizes  were  not  used  in  the  fecundity  anal- 
yses because  the  number  of  eggs  is  re- 
duced by  resorption  throughout  the  ma- 
turation period,  and  regressing  eggs  could 
not  be  distinguished  from  maturing  eggs 
in  these  fish.  Only  fish  having  distinct  re- 
cruitment and  maturing  eggs  without  in- 
tervening size  classes  of  eggs  were  used 
in  fecundity  work.  The  ovaries  from  these 
fish  were  removed  and  the  number  of 
maturing  eggs  determined  by  actual 
count. 

The    streams    from    which    collections 


^Current   address:    312    South    Grant    Avenue,    Pierre,    SD    57501. 


435 


436 


GREAT   BASIN   NATURALIST 


Vol.  35,  No.  4 


Table   1.        Selected   chemical   and  physical  features  of  streams  sampled. 


Ck)lIection 
site  no.' 


Location 
of  site 


Conductivity 
(micromhos/cm) 


Alkalinity 
(ppm  CaCOa) 


Discharge 

(C.F.S.) 


1 

Rock  Cr. 

2 

St.  Regis  R. 

3 

Big  Hole  R. 

4 

W.   Gallatin  R. 

5 

Madison  R. 

6 

Baker  Cr. 

7 

O'Dell  Cr. 

'8 

L.  Prickley  Pear  Cr. 

9 

E.  Gallatin  R. 

10 

Shields  R. 

11 

Flagstaff  Cr. 

12 

Beaverhead  R. 

13 

16  Mile  Cr. 

14 

So.  F.  Musselshell  R. 

15 

Little  Blackfoot  R. 

16 

Bluewater  Cr." 

17 

Big  Horn  R. 

18 

Bluewater  Cr.^ 

70 

49 

169 

80 

51 

555 

207 

117 

1,125 

230 

T18 

791 

249 

107 

1,409 

317 

154 

70 

348 

167 

100 

358 

195 

69 

360 

195 

400 

402 

221 

159 

405 

197 

5 

521 

193 

405 

522 

195 

50 

561 

243 

83 

612 

188 

105 

798 

209 

18 

805 

188 

3,500 

1,387 

214 

28 

'See  Figure  1 

^Section  above  Bluewater  Fish  Hatchery 

'Section  below  Bluewater  Fish  Hatcherj' 


Fig.   1.  Map  showing  location  of  collecting  sites. 


were  made  were  grouped  into  classes  pri- 
marily on  the  basis  of  similarities  in  con- 
ductivities followdng  the  technique  used 
by  McFadden  et  al.  (1965).  Streams  from 
which  collections  1  and  2  were  taken  each 
had  less  than  100  units  of  conductivity 
and  alkalinity  and  formed  Class  I. 
Streams  from  which  collecting  sites  3 
through  18  were  located  had  alkalinity 
values     above     100:     thus     conductivities 


were  used  as  the  primary  indicator  of 
water  fertility.  Class  II  contained  streams 
on  which  collection  sites  3  through  11 
were  located.  These  streams  had  conduc- 
tivities ranging  from  207-405  micromhos/ 
cm  at  25  C.  Streams  of  collecting  sites  12 
through  15  had  conductivities  of  from 
521-612  and  comprised  Class  III  except 
for  the  analysis  of  size  and  age  at  sexual 
maturity  in  which  Stream  17  was  includ- 


Dec.  197; 


LOCKARD:     BROWN    TROUT 


437 


Table  2.     The  location,  date,  and  number  of  fish  collected. 


1972 

1973 

Collection 

Number 

Collection 

Number 

Total 

Collection  site 

date 

fish 

date 

fish 

fish 

1      (Rock  Cr.) 

Oct.  13 

16 

Sept.    14 

17 

33 

2     (St.  Regis  R.) 

Sept.    1 

22 

22 

3     (Big  Hole  R.) 

Oct.  9 

17 

17 

4     (W.  Gallatin  R.) 

Oct.   18 

17 

Sept.  25 

11 

28 

5     (Madison  R.) 

Sept.  21 

13 

Sept.  20 

19 

32 

6     (Baker  Cr.) 

Oct.   10 

17 

Sept.  26 

9 

26 

7     (O'Dell  Cr.) 

Sept.  22 

11 

Sept.  21 

8 

19 

8     (L.  Prickley  Pear  Cr.) 

Oct.  23 

16 

Oct.  2 

17 

33 

9     (E.  Gallatin  R.) 

Sept.  18 

15 

Oct.  4 

16 

31 

10     (Shields  R.) 

Oct.  6 

11 

Sept.  24 

6 

17 

11      (Flagstaff  Cr.) 

Oct.  9 

9 

9 

12     (Beaverhead  R.) 

Sept.  26 

14 

Oct.   17 

"9 

23 

13      (16  Mile  Cr.) 

Sept.   11 

13 

Oct.   11 

14 

27 

14     (So.  F.  Musselshell  R.) 

Sept.  24 

13 

13 

15      (Little  Blackfoot  R.) 

Oct.  1 

28 

28 

16     (Bluewater  Cr.)' 

Sept.  8 

T3 

Sept.  12 

14 

27 

17     (Big  Horn  R.) 

Sept.  15 

14 

14 

18      (Bluewater  Cr.)' 

Sept.  8 

'\3 

Sept.   12 

37 

50 

^Section    above    Bluewater    Fish    Hatchciy 
-Section    below    Bulewater    Fish    Hatchery 


ed.  The  streams  of  Class  II  and  III  were 
combined  into  Class  IV  because  their 
fish  had  similar  relationships  to  conduc- 
tivity. Class  V  was  made  up  of  Bluewater 
Creek  on  which  collecting  sites  16  and 
18  were  located.  These  collecting  sites 
were  grouped  together  primarily  because 
of  their  high  conductivities. 

The  fish  in  stream  classes  were  statis- 
tically compared  by  selected  procedures 
and  techniques  from  "Statistical  Meth- 
ods" (Snedecor  and  Cochran  1971)  and 
"Statistical  Methods"  (Arkin  and  Colton 
1972).  Additional  techniques  were  pro- 
vided by  Dr.  R.  E.  Lund,  Alathematics  De- 
partment,  Montana   State   University. 

Results 

Size  and  Age  of  Sexually  Mature 
Female  Brown  Trout 

Generally  the  attaimnent  of  sexual  ma- 
turity of  fish  is  dependent  on  size  and 
age.  Inspection  of  age  groups  within 
stream  classes  indicated  an  apparent  ten- 
dency for  a  higher  proportion  of  the 
larger  females  to  be  sexually  mature 
(Table  3).  To  test  the  linearity  of  this 
trend,  regressions  were  made  on  fish  from 
age  groups  in  stream  classes  showing  an 
increase  in  sexual  maturity  with  increas- 
ing length.  In  age  group  I,  fish  from  Class 
V  streams  showed  a  significant  positive 
linear  relationship  between  length  and 
sexual  maturity  (P=  0.001).  In  age  group 


II,  a  significantly  higher  proportion  of 
larger  fish  were  sexually  mature  in 
Stream  Classes  I,  II,  III,  and  IV  with  P 
values  of  less  than  0.05.  In  age  group 
III  +  ,  fish  from  Stream  Classes  II  and  IV 
had  significant  positive  linear  relation- 
ships between  length  and  sexual  maturity 
(P<0.05).  McFadden  et  al.  (1965)  found 
a  tendency  within  a  given  year  class  for 
a  higher  percentage  of  larger  than  smaller 
fish  to  be  sexually  mature. 

The  effect  of  age  on  the  attainment  of 
sexual  maturity  in  fish  was  determined 
by  comparing  the  proportions  of  sexual- 
ly mature  fish  between  age  groups  by  a 
technique  of  R.  E.  Lund.  Only  2  percent 
of  age  group  I  fish  8.0-13.9  inches  long 
from  Class  IV  streams  were  mature,  while 
34  percent  of  comparably  sized  fish  in  age 
group  II  were  mature.  The  difference  in 
proportions  was  significant  with  a  P  = 
0.001.  There  were  significantly  fewer  ma- 
ture 6.0-8.9  inch  fish  from  Class  V  streams 
in  age  group  I  than  in  age  group  II  (P  = 
0.08).  These  combined  probability  values 
demonstrated  a  significantly  (P=  0.001) 
higher  proportion  of  age  II  fish  were  ma- 
ture than  age  I  fish.  Significantly  more 
of  size  group  10.0-19.9  inch  fish  from 
Class  IV  streams  were  mature  at  age 
III+  than  age  II  (P=  0.001).  This  rela- 
tionship of  a  higher  percentage  of  older 
females  being  sexually  mature  than  youn- 
ger females  in  the  same  size  group  has 
been  reported  by  McFadden  et  al.  (1965). 


438 

GREAT   BASIN    NATURALIST 

Vol.  35. 

,  No.  4 

Table 

3.     Size  and 

age  of  sexually 

mature 

female 

brown  t 

rout  by 

stream 

classes 

Length 
(inches) 

Stream 

classes 

Age 

I 

II 

III 

IV 

V 

group 

#Fish 

%Mat. 

#Fish 

%Mat 

.  #Fish 

re  Mat. 

#Fish 

i%Mat.  #Fish 

-^cMat. 

I 

4.0-4.9 

0 

0 

0 

0 

5 

0 

5.0-5.9 

0 

0 

0 

0 

3 

33 

6.0-6.9 

3 

0 

0 

0 

0 

16 

31 

7.0-7.9 

4 

0 

1 

"o 

0 

1 

0 

17 

47 

8.0-8.9 

>1 

0 

7 

0 

2 

0 

9 

0 

6 

100 

9.0-9.9 

1 

0 

6 

0 

5 

0 

11 

0 

0 

10.0-10.9 

0 

11 

0 

10 

0 

21 

0 

0 

ll.O-n.9 

0 

0 

7 

0 

7 

0 

0 

12.0-12.9 

0 

0 

2 

0 

2 

0 

0 

13.0-13.9 
Total 

0 

0 

2 

50 

2 

50 

0 

II 

Q 

0 

25 

0 

28 

4 

53 

2 

47 

43 

6.0-6.9 

u 

0 

0 

0 

1 

100 

7.0-7.9 

0 

0 

0 

0 

6 

67 

8.0-8.9 

2 

0 

1 

0 

0 

1 

0 

6 

83 

9.0-9.9 

9 

33 

5 

0 

2 

0 

7 

0 

5 

100 

10.0-10.9 

6 

33 

16 

13 

9 

11 

25 

12 

1 

100 

n.0-11.9 

1 

100 

29 

31 

3 

0 

32 

28 

1 

100 

12.0-12.9 

2 

100 

23 

61 

9 

22 

32 

50 

4 

100 

13.0-13.9 

T 

100 

9 

67 

7 

57 

16 

63 

0 

14.0-14.9 

0 

19 

84 

3 

100 

22 

86 

0 

15.0-15.9 

0 

3 

67 

5 

80 

"s 

75 

0 

16.0-16.9 

0 

1 

ilOO 

4 

100 

5 

100 

0 

17.0-17.9 

0 

1 

100 

3 

100 

4 

100 

0 

18.0-18.9 

0 

0 

2 

100 

2 

100 

0 

19.0-19.9 
Total 

0 

0 

1 

100 

T 

100 

0 

21 

43 

107 

48 

48 

50 

155 

48 

24 

88 

III  + 

8.0-8.9 

0 

0 

0 

0 

I 

100 

9.0-9.9 

0 

0 

0 

0 

1 

100 

10.0-10.9 

3 

67 

1 

0 

0 

1 

-9 

0 

11.0-11.9 

8 

75 

^ 

67 

1 

100 

4 

75 

2 

100 

12.0-12.9 

1 

100 

4 

100 

2 

100 

6 

100 

0 

13.0-13.9 

5 

100 

12 

92 

6 

100 

18 

94 

0 

14.0-14.9 

2 

50 

n 

92 

4 

75 

17 

88 

1 

100 

15.0-15.9 

2 

100 

16 

100 

9 

100 

25 

100 

1 

100 

16.0-16.9 

2 

100 

14 

93 

1 

100 

15 

93 

0 

17.0-17.9 

1 

100 

8 

100 

2 

100 

10 

100 

0 

18.0-18.9 

1 

100 

5 

100 

3 

100 

8 

100 

0 

19.0-19.9 

0 

3 

100 

1 

100 

4 

100 

0 

20.0-20.9 
Total 

0 

1 

100 

0 

1 

100 

0 

25 

84 

80 

94 

29 

97 

109 

95 

6 

100 

Grand  Total 

55 

55 

212 

59 

105 

51 

317 

57 

77 

61 

Com])arisons  were  made  of  the  pro- 
portions of  sexually  mature  female  brown 
trout  between  stream  classes  using  a  meth- 
od of  Arkin  and  Colton  (1972).  No  sig- 
nificant difference  (0.05  level)  was  found 
in  the  proportion  of  mature  females  in 
Class  II  and  III  streams  either  by  age 
group  or  grand  total  so  further  compari- 
sons by  this  method  were  made  between 
the  fish  of  Stream  Classes  I,  IV,  and  V. 

There  was  no  significant  difference  be- 
tween Class  I  and  IV  streams  in  the  pro- 
portions of  mature  females  in  age  grouji 
II;  however,  Class  IV  streams  had  a  sig- 


nificantly higher  ]:)roj)ortion  of  mature 
females  in  age  group  III+  than  did  Class 
I  streams  (P=  0.054).  Class  V  streams 
had  a  higher  proportion  of  sexually  ma- 
ture females  than  both  Class  I  and  IV 
streams  in  both  age  group  I  (P=  0.014 
and  0.001,  respectively)  and  age  group  II 
(P=    0.001   and   0.001,  respectively). 

Fisher's  randomization  test  (Bradley 
1968)  was  used  to  further  test  the  hypoth- 
esis that  maturation  increases  as  conduc- 
tivity increases.  The  probability  of  obtain- 
ing the  iiu  reased  proportions  of  mature 
lish  ill  all  age  groups  with  the  increasing 


Dec.  1975 


LOCKARD:    BROWN   TROUT 


439 


conductivities  in  Stream  Classes  I,  II, 
III,  and  V  (Table  3)  is  P=  0.00014. 

The  class  I  and  IV  streams  in  this  stud}' 
were  similar  in  conductivit}"  and  alkalin- 
ity to  the  infertile  and  fertile  streams  in 
the  studies  of  McFadden  and  Cooper 
(1962)  and  McFadden  et  al.  (1965).  In 
the  latter  study,  fish  from  fertile  waters 
attained  maturity  at  an  earlier  age  than 
those  from  infertile  waters.  This  was  at- 
tributed partially  to  a  greater  growth  rate 
of  fish  in  fertile  waters;  however,  the 
authors  also  found  higher  proportions  of 
fish  of  the  same  size  and  age  were  sexual- 
ly mature  in   fertile   streams. 

In  the  present  study,  this  latter  rela- 
tionship was  not  observed  among  fish 
from  Class  I  and  IV  streams.  Instead, 
higher  proportions  of  females  of  a  given 
size  and  age  tended  to  be  mature  in  the 
less  fertile  Class  I  streams.  The  differences 
in  age  at  maturity  between  fish  from 
Stream  Classes  I  and  IV,  therefore,  seem 
closely  related  to  differences  in  growth 
rate.  The  distribution  of  sizes  of  speci- 
mens of  given  age  groups  do  indicate 
faster  growth  rates  in  Class  IV  streams 
(Table  3).  McFadden  and  Cooper  (1962) 
also  reported  positive  correlations  between 
growth  rates  of  brown  trout  and  environ- 
mental fertilit}^ 

Class  V  streams  had  higher  conductiv- 
ity and  alkalinity  values  than  any  of  the 
streams  studied  by  McFadden  et  al. 
(1965).  In  the  more  fertile  waters  (Class 
V),  fish  matured  at  younger  ages  than  in 
less  fertile  waters  (Classes  I  through  IV); 
however,  this  early  maturity  in  Class  V 
streams  was  not  due  to  a  faster  growth 
rate  in  fertile  waters.  That  is,  the  smallest 
fish  in  each  age  group  are  found  in  the 
Class  V  streams  (Table  3).  Therefore, 
some  factor  other  than  grow1;h  rate  or 
chronological  age  apparent!}'  influenced 
the  size  and  age  at  which  sexual  maturity 
was  reached  by  fish  from  the  very  fer- 
tile  (chemically)  waters  of  Class  V. 

Fecundity 

Regression  analyses  for  the  number  of 
mature  eggs  in  a  fish  (dependent  variable) 
on  fish  length  (independent  variable) 
were  applied  to  the  fish  of  the  individual 
streams  and  stream  classifications.  T  tests 
for  the  regression  of  numbers  of  eggs  on 
lengths  were  statistically  significant  at  the 
0.05  level  for  fish  from  all  streams  and 


stream  classifications  with  most  probabil- 
ity values  being  less  than  0.01.  Flagstaff 
Creek  and  the  Big  Horn  River  were  omit- 
ted from  analyses  because  of  an  insuf- 
ficient number  of  mature  fish. 

The  regression  lines  of  numbers  of 
eggs  regressed  on  fish  length  in  stream 
classifications  are  shown  in  Figure  2. 
The  regression  lines  with  steeper  slopes 
show  a  greater  increase  in  number  of  eggs 
per  increment  of  length  than  lines  with 
lesser  slopes. 

Regression  coefficients,  slopes  of  the  re- 
gression lines,  were  calculated  for  these 
regressions  on  each  stream  and  stream 
classification  (Table  4).  The  slopes  of 
the  stream  classification  regressions  were 
tested  for  significant  differences  by  a 
method  of  R.  E.  Lund  (Table  5).  Six  of 
the  8  comparisons  of  slopes  of  regressions 
were  significantly  different  at  the  0.05 
level. 

Fish  from  Stream  Class  I  were  less  fe- 
cund than  fish  from  Stream  Class  II  (Fig. 
2  and  Table  4).  The  difference  between 
these  stream  classes  was  statistically  sig- 
nificant at  the  0.05  level  (Table  5).  This 
relationship  of  increased  fecundity  with 
increased   conductivity  is   similar  to  that 

Table  4.  Regression  coefficients  (slopes)  of 
streams   and   stream   classifications. 


Stream  or       No.  of  eggs  regressed 

on  fish  length 

classifi- 

Regression 

Std 

cation 

coefficient 

error 

N 

Rock  Cr. 

213 

22 

18 

St.  Regis  R. 

173 

47 

12 

Stream  Class  I 

254 

22 

30 

Big  Hole  R. 

284 

40 

7 

W.  Gallatin  R. 

426 

83 

14 

Madison  R. 

457 

64 

16 

Baker  Cr. 

164 

53 

13 

O'Dell   Cr. 

248 

51 

14 

L.    Prickley   Pear   Cr. 

236 

43 

16 

E.  Gallatin  R. 

218 

29 

19 

Shields  R. 

252 

43 

16 

Stream   Class   II 

325 

20 

115 

Beaverhead  R. 

172 

98 

15 

16  Mile  Cr. 

143 

48 

15 

So.   F.   Musselshell   R. 

315 

112 

6 

Little  Blackfoot  R. 

249 

65 

15 

Stream  Class  III 

286 

39 

51 

Stream  Class  IV 

318 

18 

166 

Bluewater   Cr.^ 

170 

20 

19 

Bluewater  Cr." 

113 

16 

21 

Stream   Class  V 

147 

13 

40 

^Section  above  Bluewater  Fish  Hatchery 
-Section  below  Bluewater  Fish  Hatchery 


440 


GREAT  BASIN  NATURALIST 


Vol.  35,  No.  4 


"I      I      I      I      r 

11  12  13  14  IS 

Fish  Length   (inches) 


20         21 


■Fig.  2.  The  regression  lines  of  number  of  eggs  on  length  for  fish  in  stream  classifications. 


Table  5.  Comparisons  between  stream  classes 
by  slopes  of  regressions  of  number  of  eggs  on 
fish  length. 


Slope 

Number  of  eggs 

regressed  on 

fish  length 

comparison 

T 

df 

P 

I   vs   II 

2.42 

77 

0.018* 

I    vs   III 

0.72 

46 

0.475 

I   vs  IV 

2.25 

73 

0.028* 

I   vs   V 

4.18 

47 

0.000* 

II    vs    III 

0.90 

75 

0.371 

II  vs  V 

7.61 

149 

0.000* 

III    vs   V 

3.39 

60 

0.001* 

IV  vs  V 

7.80 

174 

0.000* 

•Significant  at  the  0.05  level 


found  by  McFadden  et  al.  (1965)  in  fish 
from  infertile  and  fertile  streams  having 
conductivities  and  alkalinities  similar  to 


those  of  Class  I  and  II  streams  in  this 
study. 

Fish  from  Stream  Class  III  appeared  to 
be  more  fecund  than  fish  from  Stream 
Class  I  (Fig.  2  and  Table  4).  This  rela- 
tionship of  increased  fecundity  with  in- 
creased conductivity  was  not  statistically 
significant  at  the  0.05  level  (Table  5). 
Stream  Class  III  contained  streams  with 
higher  levels  of  conductivity  than  those 
reported  by  McFadden  et  al.   (1965). 

Fish  from  Stream  Class  IV  (Stream 
Classes  II  and  III  combined)  represent 
fish  from  a  broad  category  of  chemically 
fertile  streams  with  conductivities  from 
about  200  to  600  micromhos/cm.  In  gen- 
eral, these  fish  were  more  fecund    (P= 


Dec.  1975 


LOCKARD:    BROWN    TROUT 


441 


0.028)  than  fish  from  Class  I  streams, 
which  represent  chemically  infertile 
waters. 

Fish  from  Stream  Class  V,  which  had 
the  highest  conductivity,  had  the  lowest 
fecundity.  The  conductivity  values  of 
this  stream  were  about  three  times  great- 
er than  the  highest  values  reported  by 
McFadden  and  Cooper  (1962).  The  above 
results  suggest  some  factor  other  than 
conductivity  is  determining  the  fecun- 
dity of  fish  in  this  stream  class. 

Summary 

An  inverse  relationship  between  chemi- 
cal fertility  and  age  at  sexual  maturity 
was  found  in  brown  trout  from  streams 
of  Montana  in  this  study.  This  same  re- 
lationship between  the  chemical  fertility 
of  streams,  as  measured  by  conductivities 
and  alkalinities,  and  the  age  of  sexual  ma- 
turity of  brown  trout  from  Pennsylvania 
has  been  reported  by  McFadden  et  al. 
(1965).  They  suggested  this  relationship 
was  due  partially  to  fish  in  fertile  streams 
having  greater  growth  rates.  Growth  rate 
may  account  for  the  age  at  maturity  in 
fish  from  1 6  of  the  1 7  streams  in  this 
study,  but  cannot  explain  the  age  at  ma- 
turity in  fish  from  Bluewater  Creek.  Fish 
from  Bluewater  Creek  attained  sexual  ma- 
turity much  earlier  than  fish  from  less 
fertile  streams;  howe\er,  these  fish  from 
the  stream  with  the  highest  conductivity 
had  the  poorest  growth  rates  of  all  the 
fish  studied.  This  shows  growth  rate  was 
not  the  determining  factor  in  the  attain- 
ment of  sexual  maturity  for  fish  from 
Bluewater  Creek. 

McFadden  et  al.    (1965)    found  a  posi- 


tive relationship  between  the  chemical 
fertility  of  streams  and  the  fecundity  of 
their  fish.  A  similar  relationship  was 
found  between  chemical  fertility  and  fe- 
cundity in  fish  of  this  stud}'  from  streams 
having  conductivities  similar  to  those 
studied  by  McFadden  et  al.  (1965).  How- 
ever, fish  from  Bluewater  Creek,  chemi- 
cally the  most  fertile  stream,  were  the 
least  fecund.  Generally  the  age  at  sexual 
maturity  of  fish  from  all  stream  classes 
and  the  fecundity  of  fish  from  Stream 
Classes  I,  II,  III  and  IV  appeared  to  be 
related  to  the  chemical  fertility  of  their 
streams. 

Literature  Cited 

Arkin,  H.,  and  R.  R.  Coi.ton.      1972.     Statistical 

methods.  Barnes  and  Noble  Books,  New  York. 

344  pp. 
Bagenal,  T.  B.      1969.     The  relationship  between 

food    supply    and    fecundity    in    brown    trout 

Salmo  trutta  L.  J.  Fish  Biol.   1:167-182. 
Bradley,  J.  V.     1968.     Distribution-free  statistical 

test.   Prentice-Hall,   New  Jersey.   399  pp. 
McFadden.  J.  T..  and  E.  L.  Cooper.     1962.     An 

ecological    comparison    of    six    populations    of 

brown     trout     (Salmo    trutta).     Trans.     Am. 

Fisheries  Soc.   91:53-62. 
McFadden,  J.  T..  E.  L.  Cooper,  and     J.   K.  An 

derson.      1965.     Some  effects  of  environment 

on    egg    production    in    brown    trout     (Salmo 

trutta).  Limn,   and  Ocean.   10(l):88-95. 
Parker.     R.     R.      1961     Effects    of    formalin     on 

length  and  weight  of  fishes.  J.  Fish.  Res.  Bd. 

Canada  20(6) : 1441-1455. 
Scott.    D.    P.     il962.     Effect   of   food   quantity   on 

fecundity  of  rainbow   trout.  Salmo  gairdneri. 

,J.  Fish.  Res.  Bd.  Canada  19:715-731. 
Snedecor,   G.    W..    and    W.    G.    Cochran.      1971. 

Statistical     methods.     Iowa     St.     Univ.     Press. 

Iowa.   593  pp. 
United       States       Geological       Survey.      1972. 

Water    resources    data    for    Montana.    Part    2. 

Water  ([uality   records.  218  pp. 


SOME  RELATIONSHIPS  BETWEEN  INTERNAL  PARASITES 
AND  BROWN  TROUT  FROM  MONTANA  STREAMS 

Lawrence  L.  Lockard^,  R.  Randall  Parsons^,  and  Barry  M.  Schaplow^ 

Abstr.'^ct. —  Forty-five  percent  of  306  brown  trout  from  16  Montana  streams  were  infected  with 
one  or  more  of  the  nematodes  Cystidicoloides  salvelini,  Bulbodacnids  globosa,  Rhabdochona  sp., 
and  Eustrongylides  sp.  The  relationships  between  incidence  and  intensity  of  nematode  infections 
and  age  and  sexual  maturity  of  the  host  fish  were  studied.  Generally,  se.xuallj'  mature  female  brown 
trout  had  a  higher  rate  of  infection  and  had  more  nematodes  per  infected  fish  than  immature  fe- 
male brown  ti'out.  Higher  incidence  and  intensity  of  infection  in  sexually  mature  fish  was  attributed 
to  more  aggressive  feeding  behavior  leading  to  more  exposure  to  the  intermediate  hosts  (mayflies) 
of  the  nematode  parasites. 


Nimierous  parasitological  surveys  of 
trout  in  North  America  have  been  re- 
ported; however,  few  have  included  in- 
formation on  the  parasites  of  brown  trout 
{salmo  trutta) .  Van  Cleave  and  Mueller 
(1934)  studied  13  brown  trout  from 
Oneida  Lake,  New  York,  and  3  from  one 
of  its  tributary  streams.  Fifty-eight  Wis- 
consin brown  trout  were  studied  by  Bang- 
ham  (1946)  and  Fischthal  (1947a,  1947b, 
1950,  and  1952),  with  54  originating 
from  streams.  In  the  western  geographic 
region  of  the  United  States  the  studies  of 
Bangham  (1951),  Hugghins  (1959),  Al- 
exander (1961),  and  Fox  (1962)  include 
information  on  the  parasites  of  brown 
trout.  In  these  studies  a  total  of  55  brown 
trout  were  examined:  51  from  lakes  and 
4  from  streams.  Heckmann  (1971)  ex- 
amined 28  brown  trout  from  Montana 
for  blood  parasitism. 

This  study  is  an  attempt  to  examine 
some  relationships  between  internal  para- 
sites, brown  trout,  and  the  fish's  stream 
habitat.  It  is  based  on  the  necropsy  of  306 
brown  trout  collected  from  17  sites  on  16 
streams  in  south  and  western  Montana. 
The  specimens  were  collected  in  early 
September  through  October  (1972  and 
1973)  just  prior  to  and  during  the  spawn- 
ing seasons.  According  to  Van  Cleave 
and  Mueller  (1934),  fish  are  most  heav- 
ily parasitized  in  the  summer  season  when 
they  are  most  actively  feeding.  Since  Fox 
(1962)  found  nematode  infections  in 
Montana  brown  and  rainbow^  trout 
(Salmo  gairdneri)  highest  in  August  and 
September,  these  findings  may  be  indica- 
tive of  near  maximum  parasite  infec- 
tions for  brown  trout  in  Montana  streams. 


Methods  and  Materials 

All  fish  were  collected  by  use  of  electro- 
fishing  gear  described  by  Vincent  (1967). 
Collecting  sites  were  selected  to  reflect  a 
wide  range  of  geographic  locations  (Fig. 
1),  and  physical  and  chemical  conditions 
(Table  1).  Collections  were  taken  on  both 
sides  of  the  Continental  Divide  from 
streams  belonging  to  the  Clark  Fork  of 
the  Coltmibia,  Yellowstone,  and  Missouri 
River  drainages.  At  least  one  fall,  winter, 
and  summer  measurement  of  conductivi- 
ty and  alkalinity  was  made  on  each 
stream.  These  field  measurements  were 
averaged  with  the  available  yearly  con- 
ductivity and  alkalinity  averages  ob- 
tained from  Water  Resources  Data  for 
Montana  (USGS  1972).  Discharge  values 
were  obtained  by  averaging  available 
yearly  values  from  the  above  USGS  rec- 
ords with  values  measured  or  estimated 
by  fisheries  biologists  of  the  Montana 
Department  of  Fish  and  Game.  The 
streams  sampled  varied  in  average  dis- 
charge from  5  cubic  feet  per  second  to 
1,409  cubic  feet  per  second,  in  average 
conductivity  from  70  to  1,387  micromhos 
/cm  at  25  C.,  and  in  alkalinity  from  49  to 
221  ppm  CaCO'.  Ecologically  the  streams 
are  diverse  although  they  all  support  nat- 
urally reproducing  populations  of  browoi 
trout. 

Immediately  after  capture  the  fish  were 
preserved  in  10  percent  formalin,  washed 
in  water  and  stored  in  40  percent  isopro- 
pyl  alcohol  for  later  examination.  Parker 
(1963)  indicates  fish  shrink  about  3-4 
percent  in  length  and  gain  between  5  and 
12  percent  in  weight  when  preserved  in 
formalin.  About  three  to  eight  months 
after    collection    fish    were    individually 


^Fish    and    Wildlife    Service,    U.S.    Department    of    the    Interior,    P.O.    Bo.\   250,    Picnc,    South    Dakota   57501. 


442 


Dec.  1975 


LOCKARD,   ET   AL.:    BROWN   TROUT 


443 


Fig.   1.  Map  showing  location  of  collecting  sites. 

Table  1.  Selected  physical  and  chemical  characteristics  of    16  Montana   streams   and  the  incidence 
and  intensity  of  nematode  infections  in  brown  trout  from  them. 


Stream 

Ave.# 

Collection 

dis- 

'  # 

% 

worms 

site  no.i 

Stream 

charge 

Cond. 

Alk. 

Fish 

Par. 

par. /fish 

1 

Madison  R. 

1409 

249 

107 

13 

100 

68.7 

2 

Big  Hole  R. 

1125 

207 

117 

19 

100 

14.3 

3 

W.  Gallatin  R. 

791 

230 

118 

17 

100 

21.7 

4 

St.  Regis  R. 

555 

80 

51 

22 

14 

1.7 

5 

Beaverhead  R. 

405 

521 

193 

14 

43 

1.7 

6 

E.  Gallatin  R. 

400* 

360 

195 

15 

87 

3.9 

7 

Rock  Cr. 

169 

70 

49 

16 

63 

3.4 

8 

Shields  R. 

159 

402 

221 

11 

18 

1.5 

9 

L.  Blackfoot  R. 

105 

612 

188 

28 

21 

1.2 

10 

O'Dell  Cr. 

100* 

348 

167 

11 

46 

3.0 

11 

So.  F.  Musselshell  R. 

83 

561 

243 

13 

8 

1.0 

12 

Baker  Cr. 

70* 

317 

154 

28 

39 

1.9 

13 

L.  Prickley  Pear  Cr. 

69 

358 

195 

21 

5 

1.0 

14 

16  Mile  Cr. 

50* 

522 

195 

13 

0 

0 

15 

Bluewater  Cr.- 

28 

1387 

214 

27 

56 

2.1 

16 

Bluewater  Cr.^ 

18 

798 

209 

22 

77 

3.7 

17 

Flagstaff  Cr. 

5* 

405 

197 

16 

0 

0 

♦Estimated 

by    fisheries    biologists    of    the 

Montana    Departm  en 

of   Fish 

md    Game. 

^See  Figur 

B  1 

=Section  below  Bluewater  Fish  Hatchery 

^Section  above  Bluewater  Fish  Hatchery 

measured  and  weighed,  and  scale  samples 
were  taken  for  age  determinations.  The 
ovaries  were  removed  for  an  unrelated 
fecundity  study  (Lockard  1974)  and  con- 
sequently were  not  included  in  this  nec- 
ropsy. Also,  the  heart  and  gas  hladder 
were  excluded  because  they  were  either 
mutilated  or  lost  when  the  ovaries  were 
removed.    The    procedure    for    necropsy 


was  as  follows.  The  gills  and  external 
body  of  the  fish  were  examined  for  ecto- 
jjarasites  and  fluke  metacercaria,  although 
the  preserving  and  handling  procedures 
greatly  reduced  the  chances  of  discovering 
j)arasitic  copepods,  leeches,  or  mono- 
genetic  flukes.  After  removal  of  the  re- 
maining internal  organs,  the  liver  was  dis- 
sected into  small  pieces,  placed  in  a  jar 


444 


GREAT   BASIN    NATURALIST 


Vol.  35.  No.  4 


with  water,  and  put  on  a  mechanical 
shaker  for  5-10  minutes.  The  contents 
were  then  washed  onto  a  200-mesh  col- 
lecting screen  and  examined  in  an  il- 
luminated tray  (Barber  and  Lockard 
1973).  The  gastrointestinal  tract  was 
opened  and  its  component  parts  scraped 
and  placed  with  their  contents  in  jars 
of  water.  After  5-10  minutes  on  the  me- 
chanical shaker,  the  contents  of  each  jar 
were  examined  in  the  illuminated  tray. 
The  kidney  and  testes  w'ere  examined 
grossly,  and  observed  abnormalities 
checked  microscopically.  Recovered  nema- 
todes were  placed  in  70  percent  alcohol-5 
percent  glycerine  and  later  mounted  in 
glycerine. 

Results 

Parasites  were  found  throughout  the 
upper  digestive  tract  from  the  esophagus 
to  the  pyloric  caeca.  The  examinations 
revealed  that  45  percent  of  the  306 
brown  trout  were  infected  with  one  to 
four  genera  of  nematodes.  Table  2  lists 
the  occurrence  of  each  taxon  in  a  given 
stream.  Identification  of  Cystidicoloides 
salvelini,  Bulbodacnitis  globosa,  and  Rhab- 
dochona  sp.  was  confirmed  by  Dr.  James 
R.  Adams,  University  of  British  Columbia 
(personal  correspondence) .  The  Rhabdo- 
chona  sp.  apparently  is  a  new  species 
and  has  been  submitted  to  Dr.  Morovec 
in  Prague,  Czechoslovakia,  for  further 
identification. 

Inspection  of  Table  1  indicated  an  ap- 
parent  relationship    between    the    size    of 


stream  discharges,  the  percent  parasitism 
(incidence)  in  fish  and  average  number 
of  worins  per  parasitized  fish  (intensity). 
Regression  analyses  were  performed  to 
test  these  relationships.  The  ])ercentage 
of  fish  parasitized  regressed  on  stream 
size  was  found  to  be  correlated  (R-  =  0.46; 
T  test  P=  0.003).  The  average  number  of 
nematodes  per  parasitized  fish  regressed 
on  stream  discharge  was  found  to  be  high- 
ly correlated  (R— 0:68;  .  T  test  P= 
0.00004) . 

The  306  fish  used  for  this  study  ranged 
from  6.4  to  20.7  inches  (16.3  to  52.6  cm) 
in  total  length  and  from  0.11  to  3.90 
pounds  (49.9  to  1769.0  grams)  in  weight 
with  the  mean  length  and  weight  being 
12.3  inches  (31.2  cm)  and  0.93  pounds 
(421.8  grams).  To  test  for  a  possible 
correlation  between  length  of  fish  and  in- 
tensity of  parasitism,  a  regression  of  num- 
ber of  nematodes  per  infected  fish  on 
fish  length  was  made  on  fish  from  the 
three  collections  with  the  highest  percent 
parasitism  (Table  1).  Fish  from  each 
stream  were  tested  individually  to  com- 
pensate for  varying  intensities  of  parasit- 
ism and  differences  in  mean  size  of  fish 
between  streams.  There  was  no  significant 
correlation  at  the  0.5  level  between  num- 
ber of  parasites  and  length  of  host  in  fish 
from  the  Madison  River,  Big  Hole  River, 
and  the  West  Gallatin  River. 

The  relationship  of  fish  age  to  the  in- 
cidence and  intensity  of  parasitism  is 
shown  in  Table  3.  Differences  in  the  per- 
cent of  infected  fish  between  age  classes 


Table  2.  Occurrence  of  nematode  genera  in  brown  trout  from   16  Montana  streams. 


Rhabdo- 

Bulbodac- 

Cystidi- 

Eustrong- 

ColL 

action  site 

chona 

nitis 

coloides 

ylides 

1 

(Madison  R.) 

X 

X 

X 

X 

2 

(Big  Hole  R.) 

X 

X 

X 

3 

(W.  Gallatin  R.) 

X 

X 

X 

4 

(St.  Regis  R.) 

X 

X 

5 

(Beaverhead  R.) 

X 

X 

6 

(E.  Gallatin  R.) 

X 

X 

7 

(Rock  Cr.) 

X 

X 

8 

(Shields  R.) 

X 

9 

(L.  Blackfoot  R.) 

X 

10 

(O'Dell  Cr.) 

X 

X 

11 

(So.  F.  Musselshell  R.) 

X 

12 

(Baker  Cr.) 

X 

X 

X 

13 

(L.  Prickley  Pear 

Cr.) 

X 

14 

(16  Mile  Cr.) 

15 

(Bluewater  Cr.)i 

X 

X 

16 

(Bluewater  Cr)- 

X 

X 

17 

(Flagstaff  Cr.) 

'Section  above  Bluewater  Fish  Hatchery 
-Section  below  Bluewater  Fish  Hatchery 


Dec.  1975 


LOCKARD,   ET   AL.:    BROWN   TROUT 


445 


Table  3.   Incidence  and  intensity  of  parasitism 
in  age  classes  of  brown  trout. 


Age  class 


# 
Fish 


Infected 


Ave.  #  worms/ 
infected  fish 


I  58 

II  137 

III+  111 

Total  (ave.)  306 


50 
38 
62 

(45) 


2.4 

9.1 

21.3 

(12.8) 


were  tested  by  a  method  of  Arkin  and 
Colton  (1972)  with  results  showing  no 
significant  differences  between  age  class- 
es I  and  II,  and  I  and  III+  at  the  0.05 
level.  Although  the  difference  in  the  per- 
cent of  fish  infected  in  age  classes  II  and 
III+  was  statistically  significant  (P  = 
0.028)  it  was  not  considered  to  be  bio- 
logically significant.  As  the  age  of  the 
fish  increased  the  average  number  of 
worms  per  infected  fish  also  increased 
(Table  3).  These  differences  in  the  in- 
tensity of  parasitism  between  age  classes 
shown  in  Table  3  were  significant  at  the 
0.05  level. 

The  relationship  of  incidence  and  inten- 
sity of  parasitism  to  age  and  sexual  ma- 
turity in  brown  trout  is  given  in  Table 
4.  Significantly  (0.05  level)  more  mature 
fish  than  immature  fish  in  age  classes  I 
and  II  and  in  the  overall  total  were  in- 
fected. However,  there  was  no  significant 
difference  between  the  incidence  of  para- 
sitism in  mature  and  immature  fish  in 
age  class  III  +  .  Mature  fish  in  age  classes 
II  and  III+  and  the  overall  total  also  had 
a  significant!}'  higher  number  of  worms 
per  fish  than  immature  fish.  In  age  class  I 
there  were  no  significant  differences  in 
the  average  number  of  worms  in  mature 
and  immature  brown  trout. 

Tests  were  made  to  determine  the  re- 
lationship of  the  chemical  productivity 
of  each  stream  with  the  degree  of  parasit- 
ism in  its  fish.  No  significant  (0.05  level) 
correlations  were  found  when  percent 
parasitism  and  average  number  of  worms 


per    parasitized    fish    were    regressed    on 
stream  conductivities  and  alkalinities. 

In  the  five  streams  (Big  Hole  River, 
Baker  Creek,  Little  Prickley  Pear  Creek, 
Bluewater  Creek,  and  Flagstaff  Creek) 
from  which  the  48  males  were  collected, 
there  was  no  significant  difference  be- 
tween their  rate  of  infection  (45.8  per- 
cent) and  that  of  females  from  the  same 
streams  (48.2  percent). 

Discussion 

Incidence  of  parasitism  in  brown  trout 
varies  with  species  of  parasite  and  with 
habitat  of  the  host  fish.  Fox  (1962)  found 
30  brown  trout  from  Meadow  Lake,  Mon- 
tana, infected  with  fluke  metacercaria  of 
Bolbophorus  confusus.  Five  of  the  30 
brown  trout  had  immature  stages  of  the 
nematode  Eustrogylides  sp.  encysted  with- 
in the  body  and  1  of  the  30  had  an  un- 
identified nematode  in  its  digestive  tract. 
Van  Cleave  and  Mueller  (1934)  examined 
13  brown  trout  from  Oneida  Lake  and 
3  collected  from  Black  Creek  about  one 
mile  from  its  confluence  with  Oneida 
Lake,  New  York.  The  stream-inhabiting 
brown  trout  had  the  nematode  Cystidi- 
coloides  hardwoodi  exclusively,  whereas 
the  lake-inhabiting  brown  trout  had  the 
nematode  Spinitectus  gracilis  and  never 
C.  hardwoodi.  In  over  1,000  fish  from  the 
lake  C.  hardwoodi  was  never  taken;  thus 
it  was  probably  limited  to  streams. 

In  correlating  parasitism  to  habitat.  Van 
Cleave  and  Mueller  (1934)  conclude 
"that  in  fishes  of  wide  range  through  a 
variety  of  environmental  types,  the  type 
of  parasitism  is  more  or  less  closely  cor- 
related with  the  habitat  from  which  the 
fish  is  taken."  Bangham  (1951)  states 
"the  fact  that  there  were  not  many  dif- 
ferent types  of  habitats  limited  the  num- 
bers of  various  parasite  species."  Stream 
habitat  may  not  favor  the  intermediate 
hosts  of  many  lake  parasites,  i.e.,  cestodes 


T.\BLE  4.  Incidence  and  intensitj-  of  parasitism  related  to  sexual  maturity  and  age  of  brown  trout. 


Ave. 

#  worms/ 

Age 

# 
Fish 

Se 

X.   Mat. 

"Ji 

Infected 

infected 

fish 

class 

Se 

X.  mat. 

Sex.  imm. 

Sex.    mat. 

Sex.    imm. 

I 

58 

19 

82 

43 

2.1 

2.5 

II 

137 

51 

47 

28 

11.0 

5.9 

III  + 

111 

88 

52 

54 

23.7 

4.4 

Total 

306 

(60) 

(53) 

(34) 

(17.1) 

(4.2) 

(Ave.) 

446 


GREAT    BASIN    NATURALIST 


Vol.  35,  No.  4 


(copepods),  trematodes  (snails),  acantho- 
cephalans  ( Crustacea).  Parasitic  cope- 
pods  and  monogenetic  flukes  not  com- 
mon in  streams  may  be  favored  by  the 
lake  environment.  1'his  would  partially 
explain  wh}'  nematodes  were  the  only 
parasites  found  in  these  brown  trout 
from  cold  water  stream  habitats. 

Hugghins  (1959)  found  no  parasitism 
in  four  brown  trout  from  Black  Hills 
streams  in  South  Dakota.  Van  Cleave 
and  Mueller  (1934)  classified  the  fre- 
quency of  occurrence  of  six  species  of 
parasites  in  brown  trout  as  "occasional" 
based  on  a  scale  of  abundant,  common,  oc- 
casional, and  rare.  Forty-one  percent  of 
58  brown  trout  from  Wisconsin  streams 
and  ponds  were  parasitized  (Bangham 
1946,  and  Fischthal  1947a,  1947b,  1950, 
1952).  This  is  comparable  to  the  45  per- 
cent incidence  of  parasitism  found  in  this 
5tudy. 

The  correlation  of  size  of  stream  with 
incidence  and  intensity  of  parasitism  was 
shown  in  Table  1.  It  can  be  seen  from 
Table  2  that  large  streams  have  a  greater 
diversity  of  species  of  nematodes  than 
small  streams.  It  would  be  expected  in  a 
large  river  with  diverse  habitat  niches 
that  the  chance  of  intermediate  and  de- 
finitive hosts  for  a  certain  parasite  exist- 
ing together  would  be  greater  than  in  a 
smaller  stream  with  a  more  restricted 
type  of  habitat. 

Fox  (1962)  states  that,  in  general,  the 
longer  the  trout,  the  greater  the  number 
of  individual  parasites  and  kinds  of  para- 
sites it  contained.  This  is  related  to  longer 
exposure  time  to  parasitism,  since  the 
body  length  is  generally  determined  by 
age.  Woodbury  (1940)  summarized  12 
previous  investigations  and  reported  that 
the  longer  fish  had  greater  numbers  of 
parasites  in  nine  of  these  studies.  Con- 
versely, Langlois  (1936)  and  Hubbs 
(1927)  found  shorter  fish  the  most  heavi- 
ly parasitized.  In  this  study  no  correlation 
between  fish  length  and  intensit}^  of  in- 
fection was  found  in  fish  from  the  three 
streams  with  100  percent  })arasitism.  How- 
ever, the  fish  in  these  streams  were  not 
proportionately  representative  of  all  age 
classes;  thus  these  results  do  not  rule  out 
a  relationship  between  age  and  intensity 
of  infection.  This  study  did  not  show^  that 
older  fish  were  generally  more  likely  to 
be  infected  than  younger  fish  (Table  3). 


However,  it  can  be  seen  from  Table  3 
that  as  the  age  of  the  fish  increases,  so 
does  the  average  number  of  worms  per  in- 
fected fish.  This  could  be  related  to  the 
older  fish  having  more  exposure  time  to 
})arasites  than  the  younger  ones  do. 

The  relationships  of  incidence  and  in- 
tensity of  infection  to  feeding  aggressive- 
ness of  fish  as  expressed  by  sexual  ma- 
turity was  examined.  A  higher  percentage 
of  mature  fish  was  parasitized  than  im- 
mature fish  in  the  overall  total  and  for 
age  classes  I  and  II  (Table  4).  The  dif- 
ference between  infection  rates  between 
mature  and  immature  fish  in  the  overall 
total  could  be  attributed  to  older  mature 
fish  having  more  exposure  time  to  the 
parasites  than  younger  fish  do.  However, 
the  differences  in  infection  rates  between 
mature  and  immature  fish  within  age 
classes  I  and  II  could  be  the  result  of  the 
fastest-growing  (most-aggressive)  fish  in 
an  age  class  reaching  sexual  maturity  be- 
fore less-aggressive  fish.  Bagenal  (1969) 
fed  different  quantities  of  food  to  two 
groups  of  brown  trout  and  found  that  the 
more  aggressive  of  the  under-fed  fish  took 
more  than  their  share  of  the  food.  Conse- 
quently, the  more  aggressive  of  the 
starved  fish  had  growth  comparable  with 
that  of  the  well-fed  fish.  He  also  found 
that  more  of  the  better-fed  fish  were  ma- 
ture than  the  under-fed  fish.  There  were 
no  significant  differences  (0.05  level)  be- 
tween mature  and  immature  fish  in  the 
percent  infected  for  age  class  III  +  .  Niel- 
son  (1953)  noted  that  brown  trout  in  Cali- 
fornia began  a  shift  in  feeding  habits 
from  strictly  aquatic  invertebrate  forms 
to  partial  diets  of  fish  in  the  third  year 
of  life.  An  explanation  for  the  similar  per- 
cent infections  for  mature  and  immature 
age  class  III+  fish  (Table  4)  could  be  this 
change  in  feeding  habits  in  which  the 
parasite  intermediate  hosts  (mayflies) 
would  be  preyed  upon  less  by  the  larger 
mature  fish. 

McFadden,  Cooper,  and  Anderson 
(1965)  state  that  perhaps  some  combin- 
ation of  chronological  age  and  growth 
rate  (or  the  factors  which  determine 
growth  rate),  a  "physiological  age",  de- 
termines the  age  of  sexual  maturity.  One 
factor  which  influences  growth  rate  and 
thus  age  at  sexual  maturity  is  the  feed- 
ing aggressiveness  of  a  fish.  If  sexually 
mature  fish  feed  more  actively  than  sex- 


Dec.  1975 


LOCKARD.    ET   AL.:    BROWN    TROUT 


447 


ually  immature  fish,  then  mature  fish  of 
a  given  age  would  be  larger  than  the  im- 
mature fish  in  that  age  class  and  would 
have  more  exposure  to  parasitism  through 
their  aggressive  feeding  habits.  The  fol- 
lowing is  a  comparison  of  mean  lengths 
of  mature  and  inmiature  female  brown 
trout  within  age  classes.  In  age  class  I, 
only  the  Bluewater  Creek  collections  con- 
tained mature  females,  and  they  had  a 
mean  length  of  8.0  inches  while  the  im- 
mature females  from  Bluewater  Creek 
were  7.5  inches  in  mean  length.  Mature 
female  fish  from  all  streams  of  age  class- 
es II  and  111+  had  mean  lengths  of  13.3 
and  15.1  inches,  respectively;  while  the 
immature  females  from  these  age  classes 
had  mean  lengths  of  11.0  and  12.9 
inches,  respectively.  Thus  sexually  ma- 
ture fish  v\dthin  an  age  class  are  larger 
than  sexualh'  immature  fish.  This  could 
be  the  result  of  more  aggressive  feeding 
habits.  The  most  actively  feeding  fish 
would  have  more  chances  to  encounter 
the  intermediate  hosts  (mayflies,  Hoff- 
man 1967)  of  these  nematode  parasites 
{Cystidicoloides  sp.  and  Rhabdochona  sp. 
were  99.4  percent  of  all  worms  found) 
and  thus  more  of  them  would  be  infected. 

This  theory  to  explain  differences  in 
infection  rates  between  mature  and  im- 
mature fish  within  age  classes  is  rein- 
forced by  inspection  of  the  average  num- 
ber of  worms  per  parasitized  fish  in  age 
classes  (Table  4).  The  mature  fish  in  age 
classes  II  and  III+  and  for  the  overall 
total  have  a  higher  average  number  of 
worms  per  fish  than  the  immature  fish. 
This  suggests  that  the  more  actively  feed- 
ing mature  fish  within  an  age  class  are 
exposed  more  to  parasitism  by  their  feed- 
ing habits  than  less  actively  feeding  im- 
mature fish. 

In  age  class  I  (Table  4)  there  was  no 
significant  difference  (0.05  level)  be- 
tween mature  and  immature  brown  trout 
in  the  average  number  of  worms  j)er  in- 
fected fish,  possibly  because  exjiosure  time 
was  too  short  for  significant  differences 
to  be  expressed . 

Fox  (1962)  found  little  relationship  be- 
tween incidence  of  parasitism  and  con- 
dition of  trout.  He  states  that  possibly 
".  .  .  analysis  of  fecundity  will  show  that 
parasite  incidence  does  effect  some  host 
conditions."  Comparison  of  the  findings 
of  this   study   with   those   of  a   fecundit} 


study  on  the  same  fish  (Lockard  1974) 
indicates  that  no  general  relationship  be- 
tween parasitism  and  fecundity  exists. 
Table  1  shows  the  Madison  and  West 
Gallatin  rivers  were  highly  parasitized; 
in  the  above  fecundity  study  they  had  the 
highest  fecundity  (steepest  slopes  of  the 
regressions  of  number  of  eggs  on  fish 
length  for  a  stream) .  Thus  parasitism  ap- 
parently did  not  adversely  affect  the  fe- 
cundity of  infected  fish. 

Acknowledgments 

The  authors  express  their  appreciation 
to  Dr.  Da^'id  E.  Worley  of  Montana  State 
University  for  providing  laborator}"  space 
and  equipment,  and  to  Dr.  William  R. 
Gould  of  Montana  State  University  and 
Mr.  George  D.  Holton  of  the  Montana 
Department  of  Fish  and  Game  for  re- 
viewing the  manuscript. 


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Arkin,  H..  .-^nd  R.  R.  Colton.  1970.  Statisti- 
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SEXUAL  DIMORPHISM  IN  MALPIGHIAN  TUBULES  OF 
PTERONARCYS  CALIFORNICA  NEWPORT  (PLECOPTERA) 

Ralph  R.  Hathaway  1 

Abstr.'VCt. —  The  Malpighian  tubules  in  female  nymphs  and  adults  of  Pteronarcys  californica  New- 
port are  larger  and  whiter  than  in  males.  This  difference  is  detectable  in  nymphs  as  small  as  300 
mg  in  live  weight  (final  instar  nymphs  weigh  800  to  1600  mg  depending  on  sex),  and  is  most  pro- 
nounced in  late  nymphs  and  adults.  These  differences  are  thought  to  be  related  to  the  increased  ex- 
cretory load  imposed  by  the  synthesis  of  ooplasm  in  the  late  female  nympli  and  adult. 


The  excretory  systems  of  Plecoptera 
reflect  the  relatively  primitive  taxonomic 
level  of  this  order.  The  Malpighian  tu- 
bules are  usually  numerous  and  un- 
branched  and  have  an  uncomplicated  re- 
lationship to  the  gut,  joining  it  at  the  level 
where  the  midgut  empties  into  the  hind- 
gut.  Apart  from  this  connection,  the  tu- 
bules are  mainly  free  and  unattached,  ex- 
tending throughout  the  body  cavity.  This 
situation  is  seen  with  textbook  clarity  in 
Pteronarcys  californica. 

Much  current  interest  in  the  Malpi- 
ghian tubules  centers  on  their  ultra  struc- 
ture and  on  their  function  as  a  homeo- 
static  organ  (Berridge  and  Oschman  1969; 
Phillips  and  Maddrell  1974).  They  are 
involved  in  the  elimination  of  salts  and 
amines,  and  in  water  balance.  Most 
studies  have  been  done  on  terrestrial 
forms  in  which  water  retention  and  ion 
transport  are  achieved  in  specializations 
of  the  tubules  and,  at  times,  inclusion  of 
tubules  in  the  wall  of  the  hind  gut.  The 
only  studies  on  excretion  in  Plecoptera 
are  those  of  Colby  (1972),  who  used 
in  vivo  measurements  to  demonstrate  the 
formation  of  a  dilute  urine  in  the  aquatic 
nymphs  of  Pteronarcys  californica.  There 
have  been  no  detailed  morphological  stud- 
ies of  Plecopteran  Malpighian  tubules. 

In  a  study  of  the  growth  of  internal 
organs  (Branham  and  Hathaway  1975), 
I  have  observed  that  there  is  a  distinct 
sexual  dimorphism  in  the  appearance  of 
Malpighian  tubules  in  P.  californica, 
which  has  not  to  my  knowledge  been 
reported  earlier. 

Methods 

Stoneflies  were  collected  in  the  Provo 
River  in  Utah  and  transported  alive  to 
the  laboratory.  Several  hundred  fresh  and 
preserved   animals   from   all    size   classes 

^Biology  Department,   University  of  Utah,   Salt   Lake  City,   Utah 


were  examined.  The  dimorphisms  were 
generally  well  preserved  in  animals  fixed 
in  70  percent  ethanol  and  other  killing 
and  fixing  fluids.  A  few  animals  were 
used  for  histological  preparations  in  which 
10/t  sections  were  stained  with  Delafields 
hematoxylin.  Weights  of  live  animals 
were  taken  on  an  analytical  balance  after 
thorough  blotting  to  remove  excess  water. 

Observations 

Male  nymphs  in  the  final  instars  weigh, 
on  the  average,  about  800  mg,  whereas 
the  corresponding  female  nymphs  are 
about  twice  as  big,  or  1600  mg.  The  Mal- 
pighian tubules  in  the  female  nymphs 
in  all  size  classes  above  300  mg  are  larger 
and  whiter  than  those  in  the  males.  This 
difference  becomes  more  pronounced  as 
the  animals  grow  larger  and  is  at  its  maxi- 
mum in  the  final  instars  and  adults  (com- 
pare Figs,  la  &  c).  The  differences  are 
not  obvious  in  animals  weighing  less  than 
300  mg. 

It  is  at  about  300  mg  of  body  weight 
that  the  body-size  differences  between 
male  and  female  nymphs  becomes  dis- 
cernable;  therefore  the  onset  of  differ- 
ences in  the  Malpighian  tubules  corre- 
sponds with  this  event.  It  is  not  merely 
size,  however,  that  is  involved  in  the  sex- 
ual dimorphism  in  the  Malpighian  tu- 
bules. In  females  the  tubules  are  boldly 
conspicuous  because  of  their  intense 
whiteness,  whereas  the  male  tubules  are 
at  first  difficult  to  find  because  of  their 
translucent,  colorless  or  pink-tinged  ap- 
pearance. The  female  tubules,  moreover, 
are  of  more  uniform  diameter  throughout 
their  length,  while  the  diameter  of  male 
tubules  is  irregular.  Females  that  are 
smaller  and  at  least  a  year  yoimger  than 
final  instar  males  still  have  tubules  that 


449 


450 


GREAT   BASIN   NATURALIST 


Vol.  35,  No.  4 


are  larger  and  whiter  (compare  Figs,  la 
&b). 

About  20  percent  of  the  tubules  in  ani- 
mals of  both   sexes   are  connected   ante- 


riorly to  the  midgut  by  their  distal  ends 
(Fig.  la).  This  union  is  formed  from  a 
delicate  strand  of  solid  connective  tissue, 
so  there  is  no  confluence  at  this  point  be- 


Fig.  1:  Malpighian  tubules  in  Pteronarcys  californica:    (a)  Male  final-year  njrmph.  Live  wt.   =  840 

mg.  Isolated  gut  with  anterior  end  down.  Some  of  the  tubules  are  anchored  to  the  gut  by  their  dis- 
tal ends.  Scale  bar  equals  2  mm.  (b)  Female  penultimate-year  nymph.  Live  wt.  =  620  mg.  Isolated 
gut  with  anterior  end  down.  Tubules  are  larger  and  whiter  than  in  (a).  Anchored  distal  ends  of  some 
of  the  tubules  are  again  visible.  Magnification  same  as  in  (a),  (c)  Female  final-year  nymph.  Live 
wt.  =  1560  mg.  Isolated  gut  with  anterior  end  down.  Magnification  same  as  in  (a)  and  (b).  (d)  Fe- 
male final-year  n>-mph.  Cross  sections  of  tubules.  Scale  bar  =  0.1  mm.  (e)  Male  final-year  nymph. 
Cross  sections  of  tubules.   Magnification  is  same  as  in   (d). 


Dec.  1975 


HATHAWAY:    SEXUAL  DIMORPHISM    IN    PLECOPTERA 


451 


tween  the  lumina  of  the  gut  and  the  tu- 
bules. The  lumina  of  these  tubules  drain 
into  the  gut  at  the  same  level  as  all  the 
other  tubules,  i.e.  at  the  point  of  junction 
between  the  midgut  and  the  hindgut.  Be- 
tween their  proximal  and  distal  ends, 
these  tubules  are  totally  free  of  the  gut 
wall;  the  only  obvious  difference  from  the 
majority  of  the  Malpighian  tubules  is 
that  their  distal  ends  are  anchored  to  the 
midgut  at  a  point  about  two-thirds  of  the 
way  from  its  anterior  end,  while  the  dis- 
tal ends  of  the  other  tubules  are  free. 

Discussion 

The  differences  described  above  must 
have  a  functional  basis  relating  to  sex. 
Gonadal  development  can  be  seen  in  both 
sexes  as  they  begin  their  final  year  of 
nymphal  life.  At  this  time  males  weigh 
about  500  mg  and  females  weigh  900  to 
1000  mg.  Gonaflal  enlargement  and 
sperm  production  are  initiated  early  in 
the  male;  maximum  gonadal  size  is 
reached  midwa\'  through  the  final  year. 
Corresponding  events  in  the  female  do  not 
occur  until  the  final  six  or  eight  weeks 
of  nymphal  life.  During  this  period  a 
large  part  of  metabolic  effort  in  females 
is  directed  toward  the  synthesis  of  oo- 
plasm. There  is  evidence  that  oogene- 
sis utilizes  substances  from  the  fat  body 
(Branham  and  Hathaway  1972).  This 
effort  continues  into  the  adult  stage,  in 
which  gamete  production  in  the  ovaries 
can  take  place  for  at  least  four  weeks. 
Thus,  one  of  the  more  obvious  explana- 


tions for  sexual  differences  in  Malpighian 
tubules  is  that  the  redirection  of  metabo- 
lism associated  with  egg  production  re- 
el uires  more  elaborate  excretory  organs. 

Another  ])ossibility  is  that  the  Malpi- 
ghian tubules  in  the  female  are  involved 
in  a  secondary  function  such  as  the  stor- 
age of  energy-rich  substances.  The  pri- 
mary storage  organ  is  the  fat  body,  but 
this  decreases  in  size  during  the  time  of 
egg  production.  Thus  the  tubules  may 
serve  to  store  additional  reserves  to  be 
utilized  during  the  non-feeding  late- 
nymph  and  adult  stages. 

It  seems  likely  that  the  observations  re- 
corded here  have  some  general  signifi- 
cance, especially  in  Plecoptera.  Sexual  di- 
morphism in  Malpighian  tubules  also  oc- 
curs in  Pteronarcella  hadia  (Pteronarci- 
dae),  which  differs  in  many  life  historical 
respects  from  P.  calif ornica.  Further  ob- 
servations on  other  species  would  prob- 
ably be  rewarding. 

Literature  Cited 

BeRIUDGE.     M.     .]..    AND    ,1.     L.     OSCHMAN.        1969.       A 

structural  basis  for  fluid  secretion  by  Mal- 
pigbian  tubules.   Tissue   and  Cell   1:247-272. 

BiiANHAM,  .r.  M..  AND  R.  R.  Hathaw.xy.  1975. 
Sexual  differences  in  the  growth  of  Pteron- 
arcys  californica  Newport  and  Pteronarcella 
badia  (Hagen)  (Plecoptera).  Can.  J.  Zool. 
5^:501-506. 

Coi.BY.  C.  1972.  Salt  and  water  ijalance  in 
stoneflv  naiads.  Pteronarcys  californica  New- 
j)ort.   Comp.   Biochem.   Physiol.   41:851-860. 

Piiii.LiPS,  .J.  E.,  AND  S.  H.  P.  Maddrell.  1974. 
Active  transport  of  magnesium  by  the  Mal- 
pighian tubules  of  the  larvae  of  the  mosquito, 
Aedes  campeslris.  J.  Exp.   Biol.  61:76-771. 


NEW  RECORD  OF  THE  BAT  PLECOTUS  PHYLLOTIS 
FROM  UTAH 

Richard  M.   Poche' 


Abstract. —  In  June  1974  a  scrotal  male  Mexican  big-eared  bat  [Plecotus  phyllotis)  was  netted 
near  Hurricane.  Utah.  This  account  represents  the  third  report  of  the  species  from  Utah  and  e.x- 
tends  the  range  approximately  330  km  west  of  previous  accounts.  Adjacent  state  records  are  located 
approximately    64   km    to    the    southeast    in    Arizona    and   80   km   southwest   in   Nevada. 


The  Mexican  big-eared  bat  has  been  re- 
ported from  Utah  by  Black  (1970)  and 
Armstrong  (1974).  Both  records  were 
from  San  Juan  County  about  8  km  N 
Blanding  and  65  km  NW  Monticello,  re- 
spectively. A  specimen  was  reported 
from  Pipe  Springs  National  Monument 
area  in  Arizona  by  Genoways  and  Jones 
(1967  ).  and  one  from  the  Mesquite,  Neva- 
da, region  by  M.  J.  O'Farrell  (personal 
communication). 

On  25  June  1974  a  scrotal  male  Mexi- 
can big-eared  bat  was  obtained  in  <;  mist 
net  over  Gould  Wash,  NW  ^4,  SW 
14,  sec.  19,  T.  42  S,  R.  12  W,  Washington 
Co.,  Utah.  The  locality  lies  approximateh' 
7  air  km  southeast  of  the  town  of  Hurri- 
cane. The  capture  of  this  speci.men  ex- 
tends the  range  in  Utah  approximately 
330  km  to  the  west  and  represents  only 
the  third  report  from  that  state.  The 
Arizona  report  is  from  an  area  about  64 
km  southeast  of  Hurricane  and  the  Ne- 
vada account  80  km  to  the  southwest. 
General  similarity  in  habitat  in  the  area 
between  Hurricane  and  the  southeastern 
reports  suggests  that  P.  phyllotis  probably 
inhabits    most    of    southern    Utah. 

The  Mexican  big-eared  bat  obtained 
near  Hurricane,  Utah,  was  netted  over 
one  of  three  small  ponds  remaining  in 
the  wash,  striking  the  net  approximately 
1  m  above  the  surface  of  the  water  be- 
tween 0100  and  0530  hours.  It  was  ex- 
amined for  ectoparasites,  of  which  there 
were  none,  marked  (with  a  3  mm  hole 
in  the  plagiopatagium,  near  the  pollex, 
between  the  4th  and  5th  phalanges  of 
the  right  wing),  and  released.  The  bat 
flew    west    along    the    wash    and    disap- 

'School    o{    Forcsty    &    Cnnscrvnlinn.     University    of    Cnlifom  i 


peared  into  the  canyon.  Six  male  scrotal 
Pipistrellus  hesperus  also  were  captured 
in  the  same  net  and  were  released. 

Gould  Wash  drains  into  the  Virgin 
River  14  km  northwest  of  the  netting 
site.  Vegetation  in  the  wash  includes 
Tamarix  and  Salix.  and  that  on  the  ad- 
jacent upland  is  comprised  primarily  of 
blackbrush  (Coleogyne  ramosissima) ,  and 
infrequent  pinon  pine  {Pinus  edulis)  and 
juniper  [Juniperus  osteosperma) .  Grass 
cover  is  predominantly  Bromus  and  Hi- 
laria.  Average  elevation  for  the  area  is 
1,250  m  with  annual  precipitation  less 
than  200  mm.  The  Hurricane  Cliffs  are 
located  5  km  west  of  the  collection  area 
and  descend  over  500  m  to  Warner  Val- 
ley, which  is  covered  with  creosotebush 
{Larrea  tridentata) ,  blackbrush,  and 
snakeweed  {Gutierrezia  microcephala) . 
The  surrounding  region  consists  of  num- 
erous steep  rises  and  mesas,  canyons,  and 
cliffs. 

I  thank  Geoff  Baillie  for  assistance  with 
netting  activities.  David  Armstrong  pro- 
vided helpful  comments  in  reviewing  this 
paper.  This  study  was  financed  by  the 
Nevada   Power   Company. 

LiTF,R.'\TURE  Cited 

Armstrong.  D.  A.  1974.  Second  record  of  the 
Mexican  big-eared  bat  in  Utah.  Southwest- 
ern  Nat.    19(1):114-115. 

Black.  H.  L.  1970.  Occurrence  of  the  Mexican 
big-eared  bat  in  l^tah.  J.  Mammal.  15(1); 
190. 

Genoways.  H.  H..  and  J.  K.  Jones.  Jr.  1967. 
Notes  on  distribution  and  variation  in  the 
Mexican  big-eared  bat.  Plecotus  phyllotis. 
Southwestern   Nat..    12:477-480, 


Beikcle 


047nj, 


452 


INDEX    TO    VOLUME    35 

The  genera  and  species  described  as  new  to  science  in  tiiis  volume  appear  in  bold  type 
in  this  index. 


Additional  records   of  reptiles   from   Jalisco, 

Mexico,  p.  317. 
Allred,  Dorald  M.,  article  by,  p.  405. 
Amphicranus  parilis,  p.  31. 
Andersen,    Ferron    L.,    and    Paul    R.    Roper, 

article  by,  p.  203. 
A  new  combination  of  Penstemon   (Scrophu- 

lariaceae),  p.  434. 
Arachnids  as  ecological  indicators,  p.  405. 
Araptiis  attenuatus,  p.  30. 
Araptus  consobrinus,  p.  394. 
Araptus  fossifrons,  p.  30. 
Araptus  micaceus,  p.  395. 
A  revision  of  the  nearctic  species  of  Clino- 
helea    Kieffer    (Diptera:    Ceratopogonidae), 
p.  275. 
A  revision  of  the  Phacelia  Crenulatae  group 
( Hydrophyllaceae )  for  North  America,  p.  127. 
Arndt,    Rudolf    G.,    Philip    A.    Medica,    and 

James  R.  Dixon,  article  by,  p.  317. 
A  systematic  study  of  Coenia  and  Paracoenia 

(Diptera:  Ephydridae),  p.  65. 
Atwood,  N.  Duane,  article  by,  p.  127. 
Atwood,    N.    Duane,    Stanley   L.    Welsh,    and 

James  L.  Reveal,  article  by,  p.  327. 
Avery,  David  F.,  Charles  Fanghella,  and  Wil- 

mer  W.  Tanner,  artcile  by,  p.  245. 
Baker,  M.  F.,  D.  W.  Nichols,  and  H.  D.  Smith, 

article  by,  p.  191. 
Basidomycetes  that  decay  junipers  in  Arizona, 

p.  288. 
Bleich,  Vernon  C,  and  Orlando  A.  Schwartz, 

article  by,  p.  62. 
Bock,  Carl  E.,  and  Larry  W.  Lepthien,  article 

by,  p.  269. 
Body  size,  organ  size,  and  sex  ratios  in  adult 
and  yearling   Belding  ground  squirrels,   p. 
305. 
Bombylius  abdominalis,  p.  414. 
Bombylius  aestivus,  p.  416. 
Bombylius  auriferoides,  p.  417. 
Bombylius  heximaculatus,  p.  410. 
Bombylius  lancifer  kanabensis,  p.  412. 
Bombylius  lassenensis,  p.  416. 
Bombylius  montanus,  p.  413. 
Bombylius  nigriventris,  p.  410. 
Borchert,   Mark,   and  Donald  H.   Owings,   p. 

402. 
Breeding  range  expansion  of  the  starling  in 

Utah,  p.  419. 
Branham,  Joseph  M.,  Arden  R.  Gaufin,  and 

Robbin  L.  Traver,  article  by,  p.  51. 
Calocoenia  (subgenus),  p.  78. 
Campbell,  R.  B.,  W.  T.  McDonough,  and  R.  0. 
Harniss,  article  by,  p.  325. 


Gather,  Mary  R.,  and  Arden  R.  Gaufin,  article 

by,  p.  39. 
Gather,  Mary  R.,  Bill  P.  Stark,  and  Arden  R. 

Gaufin,  article  by,  p.  49. 
Clarence    Cottam,    1899-1974,    a    distinguished 
alumnus  of  Brigham  Young  University,  p. 
231. 
Clark,  Stephen  L.,  article  by,  p.  434. 
Cliriohelea  pseudonubifera,  p.  280. 
Cnesinus  electus,  p.  23. 
Cnesinus  pilatus,  p.  24. 
Coenia  alpina,  p.  83. 

Computerized      reduction      of      meteorologic 
measurements   from   irrigated   and   nonirri- 
gated  plots  in  central  Utah,  p.  203. 
Correlates    of    burrow    location    in    Beechey 

ground  squirrels,   p.  402. 
Corthylus  cecropil,  p.  31. 
Cymopterus  higginsii,  p.  377. 
Dacnophthorus,  p.  394. 

Distribution  and  abundance  of  the  black-billed 
magpie   {Pica  pica)   in  North  America,  p. 
269. 
Dixon,    James    R.,    Philip    A.    Medica,    and 

Rudolf  G.  Arndt,  article  by,  p.  317. 
Endangered,  threatened,  extinct,  endemic,  and 

rare  or  restricted  Utah  plants,  p.  327. 
Environmental  factors  in  relation  to  the  salt 
content  of  Salicornia  pacifica  var.  utahensis, 
p.  86. 
Evans,  Howard  E.,  article  by,  p.  123. 
Evolutionary  divergence  in  closely  related  pop- 
ulations of  Mimulus  guttatus   (Scrophulari- 
aceae),  p.  240. 
Evolution  of  the  sceloporine  lizards   (Iguani- 

dae),  p.  1. 
Fanghella,  Charles,  David  F.  Avery,  and  Wil- 

mer  W.  Tanner,  article  by,  p.  245. 
Gallup,  John  S.,  and  Martin  L.  Morton,  article 

by,  p.  427. 
Gaufin,  Arden  R.,  and  Mary  R.  Gather,  article 

by,  p.  39. 
Gaufin,  Arden  R.,  Bill  P.  Stark,  and  Theodore 

A.  Wolff,  article  by,  p.  97. 
Gaufin,  Arden  R.,  Joseph  M.  Branham,  and 

Robbin  L.  Traver,  article  by,  p.  51. 
Genetics,    environment,    and    subspecies    dif- 
ferences: The  case  of  Polites  sabuleti,  p.  33. 
Gilbertson,  R.  L.,  and  J.  P.  Lindsey,  article 

by,  p.  288. 
Grogan,  WilHam  L.,  Jr.,  and  Willis  W.  Wirth, 

article  by,  p.  275. 
Growth   of   Plecoptera   (stonefly)    nymphs   at 
constant,   abnormally  high  temperature,  p. 
51. 


453 


454 


GREAT   BASIN    NATURALIST 


Vol.  35,  No.  4 


Guibe,  Jean,  Hobart  M.  Smith,  and  Rozella 
B.  Smith,  article  by,  p.  109. 

Hansen,  D.  J.,  and  D.  J.  Weber,  article  by, 
p.  86. 

Harniss,  R.  0.,  W.  T.  McDonough,  and  R.  B. 
Campbell,  article  by,  p.  325. 

Hathaway,  Ralph  R.,  article  by,  p.  449. 

Higgins,  Harold  G.,  and  Tyler  A.  Woolley, 
article  by,  p.  103. 

Hughes,  Karen  W.,  and  Robert  K.  Vickery,  Jr., 
article  by,  p.  240. 

Hylastes  asperatus,  p.  24. 

Invasion  of  big  sagebrush  (Artemesia  triden- 
tata)  by  white  fir  {Abies  concolor)  on  the 
southeastern  slopes  of  the  Warner  Mountains, 
California,  p.  319. 

Ips  pilifrons  thatcheri,  p.  29. 

Johnson,  D.  Elmer,  and  Lucile  Maughan 
Johnson,  article  by,  p.  407. 

Johnson,  Lucile  Maughan,  and  D.  Elmer  John- 
son, article  by,  p.  407. 

Larsen,  Kenneth  R.,  and  Wilmer  W.  Tanner, 
article  by,  p.  1. 

Lepthien,  Larry  W.,  and  Carl  E.  Bock,  article 
by,  p.  269. 

Leptocoenia  (subgenus),  p.  81. 

Life  history  and  ecology  of  Megarcys  signata 
(Plecoptera:  Perlodidae),  Mill  Creek,  Wa- 
satch Mountains,  Utah,  p.  39. 

Lindsey,  J.  P.,  and  R.  L.  Gilbertson,  article 
by,  p.  288. 

Lockard,  Lawrence  L.,  article  by,  p.  435. 

Lockard,  Lawrence  L.,  and  R.  Randall  Par- 
sons, article  by,  p.  425. 

Lockard,  Lawrence  L.,  R.  Randall  Parsons, 
and  Barry  M.  Schaplow,  article  by,  p.  442. 

Mathis,  Wayne  N.,  article  by,  p.  65. 

McDonough,  W.  T.,  R.  0.  Harniss,  and  R.  B. 
Campbell,  article  by,  p.  325. 

Medica,  Philip  A.,  Rudolf  G.  Arndt,  and  James 
R.  Dixon,  article  by,  p.  317. 

Morphology  of  ephemeral  and  persistent  leaves 
of  three  subspecies  of  big  sagebrush  grown 
in  a  uniform  environment,  p.  325. 

Morton,  Martin  L.,  and  John  S.  Gallup,  article 
by,  p.  427. 

Morton,  Martin  L.,  and  Robert  J.  Parmer, 
article  by,  p.  305. 

Multoribates  haydeni,  p.  104. 

Nectar  composition  of  hawkmoth-visited  spe- 
cies of  Oenothera  (Onagraceae),  p.  273. 

New  mites  from  the  Yampa  Valley  (Acarina: 
Cryptostigmata:  Oribatulidae,  Passalozeti- 
dae),  p.  103. 

New  record  of  the  bat  Plecotus  phyllotis  from 

Utah.  p.  452. 
New  records  of  stoneflies   (Plecoptera)  from 

New  Mexico,  p.  97. 
New  synonymy  and  new  species  of  American 
bark  beetles  (Coleoptera:  Scolytidae),  p.  21. 
New  synonymy  and  new  species  of  American 
bark  beetles  (Coleoptera:  Scolytidae),  Part 
n,  p.  391. 


Nichols,  D.  W.,  H.  D.  Smith,  and  M.  F.  Baker, 
article  by,  p.  191. 

Notes  on  the  genus  Bombylius  Linnaeus  in 
in  Utah,  with  key  and  descriptions  of  new 
species  (Diptera:  Bombyliidae),  p.  407. 

Owings,  Donald  H.,  and  Mark  Borchert,  p.  402. 

Paracoenia  iParacoenia)  ampla,  p.  71. 

Paracoenia  {Paracoenia)  calida,  p.  73. 

Paracoenia  {Paracoenia)  wirthi,  p.   78. 

Parahauloppia  cordylinosa,  p.  105. 

Parsons,  R.  Randall,  and  Lawrence  L.  Lock- 
ard, article  by,  p.  425. 

Parsons,  R.  Randall,  Lawrence  L.  Lockard, 
and  Barry  M.  Schaplow,  article  by,  p.  442. 

Parmer,  Robert  J.,  and  Martin  L.  Morton, 
article  by,  p.  305. 

Passalozetes  moniles,  p.  106. 

Penstemon  atwoodii,  p.  378. 

Phacelia  crenulata  var.  angustifolia,  p.  158. 

Photoperiodic  responses  of  phenologically 
aberrant  populations  of  pierid  butterflies 
(Lepidoptera),  p.  310. 

Pityokteines  mystacinus,  p.  29. 

Pityophthorus  amiculus,  p.  398. 

Pityophthorus  costatus,  p.  395. 

Pityophthorus  degener,  p.  397. 

Pityophthorus  dissolutus,  p.  398. 

Pityophthorus  explicitus,  p.  399. 

Pityophthorus  inceptis,  p.  396. 

Pityophthorus  mendosus,  p.  397. 

Pityophthorus  timidulus,  p.  396. 

Poche,  Richard  M.,  article  by,  p.  452. 

Poche,  Richard  M.,  and  George  A.  Ruffner, 
article  by,  p.  121. 

Provonsha,  A.  V.,  article  by,  p.  379. 

Pseudothysanoes  concentralis,  p.  27. 

Pseudothysanoes  tumidulus,  p.  28. 

Records  of  stoneflies  (Plecoptera)  from  Ne- 
vada, p.  49. 

Rreproductive  cycle  of  the  Belding  ground 
squirrel  {Spemophilus  beldingi  beldingi): 
seasonal  and  age  differences,  p.  427. 

Reveal,  James  L.,  Stanley  L.  Welsh,  and 
N.  Duane  Atwood,  article  by,  p.  327. 

Rodent  populations,  biomass,  and  community 
relationships  in  Artemisia  tridentata.  Rush 
Valley,  Utah,  p.  191. 

Roosting  behavior  of  male  Euderma  macu- 
latum  from  Utah,  p.  121. 

Roper,  Paul  R.,  and  Ferron  L.  Andersen, 
article  by,  p.  203. 

Ruffner,  George  A.,  and  Richard  M.  Poche, 
article  by,  p.  121. 

Sawin,  H.  Lewis,  Hobart  M.  Smith,  and  Ro- 
zella B.  Smith,  article  by,  p.  100. 

Schaplow,  Barry  M.,  Lawrence  L.  Lockard, 
and  R.  Randall  Parsons,  article  by,  p.  442. 

Schwartz,  Orlando  A.,  and  Vernon  C.  Bleich, 
article  by,  p.  62. 

Sexual  dimorphism  in  malpighaian  tubules  of 
Pteronarcys  califomica  Newport  (Plecop- 
tera), p.  449. 

Scolytodes  amabilis,  p.  26. 


Dec.  1975 


455 


Scolytodes  genialis,  p.  27. 

Scolytodes  lepidus,  p.  27. 

Scolytodes  obesus,  p.  26. 

Scolytus  laetus,  p.  25. 

Scolytus  torulus,  p.  25. 

Shapiro,  Arthur  M.,  article  by,  p.  33,  310. 

Smith,  Dwight  G.,  article  by,  p.  419. 

Smith,  H.  D.,  D.  W.  Nichols,  and  M.  F.  Baker, 

article  by,  p.  191. 
Smith,  Hobart  M.,  Rozella  B.  Smith,  and  H. 

Lewis  Sawin,  article  by,  p.  100. 
Smith,  Hobart  M.,  Rozella  B.  Smith,  and  Jean 

Guibe,  article  by,  p.  109. 
Smith,  Rozella  B.,  Hobart  M.  Smith,  and  H. 

Lewis  Sawin,  article  by,  p.  100. 
Smith,  Rozella  B.,  Hobart  M.  Smith,  and  Jean 

Guibe,  article  by,  p.  109. 
Some  parasites  of  paddlefish  (Polydon  spat- 
hula)  from  the  Yellowstone  River,  Montana, 

p.  425. 
Some  relationships  between  internal  parasites 

and  brown  trout  from  Montana  streams,  p. 

442. 
Some    relationships    between    water    fertility 

and  egg  production  in  brown  trout  {Salmo 

trutta)  from  Montana  streams,  p.  435. 
Stark,   Bill  P.,  Mary  R.   Gather,   and  Arden 

R.  Gaufin,  article  by,  p.  49. 
Stark,  Bill  P.,  Theodore  A.  Wolff,  and  Arden 

R.  Galfin,  article  by,  p.  97. 
Stockhouse,  Robert  E.,  II.,  article  by,  p.  273. 
Studies  in  nearctic  desert  sand  dune  Orthop- 

tera.   Part  XV.   Eremogeography  of  Spini- 

acris  with  bioecological  notes,  p.  113. 
Tanner,  Vasco  M.,  article  by,  p.  231. 
Tanner,  Wilmer  W.,  Charles  Fanghella,  and 

David  F.  Avery,  article  by,  p.  245. 
Tanner,  Wilmer  W.,  and  Kenneth  R.  Larsen, 

article  by,  p.  1. 
The    authorship   and   date    of   publication   of 


Siren  intermedia  (Amphibia:    Caudata),   p. 

100. 
The    identity   of   Boucourt's   lizard   Eumeces 

capita  1879,  p.  109. 
The  nest  and  larva  of  Diploplectron  brunneipes 

(Cresson)     (Hymenoptera:     Sphecidae),    p. 

127. 
The  Zygoptera  (Odonata)  of  Utah  with  notes  on 

their  biology,  p.  379. 
Thysanoes  tuberculatus,  p.  29. 
Tinkham,  Ernest  R.,  article  by,  p.  113. 
Traver,  Robbin  L.,  Joseph  M.  Branham,  and 

Arden  R.  Gaufin,  article  by,  p.  51. 
Utah  plant  novelties  in  Cymopterus  and  Pens- 

temon,  p.  377. 
Urosaurus  and  its  phylogenetic  relationship  to 

Uta  as  determined  by  osteology  and  myology 

(Reptilia:   Iguanidae),  p.  245. 
Vale,  Thomas  R.,  article  by,  p.  319. 
Vickery,  Robert  K.,  Jr.,  and  Karen  W.  Hughes, 

article  by,  p.  240. 
Water  balance  and  fluid  consumption  in  the 

southern    grasshopper    mouse,    Onychomys 

torridus,  p.  62. 
Weber,  D.  J.,  and  D.  J.  Hansen,  article  by, 

p.  86. 
Welsh,  Stanley  L.,  article  by,  p.  377. 
Welsh,    Stanley   L.,    N.    Duane   Atwood,    and 

James  L.  Reveal,  article  by,  p.  327. 
Wirth,  Willis  W.,  and  William  L.  Grogan,  Jr., 

article  by,  p.  275. 
Wolff,  Theodore  A.,  Bill  P.  Stark,  and  Arden 

R.   Gaufin,  article  by,  p.  97. 
Wood,  Stephen  L.,  articles  by,  p.  21,  391. 
Woolley,   Tyler   A.,    and   Harold   G.   Higgins, 

article  by,  p.  103. 
Xyleborus  californicus,  p.  399. 
Xyleborus  incultus,  p.  400. 
Xyleborus  molestulus,  p.  400. 
Xyleborus  tristiculus,  p.  401. 
Zygoribatula  apletosa,  p.  103. 


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TABLE  OF  CONTENTS 

Endangered,  threatened,  extinct,  endemic,  and  rare  or  lestricted  Utah  vas- 
cular plants.  Stanley  L.  Welsh,  N.  Duane  Atwood.  and  James  L. 
Reveal    327 

Utah  plant  novelties  in  Cymopterus  and  Pensternon.  Stanley  L.  Welsh  377 

The  Zygoptera  (Odonata)  of  Utah  with  notes  on  their  biology.    A.  Provon- 

sha    379 

New   synonymy   and   new   species   of   American   bark   beetles    (Coleoptera: 

Scolytidae),   Part  II.    Stephen  L.   Wood  391 

Correlates    of   burrow    location    in    Beechey    ground    squirrels.     Donald    H. 

Owings    and    Mark    Borchert    402 

Arachnids    as    ecological    indicators.     Dorald    M.    Allred    405 

Notes  on  the  genus  Bombylius  Linnaeus  in  Utah,  with  key  and  descrip- 
tions of  new  species  (Diptera:  Bombyliidae).  D.  Elmer  Johnson  and 
Lucile    Maughan    Johnson    407 

Breeding  range  expansion  of  the  starling  in  Utah.    Dwight  G.  Smith  419 

Some    parasites    of    paddlefish    (Polydon    spathula)    from    the    Yellowstone 

River,  Montana.    Lawrence  L.  Lockard  and  R.  Randall  Parsons  425 

Reproductive  cycle  of  the  Belding  gorund  squirrel   (Spermophilus  beldingi): 

seasonal  and  age  differences.    Martin  L.  Morton  and  John  S.  Gallup  ....     427 

A  new  combination  in  Pensternon  (Scrophulariaceae).  Stephen  L.  Clark  ....     434 

Some   relationships   between   water   fertility   and   egg  production  in   brown 

trout  (Salmo  trutta)  from  Montana  streams.  Lawrence  L.  Lockard  435 

Some  relationships  between  internal  parasites  and  brown  trout  from  Mon- 
tana streams.  Lawrence  L.  Lockard,  R.  Randall  Parsons,  and  Barry  M. 
Schaplow  442 

Sexual  dimorphism  in  malpighian  tubules  of  Pteronarcys  californica  New- 
port   (Plecoptera).     Ralph   R.    Hathaway 449 

New  records  of  the  bat  Plecotus  phyllotis  from  Utah.    Richard  M.  Poche  ....     452 

Index  to  Volume  35   453 


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