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PROCEEDINGS 


OF  THE 


California  Academy  of  Sciences 


FOURTH  SERIES 


Vol.  XXV 

ALICE  EASTWOOD 
SEMI-CENTENNIAL  PUBLICATIONS 


••i^ 


Marine  Biological  Labo^tjlo/y 
FEB  2  31949 

WOODS  HOLE,  MASS. 


SAN  FRANCISCO 

PUBLISHED  BY  THE  ACADEMY 

1943-1949 


■•^\ 


— ^  ^ 


CttcCt^     ^xJ^^'-^^'T^J^ 


TO 

ALICE  EASTWOOD 

IN  COMMEMORATION  OF  FIFTY  YEARS  OF  SERVICE  AS 
CURATOR  OF  THE  DEPARTMENT  OF  BOTANY  OF  THE 

CALIFORNIA  ACADEMY  OF  SCIENCES 

1892-1942 


THIS  VOLUME  IS  DEDICATED  IN  RESPECT  AND  AFFECTION 


FOREWORD 

Imsx'OLUMEof  the  Proceedingsoi  the  Caliloi  Ilia  Academy 
of  Sc  iences  honoring  Miss  Alice  Eastwood  ^vas  initiated  in 
i()42.  on  the  occasion  of  her  comj)lction  ol  lilty  years  ol 
service  as  Curator  ol  tlie  Academy's  Department  of  Botany. 
O^sing  to  u'artime  problems  of  publication,  the  first  num- 
ber of  the  projected  volinne  was  not  isstied  until  December. 
ic)43.  The  volume  has  been  continued  through  the  suc- 
ceeding years  and  is  now  ccjncluded  ^vitll  this  number 
which.  ha}:)pily.  we  are  able  to  isstie  on  the  occasion  ol 
Miss  Eastwood's  ninetieth  birthday,  and  in  recognition  ol 
Iier  liity-seven  years  of  service  to  the  Academy. 

On  her  ninetieth  birthday  Miss  Eastwood  \vi\\.  at  her 

own  rccjuest,  retire  from  her  post  as  Cin-ator  of  Botany. 

Ihc  Academy's  Cotuicil  is  protid  to  grant  her  the  title  of 

Curator  Emeritus— a  title  which,  however,  seems  singularly 

inappropriate  to  a  gallant  lady  ^v  ho  ^vill  be  forever  young. 

The  biograpliy  of  Miss  Eastwood  that  is  here  presented 
was  prepared  by  Dr.  F.  M.  MacFarland.  President  of  the 
Academy  from  19^^^  to  i94(),  in  collaboration  with  Dr.  R. 
C.  Miller.  Director  of  the  Academy,  and  Mr.  John  lliomas 
Howell,  Assistant  Curator  of  Botany.  The  bibliography  of 
Miss  Eastwood's  writings  was  prepared  by  Dr.  MacFarland 
and  Miss  Veronica  }.  Sexton,  the  Academy's  Executive 
Eibrai  ian.  The  thankless  task  of  indexing  the  \-olume  ^vas 
performed  by  Dr.  MacFarland  alone. 

For  all  those  A\'ho  participated  in  its  preparation  and 
publication,  this  volume  represents  a  labor  of  l(^\e. 

M.   E.   LOMRARDI. 

Presidciu  ol  the  Acadcniv 


CONTENTS  OF  VOLUME  XXV 

PAGES 

Biograpliical  Skctcli  of  Alice  Eastwood IX-    XIV 

BibliogTaphy  of  the  Writings  of  Alice  Eastwood XV-XXIV 

Contributed  Papers : 

No.    1.  MuNZ,  Philip  A.   A  Revision  of  the  Genus  Fuchsia  (Ona- 

graceae)  (Plates  1-16)  Published  December  11,  1948 ....        1-138 

No.  2.  Campbell,  Douglas  Houghton.  Relations  of  the  Temperate 
Floras  of  North  and  South  America.  Published  June  1, 
1944  139-146 

No.    3.  Barneby,  R.  C.  Pugillus  Astragalorum  Alter   (Plate  17). 

Published  June  1, 1944 147-170 

No.    4.  Smith,  Gilbert  M.  Sublittoral  Marine  Algae  of  the  ^lon- 

terey  Peninsula.  Published  June  1,  1944 171-176 

No.  5.  Wolf,  Carl  B.  The  Gander  Oak,  a  New  Hybrid  Oak  from 
San  Diego  County,  California.  (Plates  18-19).  Published 
June  1, 1944 171-188 

No.    6.  Wiggins,  Ira  L.  The  Genus  Drymaria  in,  and  adjacent  to  the 

Sonoran  Desert.  (Plates  20-22) .  Published  June  1, 1944 .  .   189-214 

No.    7.  Pierce,  George  J.  Water  and  Plant  Anatomy.  Published 

June  IJ  944 215-220 

No.  8.  ]\Iason,  Herbert  L.  A  Pleistocene  Flora  from  the  McKittrick 
Asphalt  Deposits  of  California.  (Plates  23-24) .  Published 
June  1, 1944 221-234 

No.  9.  Ew^an,  Joseph.  The  Perennial  Southwestern  Datura  and  the 
Validity  of  Matthew's  Hypothesis  in  Plant  Geography 
(6  text  figures) .  Published  June  1, 1944 235-244 

No.  10.  Benson,  Lyman.  A  Revision  of  some  Arizona  Cactaceae 

(Plate  25).  Published  November  10,  1944 245-268 

No.  11.  Babcock,  Ernest  B.  Endemism  in  Crepis  (4  text  figures) 

Published  November  10, 1944 269-290 

No.  12.  Goodspeed,  T.  H.  Nicotiana  Arentsii — A  New,  Naturally 
Occurring  Amphidiploid,  Species.  (Plates  26-27).  Pub- 
lished November  10, 1944 291-306 

No.  13.  Stebbins,  G.  L.,  Jr.,  Tobgy,  H.  A.,  and  Harlan,  Jack  R.  The 
Cytogenetics  of  Hybrids  in  Bromus  II.  Bromus  Carinatus 
and  Bromus  Arizonicus.  (13  text  figures).  Published  No- 
vember 10,  1944 307-322 


PAGES 

No.  14.  McMiNN,  Howard  E.  The  Importance  of  Field  Hybrids  in 
Determining  Species  in  the  Genus  Ceanothus.  (Plates 
28-36,  3  charts,  3  histograms).  Published  November  16, 
1944 323-356 

No.  15.  Howell,  John  Thomas.  A  Revision  of  Phacelia  Section  Mil- 

titzin.  Published  December  27, 1944 357-376 

No.  16.  St.  John,  Harold.  Endemisni  in  the  Hawaiian  Flora,  and 
a  Revision  of  the  Hawaiian  Species  of  Gunnera  (Hal- 
oragidaceae)  Hawaiian  Plant  Studies  11.  (Plates  37-46). 
Published  November  15,  1946 377-420 

No.  17.  Keck,  David  D.  A  Revision  of  the  Artemisia  vulgaris  Com- 
plex ill  North  America.  (19  text  figures)  Published  No- 
vember 15, 1946 421-468 

No.  18.  Setchell.  William  A.  The  Genus  Ruppia  L.  (Plates  47-48) . 

Published  November  15,  1946 469-478 

Index  to  Volume  XXV 479-485 


BIOGRAPHICAL  SKETCH  OF 
ALICE  EASTWOOD 

Ox  .January'  19,  1851),  in  two  localities,  some  two  thousaiul  miles  apart,  there 
were  in  progress  two  separate  and  apparently  unrelated  series  of  events  that 
were  destined  to  become,  at  a  later  date,  of  considerable  mutual  importance. 
In  San  Francisco,  the  California  Academy'  of  Sciences,  an  organization  then 
less  than  six  years  old,  was  struggling  successfully  through  the  precarious 
early  years  of  its  existence.  In  Toronto,  Canada,  Alice  Eastwood  Avas  born. 

^lore  than  tliirty  years  were  to  elapse  before  the  events  referred  to  became 
signifieant  for  each  otlier.  Nevertheless,  even  before  Alice  Eastwood  was  born, 
the  avenue  that  was  to  lead  her  to  distinguished  botanical  career  had  been 
opened  by  the  "scientific  gentlemen"  (as  they  described  themselves  in  their 
fir.st  constitution)  who  made  up  the  California  Academy  of  Sciences.  As  if 
with  prophetic  insight,  Dr.  Albert  Kellogg,  one  of  the  founders  of  the  Academy 
and  its  most  prominent  early  botanist,  had  proposed  in  a  quaintly  worded 
resolution  that  was  adopted  at  the  meeting  of  August  1,  1853  :  "Be  it  resolved, 
as  the  sense  of  this  society,  that  we  highly  approve  of  the  aid  of  females  in 
every  department  of  natural  history,  and  that  we  earnestly  invite  their  coop- 
eration." Thus,  through  Dr.  Kellogg's  resolution,  as  gallant  in  intent  as  it 
was  inej)t  in  phrase,  the  California  Academy  of  Sciences  became  very  probably 
the  first  institution  in  the  world  to  formally  encourage  the  participation  of 
women  in  scientific  work,  as  it  became  later  the  first  to  admit  women  to  senior 
curatorial  positions. 

While  Alice  Eastwood  was  still  a  young  girl,  her  family  moved  from  Toronto 
to  Denver,  Colorado,  where  she  obtained  her  high  school  education,  graduating 
from  the  East  Denver  High  School  with  its  first  class  in  1879.  Her  marked 
ability  and  eager  thirst  for  knowledge  led  her  to  be  drafted  as  a  substitute 
teacher  before  she  had  finished  high  school,  and  slie  continued  as  a  regular 
teacher  for  the  next  ten  years.  Despite  the  burden  of  teaching  a  great  variety 
of  more  or  less  unrelated  subjects,  her  innate  love  of  Nature  asserted  itself 
and  the  plants,  shrubs,  and  trees  of  Colorado  appealed  to  her  with  irresistible 
charm.  At  every  opportunity  she  explored  the  plains,  valleys,  and  mountains, 
the  rugged  canyons,  streams,  and  rivers  of  the  state,  and  brought  together  a 
tangible  record  of  her  field  studies  in  the  Eastwood  Herbarium,  which  later 
was  to  become  the  pride  of  the  Denver  Museum.  Without  access  to  a  scientific 
library  or  contacts  with  other  biologists,  she  became,  herself,  a  self-taught 
botanist.  Gray's  IManual,  covering  the  flora  of  the  eastern  states  was  her 
earliest  guide  to  unraveling  the  family  and  in  part  the  generic  relationships 
of  the  Colorado  plants,  and  she  welcomed  with  delight  the  appearance  of 
Coulter's  ^Manual  of  Rocky  Mountain  Botany  in  1885,  and  she  soon  found 
ample  work  to  do  in  extending  its  application  to  the  rich  Colorado  flora. 

In  1890,  Miss  Eastwood  made  a  vacation  trip  to  southern  California,  study- 
ing especially  the  plants  of  the  San  Diego  region.  Returning  in  the  next  year, 

[  IX  1 


X  CAUFOBMA  ACADEMY  OF  SCIENCES  [Proc  4th  Ser. 

she  spent  some  niunths  as  an  assistant  in  the  li('i'l)ai-iiiiii  (if  the  California 
Academyof  Sciences.  Asa  result,  she  Avas  induced  to  leave  her  Denver  position 
in  1892.  and  to  east  her  h»t  with  the  Aca(h'iiiy.  soon  heeomiii^'  joint  ("ui-ator 
of  Botany  with  Mrs.  Katliei-iiie  IJraiiih'izce.  Thi'ci'  ye;ii-s  hitei'.  upon  t  he  retire- 
ment of  ]Mrs.  I'.randejree,  she  became  Cui-ator  and  llrjid  of  the  Dt'iiai-tinent  of 
Botany  wliich  position  she  has  fUhMl  until  the  pi'escni. 

In  the  forty-seven  years  of  its  existence,  the  (jilifoinia  Academy  of  Sci- 
ences had  grown  fnmi  its  simple  beg'innings  to  he  the  leading  center  of  research 
in  the  Natural  Sciences  of  tiie  Pacific  Coast.  Its  heH)arium,  built  up  by  the 
pioneers.  Allx-rt  Kellogg.  11.  X.  P>olander.  Harry  M.  Ilarkness,  and  E.  L. 
(Jreen,  and  continued  by  T.  S.  and  Katherine  lirandegee  and  others,  had 
become  one  of  the  largest  in  the  West,  and  was  in  constant  use  by  student 
workers.  ]\Iiss  Eastwood  at  once  entered  upon  W(  rk  to  inci-ease  its  value.  Aside 
from  her  I'ontinc  ciiralorial  duties,  she  collected  and  studied  plants  and  trees 
near  and  fai'.  The  Hay  I'cgion  was  thoroughly  explored  and  ti'ips  were  made 
to  Mt.  Shasta,  to  Del  Norte  County,  to  the  Santa  Lucia  Mountains,  and  the 
interior  valleys,  returning  with  much  new  material  for  the  herbariiun. 

Tn  those  pre-autoniobile  days,  travel  away  from  the  railroads  was  often 
difticult.  liV  stage,  on  hoi-seback,  or  afoot.  Miss  Eastwood  made  her  Avay  over 
mountain  roads  and  trails  with  unflagging  zeal. 

As  a  resnlt  of  her  activity  and  that  of  those  whom  she  inspired,  the  herl)ar- 
iiim  steadily  increased  in  size  and  scientific  value.  The  great  earth(|uake  and 
fire  of  1!H)().  however,  destroyed  much  of  San  Francisco  in  a  few  hoin-s.  and 
the  nuisenni  and  collections  of  the  Academy  shared  the  same  fate.  Through 
Miss  Eastwood's  iiutiative  and  courage,  though,  some  important  things  were 
saved — the  I'eeords  of  the  Academy  from  its  beginning,  and  twelve  liinidred 
and  eleven  invaluable  botanical  types,  irreplaceable  if  lost.  Miss  Eastwood 
had  long  recognized  their  value  and  had  segregated  them  from  the  thousands 
of  other  specimens,  so  that  when  the  emergency  came,  she  was  able  to  find 
them  without  delay  in  the  wrecked  building  and  to  transport  them  to  a  place 
of  apparent  safety — a  I'efuge  that  nevertheless  had  to  be  changed  twice  before 
the  approaching  fiames.  Tn  doing  this,  she  unhesitatingly  abandoned  her  ow'n 
personal  possessions,  her  favorite  pocket  lens  alone  being  saved.  In  a  letter 
to  Science  for  May  25,  lf)()(i,  describing  the  (h'stiMidion  of  the  Academy  and 
collections,  she  wrote:  "1  do  not  feel  the  loss  to  be  nnne,  but  it  is  a  great  loss 
to  the  scientific  woi'ld  and  an  irreparable  loss  to  California.  My  own  destroyed 
work  1  do  iu)t  lament,  foi'  it  was  a  joy  to  me  while  I  did  it,  and  1  still  can  have 
the  same  joy  in  sfjuting  it  again.  ...  To  me  came  the  chance  to  care  for  what 
was  saved  from  the  ruins  of  the  Academy,  and  with  the  hel]i  of  my  devoted 
friends  T  was  able  to  do  it." 

The  type  s]")ecimens  saved  thi'ough  ^liss  Eastwood's  courage  formed  the 
luicleus  of  the  new  collections.  But  owing  to  the  total  destruction  of  its  prop- 
ei-ty  and  the  interrujition  of  its  income,  it  was  several  years  before  normal 
aetivitv  could  be  resumed.  During  this  lime,  ]\Iiss  Eastwood  studied  in  the 


#0^  W9^ 


^%t 


ALICE    EASTWOOD 

Ahiiiil    iSSo,  ;ii   ilic  Ijfi^imiiiif^  ol   lic'i    lioUiiiii  ill  (aicci 


Vol.  XXV]  ALICE  EASTWOOD  XIIT 

j^'i-o^at  lierbaria  of  the  Siuitli.soiiiaii  Institution  at  Wasliinuton,  I  he  Xt'w  Voi-lx 
Botaiiical  Garden,  and  Harvard  I'niversity,  and  also  spent  a  ninnber  oi' 
months  in  the  Kew  Gardens,  the  British  Museum,  tlie  University  of  Cam- 
bridge, and  the  Jardin  des  Plantes. 

With  tlic  new  site  for  its  bnihlini^s  in  (iokh'ii  Gate  Park  assured  and  con- 
struction begun,  tlie  Academy's  normal  scientific  activity  was  resinned.  Miss 
Eastwood  returned  in  June  1912,  and  at  once  vigorously  began  the  replace- 
ment of  the  herbarium  of  the  Department  of  Botany  and  tlie  rebuilding  of 
its  library;  collecting  trips  were  made  at  every  opportunity.  Through  the 
cooperation  of  Professor  Charles  S.  Sargent  of  the  Arnold  Arboretum,  she 
made  a  trip  to  Alaska  in  1914,  collecting  and  studying  especially  the  willows 
of  the  Yukon  region  for  his  Sylva  of  North  America .  Later,  studies  were  also 
made  by  her  in  southern  California,  Arizona,  and  New  Mexico  for  Professor 
Sargent.  She  made  other  extensive  trips  to  all  parts  of  California,  to  Oregon, 
Nevada,  Utah,  Arizona,  New  Mexico,  and  Lower  California,  returning  with 
abundant  material  for  the  rapidly  growing  herbarium.  Additional  collections 
made  by  others  came  from  Mexico,  Central  America,  South  America,  and  the 
Pacific  Islands. 

Especial  attention  has  also  been  given  to  securing  specimens  of  the  exotic 
plants  which  have  been  introduced  into  California  gardens  and  parks,  a  field 
often  neglected  in  herbaria,  but  of  great  importance  for  com]iarative  study. 

Miss  Eastwood's  publications  as  shown  in  the  accompanying  bibliography 
of  over  three  hundred  titles  have  been  many  and  varied.  The  majority  of  them 
are,  of  course,  technical  botanical  studies,  but  others  are  of  more  general 
interest  inculcating  an  appreciation  of  the  out-of-doors  and  the  conservation 
of  its  ti-easures.  In  addition,  the  admirably  edited  "Leaflets  of  Western 
Botany"  published  by  her  and  Mr.  J.  T.  Howell,  Assistant  Curator  of  Botany, 
affords  a  prompt  means  of  publication  for  shorter  articles  by  western  botanists. 

It  is  worthy  of  particular  remark  that,  while  Miss  Eastwood  had  written 
more  than  one  hundred  papers  by  the  time  she  was  fifty  years  of  age,  and 
was  considered  sufficiently  distinguished  to  be  starred  in  the  first  edition  of 
Americcni  Men  of  Science  (1906),  two-thirds  of  her  published  writings  have 
been  issued  since  that  time,  and  fifty-five  of  them  since  her  eightieth  birthday. 

Miss  Eastwood  has  taken  a  prominent  part  in  promoting  the  welfare  of 
many  scientific  organizations  interested  in  plant  life.  She  has  been  the  active 
head  of  the  California  Botanical  Club  (of  San  Francisco)  since  its  second 
year  in  1892.  She  has  participated  in  the  work  of  the  California  Floral  Society, 
the  San  Francisco  Garden  Club,  the  California  Spring  Blossom  and  Wild 
Flower  Association,  the  American  Fuchsia  Society,  the  California  Horticul- 
tural Society,  and  is  a  member  of  the  Science  Club,  the  Alpine  Club,  the 
Tamalpais  Conservation  Club,  and  other  organizations  devoted  to  the  out-of- 
doors  and  its  conservation.  She  became  a  Member  of  the  California  Academy 
of  Sciences  in  1892,  a  Life  Member  in  1917,  was  elected  a  Fellow  of  the  Acad- 
emy on  February  25,  1928,  was  declared  a  Patron  on  December  .{l,  1930,  and 


XIV  CAUFOSNIA  ACADEMY  or  SCIENCES  [Proc.  4th  Ser. 

was  iiiiaiiimously  elected  an  Honorary  ^Member  of  the  Academy  at  its  Annual 
Meeting  on  February  18,  1942. 

She  has  represented  the  Academy  in  many  scientific  meetinos  at  home  and 
abroad.  Among  these  latter  are  the  Internatioiuil  Congress  held  in  Cambridge 
ill  l!t.")().  in  Holland  in  1!>.'}5.  and  the  Centennial  Anniversary  of  the  Jardin 
des  riantes  in  the  same  j'ear. 

Xdtwitlistanding  the  ]ires.sure  of  her  scientific  activities.  Miss  Eastwood 
lias  never  been  too  busy  to  help  others,  to  answer  the  countless  questions  that 
come  to  her  daily,  to  extend  willingly  to  others  the  richness  of  her  knowledge. 
Her  frank,  direct  approach  to  every  problem  and  her  modest,  kindly  spirit 
have  won  for  her  the  esteem  and  love  of  countless  loyal  and  devoted  friends 
who  wish  for  her  many  more  years  of  productive  activity  in  the  field  she  loves. 


BIBLIOGRAPHY  OF  THE  WRITINGS  OF 
ALICE  EASTWOOD 

181U.  Thf  luorning  glory  liusli,  liKiiiund  hpiophylla.  West.  Amer.  Scientist,  7:119.  (Feb.) 
LcKcocrinum  montanum.  West.  Amer.  Scientist,  7:141.   (Mar.) 
The  common  shrubs  of  sonthwest  Colorado.  Zoe,  2  :  102-104.  (July) 
The  fertilization  of  geraniums.  Zoe,  2:112.  (July) 
The  Mariposa  lilies  of  Colorado.  Zoe,  2:201-203.  (Oct.) 
Additions  to  the  flora  of  Colorado,  I.  Zoe,  2:226-333.  (Oct.) 

1892.  The  loco  weeds.  Zoe,  3:53-.58.  (April) 

Eeview  on  "Outlines  of  lessons  in  botany  for  the  use  of  teachers,  and  mothers  studying 

with  their  children"  by  Jane  H.  Newell.  Zoe,  3  :78.  (April) 
Notes  on  some  species  of  the  genus  Oenothera.  Zoe,  3  : 248-252.  (Oct.) 

1893.  Notes  on  the  cliff  dwellers.  Zoe,  3  :375-376.  (Jan.) 

General  notes  of  a  trip  througli  southeastern  Utah.  Zoe,  3 : 354-361.  (Jan.) 

Notes  on  some  Colorado  plants.  Zoe,  4:2-12.  (April) 

Additions  to  the  flora  of  Colorado,  II.  Zoe,  4:16-20.  (April) 

A  popular  flora  of  Denver,  Colorado.  San  Francisco,  Zoe  Publ.  Co.,  May,  1893.  57  pp. 

List  of  plants  collected  in  southeastern  Utah,  with  notes  and  descriptions  of  new 

species.  Zoe,  4:113-127.  (July) 
Field  notes  at  San  Emidio.  Zoe,  4:144-147.  (July) 
Botanical  notes.  Zoe,  4:286-287.  (Oct.) 
Gilia  supcrha.  Phacelia  nudicaiijiti.  Zoe,  4:296.  (Oct.) 

1895.  Additions  and  corrections,  January  14,  1895.  3  pp.  [To:  A  popular  flora  of  Denver, 

Colorado.  1893] 
Two  new  species  of  Aquilegia  from  tlie  Upper  Sonoran  Zone  of  Colorado  and  Utah. 

Proc.  Calif.  Acad.  Sci.,  (2),  4:559-562,  pis.  18-19.  (Mar.) 
On  heteromorphic  organs  of  Sequoia  sempervirens.  Proc.  Calif.  Acad.  Sci.,  (2),  5  :170- 

176,  pis.  15-18.  (May) 
Leucocrinum  montanum.  West  Amer.  Scientist,  9,  78:69.  (June) 
Observations  on  the  habits  of  Nemophila.  Erythea,  3  :  151-153,  pi.  3.  (Nov.) 

1896.  New  localities  for  west  American  plants.  Erythea,  4:32.  (Feb.) 
Pelargonium  anceps,  Ait.  Erythea,  4:34.  (Feb.) 

New  localities  for  two  introduced  plants.  Erythea,  4:34-35.  (Feb.) 

Ephedra  viridis  Coville.  Erythea,  4:71.  (April) 

Trillium  sessile.  Erythea,  4:71.  (April) 

On  Dr.  D.  Prain's  Account  of  the  genus  Argemone.  Erythea,  4 : 93-96.  (.June) 

New  stations  for  two  introduced  plants.  Erythea,  4 :99.  (June) 

Arbutus  Memiesii  in  San  Francisco  County.  Erythea,  4:99.  (June) 

Report  on  a  collection  of  plants  from  San  Juan  County,  southeastern  Utah.  Proc. 

Calif.  Acad.  Sci.,  (2),  6:270-329,  pis.  44-46.  (Aug.) 
The  alpine  flora  of  Mt.  Shasta.  Erythea,  4:136-142.  (Sept.) 
Geranium  parviflorum  Willd.  Erythea,  4:145.  (Sept.) 
Scolymus  Hispanicus  L.  Erythea,  4:145.  (Sept.) 
New  stations  for  two  plants.  Erythea,  4:151.  (Oct.) 
Abies  bracteata.  Erythea,  4:174-175.  (Nov.) 
Centaurea  calcitrapa  L.  Erythea,  4:175.  (Nov.) 

1897.  A  guide  for  the  analysis  and  description  of  flowering  plants.  Prepared  for  the  Botani- 

cal Club,  California  Academy  of  Sciences.  San  Francisco,  George  Spaulding  &  Co. 
8  pp. 

[  XV  ] 


XVI  CALIFOKXIA  ACADEMY   or  SCIENCES  [Pkoc.  4th  Ser. 

[Ht-vit-w  ofj  Volinne  X  of  the  "Hilva  of  North  Aiiiorii-ii,"  l)y  Sargent.  Krythea,  5:43- 

44.  (March) 
[Review    of]    "L.-ilmratory    ]iractic('    for    lieyiiincrs    in    liotnny."    Erythea,    Si.lfl-Cil. 

(April  I 
Tlie  coiiifcrac  of  the  Santa  T.m-ia  Mountains.  Krytln-M,  5:71-74.  (June) 
Caiiforniaii  licrli  loic — J,'li(t  nniu.s  CalifoniicKs.  I']iytlica,  5  :98.  (Sept.) 
Studies  in  the  herliaiium  and  1h<'  titd.l.  I.  I'rnc.  ( 'alif.  Acad.  Sci.,  (3),  Botany,  1  :71-88, 

pi.s.  ()-7.  (Nov. ) 

1898.  Ferns  of  the  Yoseniite  and  the  neifrlil.oriny  Sierras.  Krythea.  (i:  14-15.  (Feb.) 
[Review  of  J  The  second  fascicle  of  ''IIow  ell's  Flora  of  northwest  America."  Erythea, 

6:58-60.  (May) 
Plant  inhabitants  of  Nob  Hill,  San  Francisco.  Erythea,  6:61-67.  (June) 
Notes  on  tin-  flora  of  Marin  County,  California.  Erythea,  6:72-75.  (July) 
Is  X(  riiiiliiillinu  frnar  n  septennial.'  Erythea,  6  :  75-76.  (July) 
I'l/iiihi  iiiiiKir  \j.  in  California.  I''iytliea,  6:93.  (Sept.) 
Studies  in  the  heriiariiini  and  tlic  Held,  I  1 .  i'roc.  Calif.  .Acad.  Sci.,  (3),  Botany,  1  :89- 

14(;,  jds.  S-l  I.  (Sept.) 
I'lants  in  Hower  in  November  and  December,  1897.  Erythea,  6:114-115.  (Nov.) 

1899.  New  localities  for  rare  California  plants.  I'^rythea,  7:76-77.  (Aug.) 
I'ariiti.sfiia  califoniica  Greene.  Erythea,  7:84.  (Sept.) 

Ai(  ntiiiii  iKiliitliciild.  Krythea,  7:149.  (Nov.) 

Introduced  plants  in  Placer  County,  California.  Erythea,  7  :  150.  (Nov.) 

Sedge  used  in  Indian  basket  making.  Erythea,  7:15U.  (Nov.) 

.Mr.  Ilendrie's  gifts  to  the  California  Academy  of  Sciences.  Erythea,  7  :15U.  (Nov.) 

Migratory  plants  in  A  l.i  nieda  County.  Erythea,  7:175-176.  (Dec.) 

.\n(itlici-  int  i(H|iii-ed  plant.  Krythea,  7:176.  (Dec.) 

190(1.  ...  Krv  and  tidia.  Rocky  Mountain  cilitidn.  I'i-cparcd  Ijv  .Mice  Kastwdiid.  I'.oston, 
(;iiiii  \-  Cinnpany,  |  (-19001  1311  p.  tigs.  1-  I  I.  At  head  of  title:  P.ergen's  Foundations 
of  Jjiitany.  At  lie;i<l  of  cover-title:  Bergen's  Klenients  of  botany. 

Notes  on  ('ii/in  s.^ii.s  M acnahkma.  Zee,  5:11-13.  (.Tune) 

.\iliilli  (/Iti  i.iiiiild.  Zoe,  5:28-S0.  (July) 

On  the  occurrence  of  IiJi(i(Ki<liolas  liedj/pnois  All.  ( Ifi  ilii/nuiis  /kiI i/iinir/iliid  DC.)  in 
North  AuH'rica.  Zoe,  5:35-36.  (July) 

Finns  Lmiihi  rluina  on  Mt.  St.  Helena.  Zoe,  5:36-37.  (July) 

Arcloshijiliiiliis  as  a  host  plant  for  Phoradi  ndvoii.  Zoe,  5  :37.  (July) 

ScorzoiK  ra  hispanicd  1^.  Zoe,  5:37.  (July  i 

Some  plants  of  Mendocino  County  new  to  the  tlora  of  California.  Zoe,  5:58-()(i.  (Aug.) 

Rediscovery  of  Tliermopsis  macrophylla  IT.  &  A.  Zoe,  5  : 7()-78.  (Sept./Oct.) 

New  sjiecies  of  California  jdants.  Zoe,  5:80-90.  (Sept./Oct.) 

llMi  1 .  .  .  .  Kc\'  iiikI  tloia.  Pacific  Coast  editi(ni.  I'i('|iarrd  iiy  .Mice  Fastwood.  1  ioslon,  ( i  inn 
&  ('oni|iaiiv,  liiol.  |cl897,  1901].  l!»!l  p.  4  ligs.  At  head  of  title:  Bergen's  Botany. 

.  .  .  K(\  and  tidia.  Pacific  Coast  edition.  i're])ared  liy  .Mice  Eastwood.  Boston,  dinn  & 
(  (iiiilKiiiN ,  |iis:i7,  PHil  |.  199  p.  4  figs.  At  head  of  title:  Bergen's  Botany.  liound 
witli:  "  l''oundations  of  botany,"  by  .Fosepli  )'.  PxTgcii.  Pioston,  (iinn  lV  ('oiii|iany, 
I  cl901J.  412  p.  front.,  illns.,  plates. 

Asrirpias  Kofolo.  Zoe,  5:98-99.  (Feb./Apr.) 

A  study  of  Ertj.sinium  (jrandiflorum  Nutt.  and  the  species  aggregated  undi  r  it.  Zoe, 
5:100-103.  (Feb./Apr.) 

Alriph  .1-  srniibarcdtnm  R.  Br.  in  Marin  County.  Zoe,  5: 13()-137.  (T-'eb./Apr.) 

Iliihrnarid  nidril inia  (Jreene.  Zoe,  5:137.  (Feb./Apr.) 

Plants  used  for  poisoning  fish.  Zoe,  5:136.  (Feb./Apr.) 


Vol.  XXV]  WBITINGS  OF  ALICE  EASTWOOD  XVII 

Some  sniJiU-flowered  species  of  Neniupliila  from  the  Pacific  Coast.  Jiulleliii  of  the 
Torrey  Botanical  Club,  28:137-160,  pis.  15-20.  (Mar.) 

An  undescribed  species  of  Paron/icliid  from  California,  r.ulletin  of  tlic  'I'm  rey  Botani- 
cal Club,  28  :2S8-289.  (May  ) 

Notes  on  Californian  species  of  Delphinium.  UuUetin  of  the  Torrey  liotanical  Club, 
28:667-674.  (Dec.) 

1902.  From  Bedding  to  the  snow-clad  peaks  of  Trinity  County  ;  also.  List  of  trees  and  shrubs 

seen  en  route.  Sierra  Club  Bulletin,  4:39-58,  pis.  LV-LVI.  (Jan.) 
Some  new  species  of  California  plants.  Bulletin  of  the  Torrey  Botanical  Club.  29 : 7.j- 

82,  pis.  6-7.  (Feb.) 
A  descriptive  list  of  the  plants  collected  by  Dr.  F.  E.  Blaisdell  at  Nome  City,  Alaska. 

Botanical  Gazette,  XXXIII :  126-149,  figs.  1-3,  Feb.  21,  1902;  199-213,  figs.  4-5, 

Mar.  22,  1902  ;  284-299,  figs.  6-10,  Apr.  18,  1902. 
Some  new  species  of  Pacific  Coast  Ribe.s.  Proc.  Calif.  Acad.  Sci.,  (3),  Botany,  2:241- 

254,  pis.  23-24.  (Apr.) 
New  species  from  the  Sierra  Nevada  Mountains  of  California.  Proc.  Calif.  Acad.  Sci., 

(3),  Botany, 2:285-293.  (June) 
A  flora  of  the  South  Fork  of  Kings  River  from  Millwood  to  the  head  waters  of  Bubbs 

Creek.  Sierra  Club.  Publ.,  No.  27.  pp.  1-96,  illus.  (June) 
New  species  of  Nemophila  from  the  Pacific  Coast.  Bulletin  of  the  Torrey  Botanical 

Club,  29:471-474,  pi.  21.  (July) 
New  western  plants.  Bulletin  of  the  Torrey  Botanical  Club,  29:523-525.  (Aug.) 

1903.  New  species  of  Oreocarya.  Bulletin  of  the  Torrey  Botanical  Club,  30  :238-246.  (April ) 
New  species  of  western  plants.  Bulletin  of  the  Torrey  Botanical  Club,  30:483-502. 

(Sept.) 
Notes  on  Garrya  with  descriptions  of  new  species  and  key.  Botanical  Gazette,  36:456- 
463.  (Dec.) 

1904.  Description  of  a  light,  compact  arrangement  of  bedding  and  personal  effects  for  a 

camping  trip.  Sierra  Club  Bulletin,  5:77-78.  (Jan.) 
Hans  Herman  Behr.  Science,  19:636.  (April) 

Some  new  species  of  western  Polemoniaceae.  Botanical  Gazette,  37:437-447.  (June) 
A  new  GUia.  Botanical  Gazette,  38:71-72.  (July) 

1905.  Report  of  the  Committee  appointed  to  prepare  and  present  an  account  of  the  life  and 

services  of  Doctor  Hans  Herman  Behr.  Read  before  the  California  Academy  of 
Sciences,  March  21,  1904.  San  Francisco,  The  California  Academy  of  Sciences, 
1905.  7  pp.  front,  (port.)  F.  Gutzkow,  George  Chismore,  Alice  Eastwood,  Committee. 

New  species  of  western  plants.  Bulletin  of  the  Torrey  Botanical  Club,  32:193-218. 
(April) 

A  handbook  of  the  trees  of  California.  Occasional  Paper,  No.  9,  Calif.  Acad.  Sci.  86 
pp.,  57  pis.  (July) 

1906.  New  species  of  California  plants.  Botanical  Gazette,  41 : 283-293,  2  figs.  (April) 

The  California  Academy  of  Sciences.  Science,  N.S.,  23  : 824-826.  May  25,  1906.  [Special 
article  quoting  Miss  Eastwood's  letters  describing  the  destruction  from  the  earth- 
quake and  fire  of  1906] 

The  earthquake  and  the  California  Academy  of  Sciences.  Torrey,  6:120-123.  (June) 

1907.  PeltipJiyUum  pelfatum.  ^luhlenbergia,  3  :78.  (Aug.) 
EaHlardella  muirii  Gray.  Muhlenbergia,  3:78.  (Aug.) 
Truutvetteria  grandis  Nutt.  Muhlenbergia,  3:78.  (Aug.) 

Cypripedium  fasciculaium   in  Santa  Cruz  County,  California.   .Mulilcnlicrgia,  3:97. 

(Sept.) 
Notes  on  California  i)lants.  ^Muhlenbergia,  3:131.  (Dec.) 


XVIII  CALIFOBXIA  ACADEMY  OF  SCIEXCES  [Proc.  4tu  Sku. 

1908.  Extensions   of   range.    Lewisia    redivivn   ruisli.    J^c\vi.si;i    Icaiia    (Porter  i    l^oliiiison. 

Muhlenbergia,  3  :147.  (.I.iii. 

1909.  Sonu'   iiiiilesc-rilKMl   species  of   Mcxiciii    I'liancroganis.   i'roc.   Aincr.    Aciul.    Aits,   Sci., 

44:603-608.  (May)  (Coiitrili.  (ir.i.v  ll(rl.:ii  iiim,  N.S.  No.  36) 
Synopsis  of  tlie  INfexican   and   Central   Aiui'rican   s]iecies  of  Cfisfilleja.  Proc.    Anier. 
Acad.  Arts,  Sci..  44  : .')f;:'.-.";91 .  CMay  i  (Contril..  (iray  Ilcrl.ariinn.  N.S.  Xo.  'M'>  > 

11*11.  liartweg's  Explorations  in  California, — 1;  11;  111;  1\'.  M  iililiiilicrKia,  7:2U-li4;  '25- 
32;  37-43;  49-50.  (Mar./May) 
[Letter]  Die  Zerstorung  der  Academie  dcr  Wissenscliaftfii  in  San  Fraiuisco.  Kihiig- 
lidu's  Oyinnasinni  zn  Soran.  Beilagc  znni  .lalircsheridit.  No.  lUli:lli-12.  Translation 
of  a  letter  to  Dr.  Angust  Brand,  dated  May  14,  190(),  2421  Ridge  Koad,  Berkeley, 
concerning  tlie  destrnetion  of  the  Academy's  Herbarium  in  the  eartlKpiake  anil  fire. 

lit  14.  The  iiiany.anitas  of  Mdinit  Tama  I] mis.  ( 'alifoi  iiia  Out  -of  I  loiirs.  p.  7.  (  A))|-il) 
The  C(<i/iiilli  IIS  itf  Mnuut  Tamalpais.  Califoiiiia  ( )ut-()f- Doors,  j).  4.  (May) 
Natnre  reclotliing  Mount  Tamalpais.  California  Out-of-Doors.  ]>.  19.  (Oct.) 
The  rosaceous  shrubs  of  ^fount  Tainal])ais.  The  Tamaljiais  Magazine,  4:6.  (Nov.) 

l!M.").  K'epiirt  of  the  1  )epai-t  nn'iit  of  Botany  for  the  year   1!I14.  I'roc.  Calif.  Acad.  Sci.,  (4), 
."'):2!t.  (Mar.  i 
How  plants  gidw  on  .Mount  'J'amalpais.  California  Out  i)f-I>oors.  ]>.  110.  (Dee.) 

l!tl(i.  K'ejioif  of  the  Department  of  Botany  for  the  year  1915.  Proe.  Calif.  .\ca(l.  Sci..  (4), 
5:208-209.  (May) 

l!tl7.  Ifeport   of  the  Department  of  Botany  for  the  year  1916.  Proc.  Calif.  .Vcad.  Sci..  (4) 
6:251-25;i  (June) 

1918.  Midwinter  flowers  on  Mount  Tamalpais.  California  Out-of-Doors.  (Jan.) 
The  shrnl)s  covering  Mount  Tamalpais.  California  Out-of-Doors.  (April  i 

K'eport  of  tlie  Department  of  Botany  for  tlie  year  1917.  Proc.  Calif.  Acad.  Sci..  (4), 

7:;'.47-;i49.  (June) 
Late  summer  on   Mount  Tamal]iais.  California  Out-of  Doors.   (Aug.) 

1919.  Kaily  spring  at  the  Grand  Canon  near  El  Tovar.  Plant  World,  22:95-99.  (April) 
Kejiort  of  the  Department  of  Botany  for  the  year  191S.  Proc.  Calif.  Acad.  Sci.,  (4), 

S  :.•{;•,()-.'];?  2.  (June) 

192n.  Peport  of  the  Dt'iiartmeiit  of  Botany  for  the  year   1919.   I'roc  Calif.  .Vend.  Sci.,   (4i, 
9:379-380.  (Aug.) 
Oaks  on  Mount  Ta)n;il]iais.  California  Out-of-Doors.  ji.  182.  (Oct.) 

1921.  Eai'ly  s|ii-iiig  on  .Mcuiiit  Tamaljiais.  ( 'a  lifoiiii  a  Out  <if  1  )(ioi-s.  p.  1  Sli.  (Jan.) 
Tainal|iais  wild  tloweis.  Trails,  I  :.j()-.")4. 

A  plea  for  the  protection  of  wild  flowers.  ( 'aliforiiia  ( )ut  ot'  Doois.  (  .\pi-il  i 
Peport  of  the   Deiiartmeiit  of  P.otany  for  the  year  1920.   I'roc.  Calif.  .\cad.  Sci.,  (4), 
10:137-1:59.  (.luly  ) 

1922.  h'eport  of  the   Department   of  Botany  for  the  yeai-  1!>21.   I'roc.  Calif,  .\cail.  Sci.,   (4), 

n  :()25-(i29.  (Aug.) 

1923.  Winter  flowers  on  Mt.  Tamalpais.  California  Out-of-Doors.  (.T.an. ) 
Trees  of  Mount  Tamalpais  region.  Trails,  2:66-69. 

The  origin  of  some  lociility  and  trail  names  of  Mount  T.imalp.iis.  C.ilitonii.i  Out-of- 
Doors.  ( .\piil  I 
The  fall  flora  of  .Mcuint  Tamalpais.  California  Ont-of-Doors.  (Aug.) 
Peport  of  the  Department  of  Botany  for  the  year  1922.  Proc.  Calif.  .\c;id.  Sci.,  (4), 
11:674-676.  (Aug.) 


Vol.  XXV]  WBITIXGS  OF  ALICE  EASTWOOD  XIX 

19124.  Aix-hiliald  Meuzies  Jourii.il  of  tlie  Vancouver  expedition;  extracts  covering  the  visit 
to  California,  with  an  introduction  and  notes  by  Alice  Eastwood.  California  His- 
torical Society  Quarterly,  2  :  265-340.  (Jan.) 
Report  of  the  Department  of  Rotany  for  the  year  192.'^.  Proc.  Calif.  Acad.  Rci.,  (4), 
12:1258-1260.  (Oct.) 

1925.  I  Review  ofj   "The  Giant  Sequoia,"  by  Rodney  S.  Ellsworth.  Sierra  Chih   I'.ullctiii, 

XII:  204-205. 
The  madrona.  California  ()ut-of-Doors.  (Jan.) 
Sequoia  versus  Eucdli/pfK.s.  California  Out-of -Doors.  (April ) 
Report  of  the  Department  of  Botany  for  the  year  1924.  Proc.  Calif.  Acad.  Sci.,  (4), 

13:467-468.  (May) 
The  rose  family  on  Mt.  Tamalpais.  California  Out-of-Doors.  (July) 

1926.  The  aftergrowth  of  a  niouiitaiii  fire.  California  Out-of-Doors.  (Jan.) 

Report  of  the  Department  of  Hotany  for  the  year  1925.  Proc.  Calif.  Acad.  Sci.,  (4), 
14:536-537.  (April) 

1927.  The  madrona.  Marin  Messenger.  (Feb.) 

Report  of  the  Department  of  Botany  for  the  year  1926.  Proc.  Calif.  Acad.  Sci.,  (4), 

15:528-530.  (Mar.) 
New  species  of  Ceanothus.  Proc.  Calif.  Acad.  Sci.,  (4),  16:361-363.  (Ai)ri] ) 
X^otable  plants  on  Mt.  Tamalpais.  California  Out-of-Doors.  (Sept.) 
ISIarin  County  in  Noveml)er.  Trails,  4:3.  (Nov.) 
Toadstools  and  mushrooms.  Trails,  4:2.  (Dec.) 

1928.  Trees  of  Mount  Tamalpais  region.  California  Out-of-Doors.  (Feb.) 
Midwinter  flowers  on  Mount  Tamalpais.  Trails,  5:2.  (Feb.) 
Violets  of  Marin  County.  Trails,  5:2.  (Mar.) 

Flowers  noted  on  the  trail  to  Little  Carson  on  March  5th.  Trails,  5  :2.  (Mar.) 
More  trees  of  Mt.  Tamalpais.  California  Out-of-Doors.  (Aug.) 

Report  of  the  Department  of  Botany  for  the  year  1927.  Proc.  Calif.  Acad.  Sci.,  (4), 
16:736-738.  (May) 

1929.  The  Eastwood  place  on  Mt.  Tamalpais,  Summit  Avenue,  Mill  Valley.  California  Out- 

of-Doors.  (Apr.) 
Report  of  the  Department  of  Botany  for  the  year  1928.  Proc.  Calif.  Acad.  Sci.,  (4), 

17:321-323.  (May) 
After  a  tire  in  the  Mount  Tamalpais  region.  California  Out-of-Doors.  (Sept.) 
The  escallonias  iu  Golden  Gate  Park,  San  Francisco,  California,  with  descriptions  of 

new  species.  Proc.  Calif.  Acad.  Sci.,  (4),  18  :385-391.  (Sept.) 
Studies  in  the  flora  of  Lower  California  and  adjacent  Islands.  Proc.  Calif.  Acad.  Sci., 

(4),  18:39.3-484.  (Sept.) 

1930.  Report  of  the  Department  of  Botany  for  the  year  1929.  Proc.  Calif.  Acad.  Sci.,  (4), 

18:549-550.  (April) 

1931.  The  true  species  of  fuchsias  cultivated  in  California.   The  National   Horticultural 

Society,  10:100-104,  illus.  (April) 
Report  of  the  Department  of  Botany  for  the  year  1930.  Proc.  Calif.  Acad.  Sci.,  (4), 

19:436-438.  (May) 
New  species  of  plants  from  western  North  America.  Proc.  Calif.  Acad.  Sci.,   (4), 

20:135-160.  (Dec.) 

1932.  The  pittosporums  in  California n  gardens  and  parks.  Leaflets  of  Western  Botany, 

1:1-7.  (Jan.) 
ZjigopliyJlum  Fabdc/o  L.  Leaflets  of  Western  Botany,  1:8.  (Jan.) 
O.vnJis  crrnun  Thunb.  Leaflets  of  Western  Botany,  1:8.  (Jan.) 


XX  CALIF<il;.\I.\   .IC.II>J:M)()F  SCIEXCES  [Proc.  4th  Ser. 

A  new  Califoniian  Montia.  Li-aflets  of  Western  Botany.  1:11.  (April ) 

Gentiana  tenelUi  in  California.  Lcaticts  df  West  c  in  liotany,  1 :1().  (April » 

Another  station  for  Zyyophijlhun  Fabiujo  L.  Lealiets  of  Western  Botany.  1  : 1  (i.  (  April) 

Vfrbenn  honarien.sis  L.  Leatlets  of  Western  Botany,  1 :21.  (Sept.~) 

Cyprip( dium  fasciriilat inn  in  California.  Lealiets  of  Western  Botany.  I:L*i'.   (Sept.) 

Bottle-brushes  in  C.ilitdrni.m  j^r.-irdciis  anil  parks.  Leaflets  of  Western  Botany,  I  :9-ll, 

April  4.  1932;  1:17-19,  Sept.  21.  l!t.!2:   I  :25-2(),  Nov.  18,  l!t:',2:  1:3:^-84,  .Ian.  IP, 

1933  :  1 :45-48,  Mar.  2(i,  1933. 
The  fetid  adder's  tonfjue.  Leaflets  of  Western  Botany,  1  :24.  (Sept.) 
Concerning  Pitfosporinii  (hiiilinipliiilloides.  Leaflets  of  Western  Botany,  I  :24.  (Sept.) 
Tlic  tree  ferns  in  ( ioliirii  { l.-ilc  Park,  liay  Cities  ( la nlcn  .M imt lily,  1  :2.  2  iigs.  (Nov.) 

1933.  The  small  ferns  of  Marin  ( 'diinty.  California  Out-of-Doors.  (Jan.) 

Taxus  brevifoJid  in  Marin  Cduiity,  C.ilifoniia.  Leaflets  of  Western  Botany,  I:38-39l 

(Jan. ) 
A  new  yei-iia  santa.  Leaflets  of  Western  Botany,  1  :4ll-41.  (Jan.) 
.\n  nndescrilied  lily  from  the  Sierra  Nevada  of  California.  Leaflets  of  Western  Botany, 

I  : 41-43.  (Jan.) 
J.I  iliou  as  a  poisonous  jilant.  Leaflets  of  Western  Bot.any,  1:43.  (Jan.) 
A  new  Miniiiliis.  Leaflets  of  Western  Botany,  1 :44.  (Jan. ) 
Flowers  alonjT  the  hij^hwa.vs.  Bay  Cities  Garden  Monthly,  2:2,  8.  (Feb./Mar.) 
Two  new  species  of  w-esfern  CaVnnn.  Leaflets  of  Western  Botany,  1 :5.5-56.  (Mar.) 
A  new  Californian  FritUlaria.  Leaflets  of  Western  Botany,  1 :55.  (Mar.) 
Juniperus  chincnsis  variety  Toruluaa,  an  undescribed  variety  of  Chinese  .juniper.  Bay 

Cities  Garden  Monthly,  2:2,  7.  1  fig.  (April) 
Bottle-brushes  recentl.v  introdvu-ed  into  California.  Leaflets  of  Western  Botany,  1:.57. 

(Aug.) 
New  manzanitas  from  Santa  Cruz  Island,  California.  Leaflets  of  Western  Botany, 

1:61-03.  (Aug.) 
.M;inz;Miitas.  C;i  lifiirnia  Out-of-Doors.  (Sept.) 
Madrone  and  manzanita.  Trails,  10:2.  (Oct.) 

Escallonias  in  Golden  (}ate  Park.  Leaflets  of  Western  Botany,  1 :65-69.  (Nov.) 
New  species  of  California  ArctostupJnjlos.  Leaflets  of  Western  Botany,  1:73-80. 

(Nov.) 

1934.  Chrirantliiis  iiraiiiliflitnis,  a  correction.  I^eaflets  of  Western  l>ot:iny,  1:87—89.   (.Tan.) 
A  kev  to  till'  i-iiniiniin  fiiinilies  of  flow  erinji'  plants  in  Ca  lif  ornin ,  ;inil  A  yuiile  fur  the 

analysis  and  description  of  flowering  jilants.  Prepared  for  and  puldislied  by  the 

Ciilifornia   Botanical  Club,  California  Academy  of  Sciences.  San  Francisco,  The 

.lanu's  H.  Barry  Company,  1934.  16  pp. 
I'lsi-allonias  in  (iolilen  (rate  Park — 11.  Leaflets  of  Western  P>otany,  1  :81-82.  (Jan.) 
Gentiana  Copelandi  Eastwood.  Leaflets  of  Western  Botany,  1 :96.  (Jan.) 
A  revision  of  tlie  genera  formerly  ini-luded  in  ArciostaplniJos.  Leaflets  of  Western 

Botany,  1:97-100.  (Mayj 
A  new  species  of  Solanum.  Leaflets  of  Western  I'.utany,  1 :104.  (May  i 
A  revision  of  Arcinslnpli jilns  with  key  and  descriptions.  Leaflets  of  Western  r.otany, 

1:10.^-127.  (Aug.j 
Glailioliis  .scgetiim    Ker-Gawl.    in    Si ma    County,   California.    Leaflets   of    Western 

Botany,  1:131.  (Nov.) 
New  species  of  Liliaceae.  Leaflets  of  Western  Botany,  1:132-133.  (Nov.) 
A  new  species  of  Tnliumii  from  British  Cohinil)i;i.  L(>;iflets  of  Western  P.otany,  1:139. 

(Nov.) 
New  species  of  2'' /"("I'l"'"'-  Leaflets  of  Western  lintany,  1:139-141.  (Nov.) 


Vol.  XXV]  WRITINGS  OF  ALICE  EASTWOOD  XXI 

19'do.  Lrptospffmum  and  allied  gviiera  in  GuidL'n  Gate  Park.  Leari(.4s  of  Western  Botany, 
1:169-170.  (Mar.) 
New  species  of  Calif ornian  Caslillcja.  Leaflets  of  Western  Botany,  1:174-176.  (Mar.) 
A  new  loco-weed  from  Oregon.  Leaflets  of  Western  Botany,  1 :178-179.  (May) 

1936.  New  lioreal  species  of  Castillejn.  Leaflets  of  Western  Botany,  1 :  193-195.  (Feb.) 
CasiiUcja  multisecta  Eastwood.  Leaflets  of  Western  Botany,  1:195.  (Feb.) 
Adenoplea  madagascariensis.  Leaflets  of  Western  Botany,  1 :197.  (Feb.) 

A  new  Townsendia  from  Utah.  Leaflets  of  Western  Botany,  1 : 206-207.  (Feb.) 

Tlie  most  minute  Mimnhis.  Leaflets  of  Western  Botany,  1:207.  (Feb.) 

The  shrubby  malvastruins  of  California  with  descriptions  of  new  species  and  a  key 

to  the  known  species.  Leaflets  of  Western  Botany,  1 : 213-220.  (April) 
Osmanthus  aurantiacus  (Makino)  Nakai.  Leaflets  of  Western  Botany,  1:224.  (April) 
A  new  Eibes  from  the  Sierra  Nevada.  Leaflets  of  Western  Botany,  1 : 230-231.  (Aug.) 
Malvastnim  fragrans  Eastwood.  Leaflets  of  Western  Botany,  1 :232.  (Aug.) 
Variations  in  Castilleja.  Leaflets  of  Western  Botany,  1:235-238.  (Nov.) 

1937.  New  species  of  Avestern  plants.  Leaflets  of  Western  Botany,  II :  7-9.  (Jan.) 
Quest  for  lilies.  Leaflets  of  Western  Botany,  11:27-29.  (April) 

Xolina  Parry i  in  Ventura  County,  California.  Leaflets  of  Western  Botany,  11:29. 

(April) 
New  species  of  Dodecatheon.  Leaflets  of  Western  Botany,  II  :36-38.  (Aug.) 
New  localities  in  North  America  for  Sphaerophysa  Salsitla.  (Pall.)  PC.  Leaflets  of 

Western  Botany,  11:38.  (Aug.) 
Zjigadenus  fontanus,  a  new  species  from  Mt.  Tamalpais.  Leaflets  of  Western  Botany, 

11:41-42.  (Aug.) 
A  brief  history  of  the  fuchsia.  Journal  of  San  Mateo  Garden  Club.  (Sept. /Oct.) 
Notes  on  Schizococcus,  with  a  key  to  the  species.  Leaflets  of  Western  Botany,  II : 49-50. 

(Nov.) 
New  species  of  western  plants.  Leaflets  of  Western  Botany,  II  :54-56.  (Nov.) 

1938.  The  trees  of  Mt.  Tamalpais.  California  Out-of -Doors.  (Jan.) 

Two  new  wallflowers.  Leaflets  of  Western  Botany,  II :  73-74.  (Jan.) 

A  key  to  the  common  families  of  flowering  plants  in  California,  and  A  guide  for  the 

analysis  and  description  of  flowering  plants.  Prepared  for  and  published  by  the 

California  Botanical  Club,  California  Academy  of  Sciences.  San  Francisco,  The 

James  H.  Barry  Company,  1938.  Second  edition  (revised).  17  pp. 
Excerpt  from  Illustrated  dictionary  of  botanical  terms,  by  John  Lindley,  1848.  With 

introduction  by  Alice  Eastwood.  Eeprinted  1938.  [This  booklet  is  photolithographed 

from  Lindley's  "Introduction  to  Botany,"  Fourth  Edition,  Book  I,  p.  319  and  Book 

III,  pp. 347-383,  346] 
The  cestrums  cultivated  in  California.  Golden  Gardens,  6:8.  (Feb.) 
The  perennial  lupines  of  California,  I.  Leaflets  of  Western  Botany,  II : 81-86.  (April) 
The  tobacco  collected  by  Archibald  Menzies  on  the  northwest  coast  of   America. 

Leaflets  of  Western  Botany,  II : 92-94,  pi.  II,  figs.  1-2.  (April) 
Allium  lacunosum  Wats.  var.  micranthum  Eastwood.  [Footnote  to  "A  Botanical  visit 

to  the  Vancouver  Pinnacles,  by  John  Thomas  Howell."]  Leaflets  of  Western  Botany, 

11:101.  (April) 
Jamrs-ia  americana  var.  caJifornica   (Small)    Jepson.   Leaflets  of  Western  Botany, 

11:102.  (April) 
Two  new  Scrophulariaceae.  Leaflets  of  Western  Botany,  II :  104.  (April) 
New  species  of  Liliaceae.  Leaflets  of  Western  Botany,  II :  109-112.  (July) 
The  yellow-flowered  perennial  lupines  of  the  Pacific  States.  Leaflets  of  Western 

Botany,  11:125-128.  (Nov.) 
A  new  variety  of  Delphinium  calif ornicum.  Leaflets  of  Western  Botany,  11:137-138. 

(Nov.) 


XXII  CALIFORXIA  ACADEMY   (U-SCIKXCES  |  PKOf .  4tii  Ser. 

1939.  rereiiiiial   lupiiius   of  the  I'acilic   Htatcs.   Leatii-ts  of    Wcstcni   iiotaiiv,  11 :  14(]-15U ; 

180-183.  (Feb.,  Apr.) 
A  new  plilo.x  from  Oregon.  Leaflets  of  Western  Botany,  II  :175.  (Feb.) 
New  California  plants.  Leaflets  of  Western  Botany,  II  :18()-188.  (April  i 
Little  known  exdtics  in  Califoi  iii;i.  (Joldi'ii  (jardens,  7  -J).  (May  ) 
Some   unilescribed   nortlifrn  ( 'alifcirnian  valerians.  Leaflets  of  Western    r.dt.'iiiy.   11: 

196-199,  figs.  1-4.  (July) 
Liipiniis  fJanauK  on  Mt.  Dana  :ind  ,it  ad.j.-ieent  loc.-ilities.  Leaflets  of  Western  Botany, 

11:2(11-2(11'.  (.Jnlyj 
Eugenia  »ii/rcianthes.  Leaflets  of  Western  Botany,  II  :"Jl-i-21;l.  (Nov.) 
Two  new  lupines.  Leaflets  of  Western  Botany,  II : 215-216.  (Nov.) 
Eri/simiim  filifoliiiin  Eastwood.  Leaflets  of  Western  Botany.  11:144.  (Nov.) 
Early  botanical  explorers  on  the  Paeifie  Coast  and  the  trees  they  found  there.  C.-ili- 

fornia  Historical  Society  Quarterly,  18:335-340.  (Dec.) 

1940.  The   stercnlias   cultivated    in   California.   Leaflets    (.f    Western    Botany,    11:217-218. 

(Jan.) 
New  s])ecies  of  Liipinu.s.  Leaflets  of  Western  Botany,  1 1  :226-228.  (Jan.) 
Studies  in  Ca.stiUrJa.  1.  Ca.stilleja  in  the  Marble  Mountains,  Siskiyou  County,  Cali- 
fornia.  2.   Miscellaneous  new  species.   Leaflets   of   Western   Botany,   11:241-245. 
(April) 
Tlie  Lupiinis  Bmrcri  aggregate.  Leaflets  of  Western  Botany,  II  :249— 253.  ( Ai>ril ) 
New  western  plants.  Leaflets  of  Western  Botany,  II : 263-209.  (July) 
New  western  plants— II.  Leaflets  of  Western  Botany,  1 1  :281-285.  (Nov.) 
Report  of  the  Dej)artment  of  Botany  for  the  years  1931,  1932.  Proc.  Calif.  Acad.  Sci., 

(4),  20:5(17-508.  (Dec.) 
Carl  Purdy.  Herbertia,  7:33-34. 

1941.  Late  (lowering  exotics  in  the  San  Fr;tncisco   I>ay   Region.  Golden  Gardens,  9:8,  11. 

(Jan.) 

New  western  plants — III.  Leaflets  of  Western  Botany,  111:18-24.  (Feb.) 

The  Lupiiiii.s  hitifoliii.s  aggregate.  Leaflets  of  Western  Botany,  ITT  :41— 43.  (April ) 

On  the  golden  anniversary  of  the  California  Botanical  Club.  Tjeaflets  of  Western 
Botany,  111:2(5-27.  (April) 

Tlie  islands  of  southern  California  and  a  list  of  the  recorded  jilants — T.  L(>aflets  of 
Western  Botany,  TIT  :27-36.  (April  j 

California,  the  land  of  flowers.  12th  Annual  Califiuiiia  Spring  (iardi'ii  Show  Year- 
book, Ajuil  .Id  to  .May  4  (incl.)  1941.  p.  29. 

The  islands  of  southern  California  and  a  list  of  the  i-ecordeil  jil.-ints — II.  I>eaflets  of 
Western  Botany,  Til  :54-78.  (July) 

New  species  of  Ca-stilleja.  Leaflets  of  Western  llotaiiy.  I  1 1  :S7-92.  (Nov.) 

1942.  New  species  of  CastiUiju.  I^eaflets  of  Western  Botany,  111:11  (i-1 1  7.  (Jan.) 

A  botanical  ex|iediti(in  to  Log  S]iring  Ridge.   Leaflets  of  Western   I'.dtany,   111:121— 

126.  (April; 
Two  varieties  of  TriUhia  p(  duncnlan.s.  Leaflets  of  Western  Botany,  111:137-138. 

(April) 
New  western  |il;ints    -IV.  Leaflets  of  Western  P.ot.any,  HI  :  l.')7-l  (id.  (.Inly) 
A  plea  for  preserv.ation  of  a  magnificent  ;izalea  grove  in  Humboldt  County.  National 

League  for  Women's  Service  Magazine,  1():9.  port.,  illus.  (Aug. ) 
New  species  and  varieties  of  Lupiiius.  Leaflets  of  Western  Botany,  111:169-175. 

(Nov.) 
SohiiiK)}!  hiiiridliil lint  in  California.  Leaflets  of  Western  Butjiny,  III  :  1  89-190.  (Nov.) 

194:'..  fiili<i  sdllirold  Kastwood.  Leaflets  of  W^^stern  T')Otany,  111:  199.  (Feb. ) 
New  western  lupines.  Leaflets  of  Western  Botany,  111  :202-204.  (Feb.; 


Vol.  XXV]  WRITIXGS  OF  ALICE  EASTWOOD  XXIII 

A  new  alien  in  California.  Leaflets  of  Western  Botany,  III  :208.  (Feb.) 

I  A  review]  Wild  violets  of  North  America.  National  League  for  Women's  Service 

Magazine,  17:13.  (Mar.) 
Some  veronicas   in   Golden   Gate  Park.   Leaflets  of  Western    Botany,  111:220-222. 

(April) 
Some  alpine  castillejas  of  the  high  Sierra  of  California.  Amer.  Midland  Naturalist, 

30:40-4(1,  1  pi.  (July) 
Variations  in  Castillcja  plagiofnma.  Leaflets  of  Western  Botany,  111:248-249.  (July) 
Two  new  species  from  Baja  California.  Leaflets  of  Western  Botany,  111:257.  (Nov.^ 
A  new  Calif ornian  rose.  Leaflets  of  Western  Botany,  III : 262.  (Nov.) 

1944.  A  collection  of  poimlar  articles  on  the  flora  of  Mount  Tamalpais.  |  San  Francisco], 

Published  l)y  the  autlior,  |  1944]  32  pp.  Reprinted  from  "California  Out-of -Doors," 

the  official  publication  of  the  Tamalpais  Conservation  Club. 
A  new  Manzanita  from  Baja  California.  Leaflets  of  Western  Botany,  IV:4-5.  (Feb.) 
The   botanical  colk'ctions   of  Chamisso   and  Eschscholtz  in  California.  Leaflets  of 

Western  Botany,  IV:  17-21.  (April) 
A  group  of  lupines  related  to  Luphiu.s  omisfitx.  Leaflets  of  Western  Botany,  IV:41- 

43.  (July) 
Why  is  a  Fuchsia  ?  American  Fuchsia  Society.  Bulletin,  no.  44-7  : 3.  (July  ) 
A  new  lupine  from  Mt.  Rainier.  Leaflets  of  Western  Botany,  IV  :43-44.  (Jul}') 
New  western  plants — V.  Leaflets  of  Western  Botany,  IV:63-()4.  (Nov.) 

1945.  Notes  on  Castanopsis,  L'oiiniei/a  and  tlic  sjiring  star-flower.  .lourn.  Calif.  Hort.  Soc, 

6:195-196.  (Jan.) 
New  western  plants — VI.  Leaflets  of  Western  Botany,  IV:  148-150.  (Mar.) 
The  wild  flower  gardens  of  San  Francisco  in  the  1890's.  Leaflets  of  Western  Botany, 

IV:  153-156.  (May) 
New  varieties   of  two  well-known  California  plants.   Leaflets   of  Western  Botanv, 

IV:  166-167.  (May) 
Tentative  key  and  discussion  of  the  lupines  with  spurred  calyx.  Leaflets  of  Western 

Botany,  IV:  186-194,  1  pi.  (Aug.) 
An  account  and  list  of  the  plants  in  the  Brackenridge  Journal.  California  Historical 

Society  Quarterly,  24:337-342.  1  pi.  (Dec.) 

1946.  The  Islay  cherry.  California  Folklore  Quarterly,  V:  102-104.  (Jan.) 

The  lure  of  Arizona  rock-ferns.  American  Fern  Journal,  36  :49-51.  ( Apr./June) 

A  tentative  key  to  the  small-flowered  lupines  of  the  western  United  States.  Leaflets 

of  Western  Botany,  IV: 217-222,  1  pi.  (Jan.)  ;  251-254.  (May) 
The  prostrate  oaks  of  Marin  County,  California.  Leaflets  of  Western  Botany,  IV : 241- 

242.  (May) 
A  new  lupine  from  Bartlett  Mt.  [In  "On  the  botany  of  Bartlett  Mountain,"  by  John 

Thomas  Howell.]  Leaflets  of  Western  Botany,  IV: 259.  (May) 
Notes  on  Arctostaphylos  Uvn-ursi.  Leaflets  of  Western  Botany,  IV:208-269.  (Aug.) 
Species  of  the  Lupiiuis  lepidus  group.  Leaflets  of  Western  Botany,  IV:  281-284. 

(Nov.) 

1947.  A  collection  of  plants  from  the  Aleutian  Islands.  Leaflets  of  Western  Botany,  V:9-13. 

(Jan.) 
Private  herbariums   donated  to  the   California   Academy   of   Sciences.   Leaflets   of 

Western  Botany,  V:4.5-48.  (July) 
Endemism  in  the  flora  of  California.  Contributions  from  the  Gray  Herbarium  of 

Harvard  University,  No.  CLXV: 55-62.  (Oct.) 
Monoecious   junipers    in    Modoc    County,    California.    Leaflets    of   Western    Botany. 

V:72.  (Oct.) 


XXIV  CAI.IFOHMA    ACADEMY   OF  SCIENCES  [Proc.  4th  See. 

1948.  I  Review]   Origins  of  the  flora  of  Califoniia.   15,v   Uoughis   11.  ('jiiindii-ll  ami   1  la   L. 

Wiggins.  Stanford  university  iiublieations  (University  series)  :  Biological  sciences, 

vol.   X,  no.    1.   Stanfniil    T'liivcrsity   Press.    1947.   Pacific    Discovery,    1,   2:IS0-31. 

(March/April) 
[Review]  The  new  world  cyiircsses.  By  Cail  i;.  WHU'  and  Willis  W.  WageiuT.  El  Aliso, 

a  series  of  papers  on  tiie  native  plants  of  California.  Vol.  1.  Rancho  Santa  Ana 

Botanic  Oardm,  A  iialiciiii.  r'alif.  Ainil  1948.  T'acific  Discovery,  1,  4:3]-.'',2.  (.July/ 

August) 
Studies  of  Pacific  coast  lilies — I.  Ijcaflets  of  Western  lJ(daiiy,  ."> :  ln;i-](i4.  (May 
Biitaiiii-al    (-(dk'ctions    |Iii    '"'riic    i-nast    of    iKJrllicast    Greenland    with    liydr(iyia|iliic 

studics  in  the  Greenland  Sea,"  by  Louise  A.  Boyd.]  American  (iroiiraphical  Society 

Special  Publication,  No.  30:270-273.  (July) 
Studies  of  Pacific  coast  lilie.s — II.  Leaflets  of  Western  Botany,  ~> :  120-123.  (Aug.) 
Studies  of  Pacific  coast  lilies— III.  Leaflets  of  Western  Botany,  5 :  133-138.  (Nov.) 
Monocciiiiis  junipci's  in  Tklodnc  T'cjunty.  ("a  lifdi-nia.  Leaflets  of  Western  P>iitaiiy,  .1:72. 

(Oct.  1 

1919.  Luiiinus  Loijliii  Gray,  a  good  species.  Leaflets  of  Western  Botany,  '> :  loo-loi).  (Jan.) 


PROCEEDINGS 

OF  THE 

CALIFORNIA  ACADEMY  OF  SCIENCES 

Fourth  Series 
\'oL.  XXV,  No.  1,  pp.  1-138,  pis.  i-iG  December  11,  1943 


ALICE  EASTWOOD 
SEMI-CENTENNIAL  PUBLICATIONS 

No.  1 

A  REVISION  OF  THE  GENUS  FUCHSIA 
(ONAGRACEAE) 

BY 

PHILIP  A.  MUNZ 
Professor  of  Botany,  Pomona  College 

INTRODUCTION 

FOR  A  LONG  TIME  a  garden  favorite,  the  genus  Fuchsia  seems  in  recent  years 
to  have  stimulated  new  interest  and  enthusiasm,  and  many  new  importa- 
tions of  botanical  species  have  been  propagated  and  are  being  used  for  hybridi- 
zation and  culture.  The  great  variation  in  habit,  in  size,  in  color,  in  climatic 
conditions  to  which  native, — all  add  to  the  possibility  for  use  both  as  an  indoor 
plant,  and  for  milder  climates,  as  a  garden  plant  which  can  be  adapted  to 
hanging  baskets,  ground  cover,  shrub,  vine  or  tree.  With  flowers  ranging  in 
size  from  a  few  millimeters  to  many  centimeters,  through  various  shades  of 
green,  red  to  purple,  it  is  not  surprising  that  it  should  command  attention. 
Strangely  enough,  so  important  a  genus  has  had  no  botanical  revision  since 
many  of  the  species  have  become  known.  For  horticultural  as  well  as  for  botan- 
ical purposes,  it  seems  desirable  to  know  what  the  species  are  and  where  they 
come  from.  To  this  end,  the  present  paper  is  presented. 

Fuchsia  was  first  made  known  to  science  by  a  monk  named  Father  Charles 
Plumier  who  described  a  species  which  he  called  "Fuchsia  triphylla  flare  coc- 
cineo"  in  his  ''Nova  Plantarum  americanarum  Genera,"  p.  14,  tab.  14,  1703. 
Linnaeus  in  1753  based  his  Fuchsia  triphylla  on  Plumier's  description  and 
figure.  For  a  long  time  there  was  confusion  as  to  the  application  of  this  name, 
it  being  used  commonly  for  species  from  Colombia,  such  as  F.  venusta,  but 
now  it  seems  pretty  well  demonstrated  that  Plumier's  plant  came  from  Santo 
Pomingo.  Fuchsia  was  apparently  first  introduced  into  England  about  the  end 

[1] 


2  CALIFORNIA  ACADEMY  OF  SCIENCES  [Proc.  4th  Ser. 

of  the  18th  Century  by  a  sailor  who  grew  it  in  a  mndow  where  it  was  observed 
by  a  niirserj'man  of  Hammersmith,  a  ^Ir.  Lee,  who  succeeded  in  buying  it  and 
propagating  it  for  the  trade.  This  seems  to  have  been  one  of  the  short-tubed 
species  such  as  F.  magellanica  or  coccinea.  Later  introductions  of  long-tubed 
ones  were  crossed  with  the  short  ones  and  our  common  garden  hybrids  have 
been  developed. 

The  name  Fuchsia  was  given  by  Plumier  in  honor  of  Leonhard  Fuchs  (the 
spelling  of  whose  surname  ought  to  help  those  who  have  difficulty  with  the 
order  of  the  letters  in  the  generic  name),  a  German  physician  and  professor 
of  medicine  at  the  University  of  Tiibingen  from  1535  to  1566.  He  was  the 
author  of  a  number  of  botanical  works,  particularly  the  great  herbal  "De 
historia  stirpium"  published  in  1542,  a  Latin  work  dealing  with  about  400 
native  German  and  100  foreign  plants.  Agnes  Arber  in  her  "Herbals,"  page 
60,  1912,  writes  "Of  all  the  botanists  of  the  Renaissance,  Fuchs  is  perhaps  the 
one  who  deserves  most  to  be  held  in  honor."  She  rates  him  as  superior  in 
scholarship,  critical  study  of  the  nomenclature  of  classical  authors,  and  in 
botanical  illustration.  She  feels  that  in  his  work  is  reached  "the  high-water 
mark  of  that  type  of  botanical  drawing  which  seeks  to  express  the  individual 
character  and  habit  of  each  species,  treating  the  plant  broadly  as  a  whole,  and 
not  laying  more  stress  upon  the  reproductive  than  the  vegetative  organs."  It  is 
no  mean  name  that  is  perpetuated  in  Fuchsia. 

On  account  of  the  great  historical  interest  of  Plumier's  volume  and  its 
rarity,  photographs  of  the  title  page,  pages  14  and  15,  containing  the  original 
description  of  the  new  genus  Fuchsia,  and  plate  14  with  its  first  published 
illustration  are  reproduced  on  Plate  16  of  the  present  account. 

In  its  fleshy  fruit  the  genus  Fuchsia  differs  from  all  other  Onagraceae;  in 
this  respect  and  in  possessing  a  corolloid  calyx  and  colored  hypanthium  it 
constitutes  a  distinct  tribe  (Raimann,  in  Engler  and  Prantl',  Die  nat.  Pflanzen- 
fam.  Ill,  7  :219,  1898) .  I  can  see  no  very  close  relationship  to  any  of  the  other 
genera  in  the  family.  Fuchsia  is  an  immensely  interesting  group  in  the  beauty 
and  variety  of  its  species,  in  the  development  of  taxonomic  differentiation,  and 
in  geographical  distribution.  In  this  paper  I  am  recognizing  about  100  species 
without  attempting  to  go  into  horticultural  and  hybrid  forms.  I  frankly  admit 
that  in  many  cases  inadequate  material  or  inaccessibility  of  types  because  of 
the  present  world  conditions  will  undoubtedly  make  necessary  a  change  of 
concept  in  the  future.  But  the  tremendous  advantage  that  has  been  given  to 
the  North  American  student  of  South  American  plants  by  the  great  accumula- 
tion of  photographs  and  fragments  of  types  at  the  Field  Museum  through  the 
long  years  of  effort  by  J.  Francis  Macbride,  now  makes  it  possible  to  come  to 
conclusions  concerning  species  described  long  ago,  which  could  not  have  been 
reached  earlier  without  work  in  many  European  herbaria. 

In  the  preparation  of  this  paper  I  have  had  from  time  to  time  access  to 
various  herbaria  in  Europe  and  South  America,  as  well  as  in  the  United  States, 
and  have  more  recently  borrowed  some  or  all  of  the  specimens  from  a  numbed- 


Vol.  XXV] 


MUNZ:  THE  GENUS  FUCHSIA 


of  the  last  named.  To  the  curators  of  these  institutions,  to  the  John  Simon 
Guggenheim  Memorial  Foundation  which  made  possible  the  journey  to  South 
America,  to  Dr.  LAnnan  B.  Smitli  of  (Jray  Herbarium  for  notes,  and  to  my  wife, 
Alice  ]\IcCulIy  Munz,  wlio  did  extensive  library  work  on  Fuchsia,  I  liereby 
express  my  appreciation.  I  submit  the  folloAving  list  of  herbaria  with  the 
abbreviations  which  are  used  in  the  citation  of  specimens : 

Botanisches  Museum,  Berlin-Daldem  (B) 

Bishop  Museum,  Honolulu  (BISH) 

Bailey  Ilortorium,  Ithaca,  N.  Y.  (BH) 

California  Academy  of  Sciences,  San  Francisco  (CAS) 

Field  Museum,  Chicago  (F) 

Lillo  Herbarium,  Tucuman,  Argentina  (LID 

Gray  Herbarium,  Harvard  University  (Gil) 

University  of  Michigan  ( MICH ) 

iMissouri  Botanical  Garden  (MO) 

New  York  Botanical  Garden  (NY) 

Jardin  des  Plantes,  Paris  (P) 

Pomona  College,  Claremont,  California  (POM) 

Museu  Nacional,  Rio  de  Janeiro  (R) 

Museo  Nacional,  Santiago,  Chile  (SGO) 

El  Darwinion,  San  Isidro,  Argentina  (SI) 

Sao  Paulo,  Brazil  (SP) 

Ignited  States  National  Herbarium  (US) 

Dominion  Museum,  Wellington,  New  Zealand  (WELT). 

Taxonomic  Characters  in  the  Genus  FUCHSIA 

For  the  most  part  the  characters  which  I  am  using  are  the  traditional  ones : 
presence  or  absence  of  petals;  proportions  of  the  hypanthium;  sepals  erect, 
divergent,  spreading  or  reflexed ;  leaves  opposite,  alternate  or  whorled ;  pu- 
bescence ;  flowers  erect  or  not ;  stamens  included  or  exserted ;  shape  of  berry; 
pubescence  on  style;  shape  of  leaves ;  length  of  petioles,  pedicels,  etc. ;  position 
of  flowers,  whether  solitary  or  axillary  or  in  inflorescences ;  number  of  veins  in 
leaves ;  color  of  various  parts ;  stipules.  Since  the  genus  has  never  been  keyed 
out  nor  anything  like  a  complete  treatment  made  of  it,  I  have  had  to  use  pretty 
much  my  own  judgment  in  choice  of  characters  and  have  attempted  to  select 
such  as  could  be  seen  with  the  least  difficulty.  All  measurements  and  propor- 
tions have  of  course  been  taken  from  pressed  specimens  and  this  fact  must  be 
kept  in  mind  if  one  tries  to  identify  fresh  material  with  my  keys. 

Relationships  Within  the  Genus 

I  should  think  the  species  included  in  the  section  Qucliisio  might  he  taken 
as  the  more  primitive  ones  in  the  genus.  The  short  hypanthium,  erect  stamens, 
many-seeded  fruit,  mostly  spreading  sepals  seem  characteristic  onagraeeous 
features.  Of  the  several  species,  F.  magcllanica  with  its  sepals  quite  free  seems 


4  CALIFOBXIA  ACADEMY  OF  SCIEXCES  [Proc.  -Ith  Ser. 

primitive.  The  range  of  Quelusia,  with  F.  magellanica  in  southern  Argentina 
and  Chile  and  with  the  other  species  occurring  in  the  mountains  of  south- 
eastern Brazil,  is  an  interesting  one,  and  entirely  different  from  all  the  rest  of 
the  genus.  The  Brazilian  species  have  the  sepals  more  or  less  connate  at  the 
base. 

The  section  Eiifuchsia  is  the  most  inclusive  one  in  the  genus.  To  me  its  most 
primitive  species  is  F.  splendens  of  Mexico,  which  seems  near  to  Quelusia  in 
its  long  exserted  stamens,  but  has  too  long  a  hypanthium  which  is  also  peculiar 
in  being  very  wide  at  the  base.  This  species  resembles  more  closely  the  section 
EufuoJisia  than  Quelusia  in  its  large  leaves.  From  it  there  can  be  arranged  two 
series  of  species :  one  with  long  hypanthium,  the  second  with  shorter  one.  Both 
series  run  from  solitary  axillary  flowers  to  well  defined  inflorescence.  In  the 
first  series  I  would  place  nearest  to  F.  splendens  the  Guatemalan  F.  cordifolia, 
in  which  the  hypanthium  has  a  broad  base  and  the  leaves  are  wide,  but  'the 
hypanthium  is  longer  and  the  stamens  less  exserted.  From  this  through  F. 
austromontana  of  Peru  there  can  be  derived  a  group  with  subcylindric  hj'pan- 
thium  rather  wide  at  the  base,  but  with  narrow  petals :  F.  denticulata,  F.  lepto- 
poda,  F.  Woytkowskii,  and  F.  macjelalenae.  Another  group  with  wider  petals 
and  somewhat  less  broad  hypanthium-base,  but  continuing  the  subcylindric 
condition  contains:  F.  canescens,  F.  rivularis,  F.  loxensis,  F.  pallescens,  F. 
Townsendii,  F.  platypetala,  and  F.  macrostigma.  Then  comes  a  series  with 
more  ampliate  hypanthium  and  wide  petals :  F.  ayavacensis,  F.  Pringsheimii, 
and  F.  tripliylla;  these  have  axillary  flowers.  With  narrower  petals  and  ampli- 
ate hypanthium  and  with  flowers  axillary  or  subterminal  we  have :  F.  petio- 
laris,  F.  Smithii,  F.  Lleivelynii,  F.  Asplundii,  F.  venusta,  F.  Jahnii,  F.  Gehri- 
geri,  F.  simplicicaulis,  and  F.  confertifolia.  "With  the  hj-panthium  narrow  and 
long,  and  with  the  flowers  in  a  short,  terminal,  definite  inflorescence  and  with 
leaves  large,  are  :  F.  Asinazui,  F.  tincta,  F.  Mathewsii,  F.  Fischeri,  F.  Storkii, 
F.  furfuracca,  F.  Munzii,  F.  hirtella,  F.  polyaniha,  F.  corymhiflora,  F. 
ahrupta,  F.  Cuatrecasasii,  F.  Killipii,  F.  hoUviana,  and  possibly  F.  fulgens. 

The  other  great  series  which  I  would  derive  from  the  cordifolia  type  has  a 
short  hypanthium.  Here  again  one  can  begin  with  axillary  flowers  with  rather 
wide  hypanthium-base,  such  as  in  F.  decussata  and  F.  Jiypoleuca,  and  then  go 
to  species  with  a  narrower  base,  such  as  F.  scahriuscula  and  F.  verrucosa  and 
F.  sanctae-rosae,  this  last  named  having  either  axillary  flowers  or  almost  ter- 
minal inflorescence.  From  it  one  passes  very  easily  into  a  large  group  with 
definitely  terminal  inflorescence :  F.  Osgoodii,  F.  Andrei,  F.  Lehmannii,  F. 
putumayensis,  F.  Hartwegii,  F.  ovalis,  F.  asperifolia,  F.  pilosa,  F.  glaherrima, 
F.  macrophylla,  F.  sessilifolia,  and  F.  sylvatica.  These  species  of  Eu fuchsia 
are  almost  entirely  Andean,  excepting  F.  splendens,  F.  cordifolia,  and  F. 
fulgens  of  Mexico  and  Central  America,  and  F.  triphylla  and  F.  Pringsheimii 
of  the  West  Indies. 

The  section  Kierschlege.ria  of  Chile  with  its  single  species  F.  lycioides  is  an 
interesting  xerophyte  which  must  be  related  to  7^.  magellanica  in  its  short 


Vol.  XXV]  MUNZ :  TEE  GENUS  FUCHSIA  5 

hypanthium  and  small  flowers.  I  should  suppose  too  that  the  section  Skinnera 
of  New  Zealand  and  Tahiti  with  small  flowers  and  reflexed  sepals  is  of  similar 
relationship,  that  is  to  the  Patagonian  magellanica.  In  its  reduced  or  even 
obsolete  petals  it  becomes  a  very  distinct  group  that  must  have  been  long 
isolated. 

The  same  tendency  to  reduction  or  suppression  of  petals  is  seen  in  a  long- 
tubed  Andean  series  undoubtedly  related  to  Eufuchsia  rather  than  to  Quelusia 
and  for  which  I  propose  the  section  Hemsleyella.  The  species  I  have  included 
here  are  doubtless  not  all  of  the  same  origin.  In  fact,  I  doubt  that  F.  cestroides 
should  be  included  with  the  others.  They,  in  turn,  are  partly  of  the  type  with 
cylindrical  hypanthium,  for  example,  F.  macrantha  and  F.  Garlepyiana,  or 
with  ampliate  hypanthium  as  in  F.  apetala.  In  addition  to  the  suppression  of 
petals,  specialization  is  evidenced  by  such  features  as  union  of  sepals  at  their 
base,  loss  of  leaves  at  anthesis,  etc. 

Perhaps  the  most  highly  specialized  section  is  Encliandra  which  occurs 
from  Panama  to  central  Mexico.  Small-flowered,  with  small  leaves,  and  the 
shorter  series  of  stamens  bent  back  into  the  hypanthium,  this  group  also  tends 
in  A'arying  degrees  to  polygamy.  It  has  also  relatively  few  seeds  in  the  berry. 
It  may  be  perhaps  related  to  the  section  SoJiufia  (with  erect  flowers  in  large 
compound  inflorescence)  of  the  same  geographical  area  in  its  small  flowers 
and  relatively  short  hypanthium.  Just  how  these  two  groups  are  related  to 
the  rest  of  the  genus  is  not  clear ;  perhaps  the  short  hypanthium  ties  them  in 
with  the  apparently  rather  primitive  Quelusia  which  must  have  come  from 
common  stock  with  them. 

THE  GENUS  Fuchsia  Linnaeus 

Fuchsia  L.,  Gen.  PI.,  ed.  1,  27,  1737;  ed.  5,  126,  1754,-  Sp.  PL,  ed.  1,  1191,  1753.  Endlicher, 
Genera  PL,  1193,  1840.  Benth.  and  Hook.,  Gen.  PL,  1,  part  3:  790,  1862.  Raimann  in 
Engler  and  Prantl,  Die  nat.  Pflanzenfam.,  Ill,  7:219,  1898. 

Hypanthium  prolonged  beyond  the  ovary,  usually  colored,  deciduous  in 
fruit.  Sepals  4,  usually  colored,  deciduous.  Petals  4,  or  minute,  or  0,  convolute 
or  spreading,  deciduous.  Stamens  8,  usually  unequal,  the  episepalous  exceed- 
ing the  epipetalous ;  filaments  more  or  less  filiform ;  anthers  linear  to  oblong. 
Pollen  grains  single,  2-  or  3-angled.  Ovary  4-loculed ;  style  elongate ;  stigma 
capitate  or  clavate,  subentire  to  4-lobed.  Suffrutescent  to  arboreous.  Leaves 
opposite  to  alternate  or  whorled,  with  small  usually  deciduous  stipules.  Flowers 
solitary  in  the  axils  or  racemose  or  paniculate,  mostly  pendulous  and  long- 
pedicelled,  generally  showy ;  perfect  to  polygamous. 

Type  species,  F.  triphylla  L. 

Key  to  Sections  of  FUCHSIA 

A.  Petals  lacking  or  reduced  to  small  scalelike  structures;  leaves  tending  to  be  alternate. 

B.  Hypanthium  infundibuliform,  not  more  than  1.5  cm.  long ;  sepals  reflexed,  separate. 

New  Zealand  and  Tahiti 4.  Slciimera. 


6  CALIFOEXIA  ACADEMY  OF  SCIENCES  [Proc.  4th  Ser. 

BB.  Hypanthium  tubular  to  funnelforni,  mostly  more  than  2  cm.  long;  sepals  divergent 

to  suberect,  more  or  less  connate  at  base.  South  America 5.  Hemsleyella. 

AA.  Petals  present,  almost  or  quite  as  long  as  sepals;  leaves  mostly  opposite  or  whorled. 

B.  Leaves  with  a  thickened  persistent  petiole-like  base  which  becomes  spine-like ; 

sepals  reflexed  ;  flowers  small.  Chile 3.  Eierschlegeria. 

BB.  Leaf-bases  not  persisting  as  spine-like  structures. 

C.  Flowers  erect  in  terminal  cymose  panicles  arranged  subtrichotomously.  Mexico 

to  Panama 6.  Schufia. 

CC.  Flowers  mostly  pendulous,  not  in  cymes. 

D.  Flowers  tending  toward  polygamy,  small,  inconspicuous ;  stamens  short, 
the  epipetalous  reflexed;  berry  few-seeded.  Mexico  to  Panama. 

7.  Encliandra. 
DD.  Flowers  perfect,  usually  showy;  stamens  more  or  less  exserted,  erect; 
berry  many-seeded.  Mostly  South  American  species. 
E.  Hypanthium   usually   not   longer   than   the   sepals ;    stamens   long- 

exserted.  Eastern  Brazil  to  Patagonia 1.  Quelusia. 

EE.  Hypanthium  several  times  as  long  as  sepals ;  stamens  not  much  ex- 
ceeding sepals.  Andes  north  of  central  Argentina  to  Mexico  and  West 
Indies 2.  Eufuchsia. 

Section  1.  QUELUSIA  (Vand.)  DC. 

Quelusia  as  genus,  Vandeli,  F1.  Lusit.,  23,  pi.  10,  1788;  and  in  Eoem.,  Script.  Hisp.  Lusit., 

101,1796. 
Nahusia  Schneev.,  Ic,  27,  1793. 
Thilcum  Molina,  Sagg.  Chile,  ed.  2, 146, 1810. 
Quelusia  as  sect.,  DeCandolle,  Prodr.,  3 :36, 1826. 

Nectary  fused  to  base  of  hypanthium ;  hypanthium  eylindric,  usually  no 
longer  than  sepals ;  stamens  long,  exserted. 

Type  species :  Vandeli  described  the  genus  without  naming  a  species.  De 

Candolle  has  F.  coccinea  under  Quelusia,  as  well  as  other  species ;  he  included 

F.  magellanica  as  a  synonym  of  F.  coccinea.  Since  Vandeli  referred  to  petioled 

leaves,  he  evidently  had  in  mind  F.  magellanica  Lam.,  and  I  would  make  it  the 

type  species. 

Key  to  Species  of  Section  QUELUSIA. 

A.  Hypanthium  short,  3-6  mm.  long ;  younger  branches  pubescent  to  pilose. 

B.  Leaf -blades  4-10  cm.  long,  2-4  cm.  wide,  quite  entire 6.  F.  regia  var.  aJpcstris. 

BB.  Leaf -blades  1.5-15  cm.  long,  0.3-2  (2.5)  cm.  wide,  mostly  serrulate. 

C.  Young  growth  strigulosc ;  leaves  mostly  less  than  0  mm.  wide,  coriaceous. 

Itatiaya,  Brazil l.F.  Campos-Portoi. 

CC.  Young  growth  puberulent  to  pilose ;  leaves  mostly  more  than  6  mm.  wide, 

membranous. 

D.  Suffrutescent,  with  herbaceous  stems  2-5  dm.  tall;  pedicels  1-2  cm.  long; 

stems  very  densely  shaggy-pilose.  Espiritu  Santo,  Brazil.  2.  F.  Bracelinae. 

DD.  Well  developed  shrub,  1-3  m.  tall ;  pedicels  2-4  cm.  long ;  steins  puberulent 

to  somewhat  pilose.  Minas  Geraes,  Brazil .A.F.  coccinea. 

AA.  Hypanthium  longer,  mostly  8-20  mm.  long;   younger  branches  usually  glabrous  or 
nearly  so. 
B.  Leaf -blades  subcntire,  4-10  cm.  long,  2-4  cm.  Avide;  sepals  connate  at  base  for 

2-15  mm.  So.  Brazil Q.F.  regia. 

BB.  Leaf -blades  serrulate,  1-5  cm.  long ;  sepals  mostly  free. 


Vol.  XXV]  MUNZ :  THE  GENUS  FUCHSIA  7 

C.  Petioles  mostly  3-10  mm.  long ;  hypantliium  8-10  (12)  mm.  long ;  sepals  15-25 

mm.  long,  3-4  mm.  wide.  So.  Chile  and  So.  Argentina 3.  F.  magellanica. 

CC.  Petioles  mostly  10-25  mm.  long;  hypanthium  10-20  (or  more)  mm.  long; 
sepals  25-80  mm.  long,  8-11  mm.  wide.  Gardens;  naturalized  as  in  Colombia, 
Venezuela 5.F.  hybrida. 

(1)  Fuchsia  Campos-Portoi  Pilger  and  Schulze 

(Plate  1,  fig.  1) 
Fuchsia  Campos-Portoi  Pilger  and  Schulze,  Notizblatt,  12  : 470,  1935. 

Shrub,  with  woody  ascending  or  divaricate  branches,  the  younger  ones  very 
slender,  strigulose,  more  or  less  reddish ;  leaves  opposite  or  ternate  or  quater- 
nate,  elliptic-lanceolate,  acute  at  both  ends,  plainly  serrulate  with  incurved, 
gland-tipped  teeth,  coriaceous,  strongly  bicolored,  glabrous  above  or  strigulose 
on  midrib,  strigulose  beneath  especially  on  midrib  and  margins,  lateral  veins 
inconspicuous,  about  5  on  each  side  of  midrib;  blades  1.5-2.5  (3)  cm.  long, 
3-5.  (8)  mm.  wide;  petioles  2-3  (4)  mm.  long,  puberulent;  stipules  subulate, 
0.5  mm.  long,  deciduous;  flowers  solitary  in  upper  axils;  pedicels  filiform, 
strigulose,  4r-15  mm.  long ;  ovary  4-5  mm.  long ;  hypanthium  red,  broadly 
fusiform,  4-5  mm.  long,  sparsely  strigulose  without,  glabrous  within ;  sepals 
red,  somewhat  strigulose,  oblong-lanceolate,  acuminate,  18-20  mm.  long,  4-6 
mm.  wide,  connate  for  2-3.5  mm.  at  base ;  petals  violet,  obovate-cuneate, 
slightly  emarginate,  10-12  mm.  long;  episepalous  stamens  20-25  mm.  long, 
alternate  ones  slightly  shorter ;  anthers  1.5  mm.  long;  style  glabrous,  ca.  3  cm. 
long ;  stigma  subentire,  ca.  2  mm.  long ;  fruit  ellipsoid,  ca.  1  cm.  long. 

From  Itatiaya,  Serra  da  Mantiqueira,  state  of  Rio  de  Janeiro,  Brazil,  where 
it  grows  at  2100-2300  m.  alt.  Material  seen,  Pilger  and  Brade  31,  type  coll. 
Dec.  27, 1934  (POM,  R) ,  Brade  15676,  March,  1934  (POM,  R) ,  Loefgren  5775, 
March  13, 1903  (POM,  SP) .  I  am  greatly  indebted  to  Dr.  Brade  of  the  Botani- 
cal Garden  at  Rio  de  Janeiro  for  information  concerning  and  material  of 
this  species.  The  species  is  characterized  by  its  narrow  leaves,  woody  habit, 
strigulose  younger  growth,  and  fascicled  appearance  of  the  short  branches. 

(2)  Fuchsia  Bracelinae  Munz,  new  species 
(Platel,  fig.  2) 

Suffrutescent,  with  running  underground  rootstock  and  erect,  rather 
simple  stems  2-5  dm.  tall,  woody  and  with  exfoliating  epidermis  at  base ;  stems 
herbaceous,  apparently  reddish  and  densely  pilose  with  light-colored  hairs  up 
to  1  mm.  long ;  leaves  opposite,  in  3's  or  4's,  ovate-lanceolate,  green  and  some- 
what strigose  above,  paler,  reddish  and  pilose  beneath  especially  on  veins, 
rounded  at  base,  acute-acuminate  at  apex,  subentire  to  denticulate,  membran- 
aceous, distinctly  veined  (main  laterals  6-8  on  each  side  of  midrib)  ;  blades 
2-5  cm.  long,  5-14  mm.  wide ;  petioles  1-3  mm.  long,  villous ;  stipules  lance- 
deltoid,  0.5-1.5  mm.  long,  deciduous ;  flowers  axillary  ;  pedicels  pilose,  filiform, 
1-2  cm.  long;  ovary  oblong,  pilose,  5  mm.  long;  hypanthium  dark  red,  pilose 


8  CALIFOBNIA  ACADEMY  OF  SCIEXCES  [Proc.  4th  Ser. 

without,  glabrous  within,  3.5-5  mm.  long,  2-2.5  mm.  wide  at  slightly  expanded 
summit ;  sepals  rose,  19-23  mm.  long,  sparsely  pilose,  connate  at  base  for  3-4 
mm.,  then  erect-divergent,  lance-oblong  or  -elliptic,  subacuminate,  5-6  mm. 
wide,  3-veined;  petals  purple,  obovate,  11-15  mm.  long,  rounded  apically; 
episepalous  stamens  up  to  3  cm.  long,  epipetalous  ca.  3  mm.  shorter ;  filaments 
reddish ;  anthers  less  than  2  mm.  long,  reddish ;  style  reddish,  up  to  3.5  cm. 
long,  glabrous;  stigma  slightly  lobed,  narrow,  1.5  mm.  long;  immature  fruit 
oblong,  8  mm.  long,  4  mm.  wide. 

Suffrutescens;  caulibus  erectis,  2-5  dm.  altis,  herbaceis,  pilosis;  foliis  oppositis,  ternatis 
aut  quaternatis,  ovato-lanceolatis,  in  superficie  superiore  strigosis,  inferiore  pilosis,  base 
rotundatis,  apice  acuto-acuminatis,  subintegris  vel  denticulatis ;  laminis  2-5  cm.  loiigis, 
0.5-1.4  cm.  latis ;  petiolis  villosis,  1-3  mm.  longis ;  pedicellis  axillaribus,  pilosis,  filiformibus, 
1-2  cm.  longis;  ovario  oblongo,  piloso,  5  mm.  longo;  hypanthio  3.5-5  mm.  longo,  supra 
2-2.5  mm.  lato,  externe  piloso,  interne  glabro;  sepalis  roseis,  19-23  mm.  longis,  sparse 
pilosis,  lanceolato-oblongis  aut  -ellipticis,  subacuminatis,  5-6  mm.  latis;  petalis  purpureis, 
obovatis,  11-15  mm.  longis;  staminibus  2.5-3  cm.  longis,  rubris;  stylo  glabro,  3-3.5  cm. 
longo ;  bacca  oblonga,  8  mm.  longa,  4  mm.  lata. 

Type :  from  rocky,  open  campo,  Serra  de  Caparao,  State  of  Espiritu  Santo, 
Brazil,  Nov.  25,  1929,  at  2650  m.,  Ynez  Mexia  4013,  Gray  Herb.,  isotypes 
(CAS,  NY,  US) ;  distributed  as  F.  pulesccjis.  Distinct  from  F.  Campos-Portoi 
and  F.  coccinea  in  its  more  herbaceous  habit ;  from  the  former  by  its  broader 
thinner  leaves  and  more  hair ;  from  the  latter  by  its  narrower  leaves.  It  is  a 
pleasure  to  dedicate  this  species  to  Mrs.  H.  P.  Bracelin  of  Berkeley,  California, 
who  has  devoted  much  time  and  energy  to  distributing  the  large  collections 
made  in  Latin  America  by  Mrs.  Mexia. 

(3)  Fuchsia  magellanica  Lam. 

Fuchsia  magellanica  Lam.,  Encycl.,  2:5C5,  1788   ?   (cf.  Kuntze,  Eev.  Gen.  PL,  l:cxxxii, 
1891);  pi.  282,  fig.  2,  1793. 

Shrub  up  to  2.5  m.  tall,  bushy  or  semiscandent,  freely  branched,  the  ultimate 
branchlets  very  slender  (ca.  1  mm.  thick),  usually  reddish,  glabrous  to  finely 
and  sparsely  puberulent,  especially  about  the  nodes,  older  ones  exfoliating ; 
leaves  opposite  or  ternate,  sometimes  alternate,  usually  rather  crowded,  the 
blades  lance-ovate  to  rhombic-ovate  to  elliptic-ovate,  rounded  or  more  fre- 
quently somewhat  cuneate  at  the  base,  acute  to  acuminate  at  the  apex,  usually 
evidently  serrulate,  more  or  less  reddish  on  veins,  paler  beneath  than  above, 
glabrous  to  puberulent  on  margins  and  veins  of  under  surface,  15-50  mm.  long, 
■4-20  mm.  wide,  with  5-7  principal  veins  on  each  side  of  midrib;  petioles  3-10 
(15)  mm.  long,  glabrous  to  puberulent;  stipules  lance-deltoid,  barely  1  mm. 
long,  deciduous ;  flowers  1  or  2  in  upper  axils ;  pedicels  filiform,  2-5.5  cm.  long, 
glabrous  to  puberulent;  ovary  oblong-fusiform,  6-10  (12)  mm.  long,  1.5-2.5 
mm.  wide;  hypanthium  tubular,  8-10  (12)  mm.  long,  glabrous  or  nearly  so 
Avithout,  pubescent  within;  sepals  narrowly  lance-oblong,  15-25  mm.  long, 
3-4  (5)  mm.  wide,  acuminate,  mostly  glabrous;  petals  obovate,  1-2  cm.  long; 
longer  stamens  2-3.5  cm.  long,  epipetalous  4-6  mm.  shorter ;  filaments  filiform, 


Vof.  XXV]  MISZ:   Tilt:  CKMS  FICIISIA 

;rlabrous ;  anthers  2-2.5  mm.  long;  style  slemiiT,  i)ubi*.scent  at  ba.se,  mostly  3-5 
fin.  loiifj;  stijjrma  capitate,  more  or  less  lobeil,  1.5-3  mm.  long;  berry  oblong, 
1-2  cm.  long,  4-5  mm.  wide. 

This  species  is  exceedingly  varial)l(',  with  a  noiiiciu  lature  much  complicated 
by  long  culture  in  the  garden  and  much  hybridization.  Many  segregates  have 
been  prctposed  and  many  attempts  have  been  made  to  classify  such  (cf.  IMiil- 
ippi.  Bot.  Zeit.,  34:577,  187G;  Bailey,  Cyclop.  Am.  Ilort.,  3:1301,  1015,  for 
exami>les).  In  the  study  of  wild  plants  the  most  that  I  can  (ibserve  is:  (1)  a 
tendency  for  larger  flowers,  larger  thinner  leaves,  and  longer  pedicels  in  the 
more  northern  material,  (2)  opposite  tendency  in  more  southern  material  and 
(3)  a  possible  less  highly  colored  variety  described  by  Prof.  Espiuo/a.  The 
complication  involving  hybridization  with  F.  cocciiiea  and  other  species 
cannot  be  discussed  here. 

Key  to  Varieties  of  Fuchsia  magcUanica 

A.  Hvpnnthiiim  and  scpnls  deep  rod;  ovary  and  pedicels  red. 

B.  Leaf  blades  mostly  15-25  mm.  long;  pedicels  20-.1.J  mm.  long 3a.  var.  typica. 

r>B.  LcafMades  mostly  25-50  mm.  long;  pedicels  40-55  mm.  long.  31).  var.  marront'     <:. 
\A.  Ilvp.nntliium  and  sepals  pink;  ovary  and  pedicels  green He.  v.ir.  V"/         . 

(3a)  Fuch.sia  magellanica  Lam.  var.  typica  Munz,  new  name 

(Plate  1,  fig.  3) 
F.  mageJlanica  Lam.,  Encycl.,  2:565,  1788;  pi.  2S2,  fig.  2,  1793. 
F.  gracili.i  var.  muUiflora  Lindl.,  Bot.  Reg.,  13:pl.  1052,  1827. 
F.  gracilis  var.  icnella  LiNDL.,  I.e.;  F.  macrostema  var.  toicUa  DC.  Prodr..  3:38,  1828:  F. 

lencUa  Ilort.,  0.  Don  in  Loud.,  Hort.  Brit.,  155, 1830. 
F.  conica  Lindl.,  Bot.  Reg.,  13:pl.  1062,  1827 ;  F.  macrostemma  var.  conica  SwE»rr,  Brit.  Fl. 

Garden,  6:  under  pi.  216,  1833;  F.  magellanica  var.  conico  Bajley,  Cyclop.  .\m.  Ilort., 

614. 1900. 
f  F.  multiflora  LoDD.,  Bot.  Cab.,  15  :pl.  1415,  1828. 
F.  discolor  Lindl.,  Bot.  Reg.,  21  :pl.  1805, 1835 ;  F.  magcUanica  var.  discolor  Bailey,  Cyclop. 

Am.  Ilort.,  614,  1900. 
F.coccinea  var.  rohustior  UooK.,  Fl.  Ant.,  2:269,  1^47;  l>:isi<l  •m  ( '..iimiirscni  .•oil.  in  Sir:iif 

of  Magnlhacns. 
F.  chonotica  Phiu,  Linnaea,  28:687,  1856;  Bot.  Zeit.,  34:577,  figs.  4  and  5,  1«76;  F.  eoe- 

cinea  var.  rhnnotica  Reiche,  Anal.  Univ.  Chile,  98:485,  1897;  Fl.  Chile,  2:267,  1898. 
F.araucana  Piiiu,  Bot.  Zeit.,  34:577,  fig.  6,  1876. 
Dnrvalla  cucharis  CoMM.  ex  Lam.,  Encycl.,  2:565,  1788. 

Leaf-blades  mostly  15-25  mm.  long,  4-10  (12)  mm.  wide,  somewhat  cori- 
aceous, often  folded  along  the  midrib;  pedicels  mostly  2-3.5  cm.  long,  red; 
hypanthium  and  sepals  deep  red;  .sepals  1.5-2  cm.  long;  petals  purplish; 
stamens  2-3  cm.  long,  reddish. 

Type  locality,  "Commerson  a  observe  cet  arbris.seau  sur  les  montagnes  du 
Magellan,  dans  presque  tous  les  bois,  en  Janvier  17G.S."  The  range  of  this 
variety  is  in  Chile  mostly  south  of  Coneepci«'>n,  and  in  Argentina.  Hepresenta- 
tive  material,  "F.  mngrUauim  Lam.  diet.  Dorvalli  Commerson  herb.,"  Lam- 
ank  Herb.  (P)  ;  F.  jvulfiflorn.  eiilt.  Iiortul.  (Jirod.  CuMieva.  in  1837  (.\Y)  ;  /•'. 


10  CALIFOBXIA  ACADEMY  OF  SCIENCES  [Proc.  4th  Seb. 

discolor,  II.  B.  Glasgow  in  1850  (NY)  ;  F.  tenella,  cult.  Lochowitz  in  1845  (F) . 
CHILE  :  Coneepeion :  San  Vicente,  Pennell  12866  (GH).  Araiico:  Isl.  of  St. 
Mary's,  Eujliis  (US).  Malleeo:  Angol,  Kuntze  in  1892  (F).  Yaldivia  :  Corral, 
Gunckel2  (F)  ;Ceri'ode\a'Mavum,Gunckel5064  (POM)  ;  Valdivia,  B«c/ihew 
in  1896  (Gil)  ;  Pangnipulli,  Hollermaj'er  under  Werdermeyer  1364  (CAS,  F, 
GH,  NY,  US).  Llanquihue:  Ancud,  Chiloe,  Pennell  12492  (F,  GH)  ;  Isla 
Chiloe,  near  Castro,  Mexia  8009  (F  ,GH,  NY,  US)  ;  Isla  Chiloe,  Ruiz  (POM) ; 
Peulla,  Pennell  12675  (F,  GH,  NY).  ARGENTINA :  Neuquen  :  Puerto  Meyer, 
Cabrera  and  Job  210  (NY).  Rio  Negro:  Lago  Nahuelhuapi,  Ljimgner  226 
(NY).  Patagonia:  Port  Otway,  Lee  in  1888  (US)  ;  Hassler  Glacier,  Hill  in 
1872  (GH) ;  Eden  Harbor,  Blake  in  1872  (GH). 

Intergrades  with  var.  macrostema  are  many,  for  example,  with  large  leaves : 
Valparaiso,  Wilkes  Exped.  (GH,  NY,  US)  ;  road  from  Curico,  Chile  to  Argen- 
tina, beyond  Los  Queues,  Mexia  7863  (F,  GH,  NY,  US)  ;  Hualqui,  Coneep- 
eion, Kuntze  in  1892  (NY);  Aysen,  Llanquihue,  Moreira  in  1934  (POM)  ; 
Isla  Victoria,  Lago  N.  Huapi,  Argentina,  Burkart  6397  (F)  ;  Lake  Argentina, 
Santa  Cruz,  Furlong  98  (GH,  NY,  US).  Other  intergrades  in  length  of  pedi- 
cels, etc.,  are :  Casa  Panque,  Llanquihue,  Chile,  Shannon  25  (US)  ;  Camino  de 
Bariloche  al  Correntoso,  Argentina,  Cahrera  5044  (F,  GH,  NY). 

(3b)  Fuchsia  magellanica  Lam.  var.  macrostema  (E.  and  P.)  Munz,  new  comb. 

F.  macrostema  R.  and  P.,  Fl.  Peniv.  3  :88,  pi.  324,  fig.  b,  1802. 

F.  gracilis  var.  m-acrostema  Lindl.,  Bot.  Reg.,  13 :  under  pi.  1052,  1827. 

F.  coccinea  var.  macro.itema  Hook.,  Fl.  Ant.,  2  :269,  1847. 

F.  coccinea  Curtis,  Bot.  Mag.,  1  :pl.  97,  1789,  not  Ait. 

F.  decussata  R.  Grah.,  Edinb.  Phil.  Journ.,  11:206  and  401,  1824;  Sims  in  Bot.  Mag.,  51: 

pi.  2507,  1824,  not  R.  and  P.,  1802. 
F.  gracilis  Lindl.,  Bot.  Reg.,  10: pi.  847,  1824,  new  name  for  Graham  F.  macrostemma  var. 

gracilis  Sweet,  Brit.  Fl.  Gard.,  6:  under  pi.  216,  1833  plant  (decussata)  ■  F.  magellanica 

var.  gracilis  Bailey,  Cyclop.  Am.  Hort.,  614,  1900. 
F.  macrostema  var.  recurvata  Hook.,  Bot.  Mag.,  63:3521,  1836;  F.  recurvata  Niven  ex 

Hook.,  I.e. 
Thilcum  tinctorium  Molina,  Sagg.  Chile,  ed.  2, 146,  1810. 

Leaf -blades  mostly  25-50  mm.  long,  10-20  mm.  wide,  membranaceous; 
pedicels  mostly  4-5.5  cm.  long,  red ;  hypanthium  and  sepals  red  ;  sepals  19-23 
mm.  long;  petals  purplish;  stamens  30-35  mm.  long,  reddish. 

Type  locality,  "Habitat  in  Conccpcionis  Chile  locis  uliginosis,  praesertim  in 
Quebrada  de  Caranio."  Ranging  mostly  in  Chile  from  Aconcagua  to  Coneep- 
eion. Representative  material,  without  locality  ,F.  macrostemma,  Gay  77,  Herb. 
Mus.  Paris  (F).  CPIILE  :  Aconcagua  :  15  km.  south  of  Valparaiso,  ^Yest  3980 
(GH).  Santiago:  Cordillera  de  Pefialolen,  Looser  2299  (POM);  Santiago, 
Bro.  Claude-Joseph  531  (NY,  US).  O'Higgins:  ]\Ionte  la  Leona,  Rancagua, 
Bertero  280  (GH,  NY).  Curico:  Cordillera  Peteroa-Planchoa,  Canon  Rio 
Claro,  Mexia  04406  (GH).  Nuble:  Banos  de  Chilian,  Pennell  12473  (F,  GH, 
US).  Coneepeion :  Talcahuano,  C.  and  I.  Skottsberg  1454  (NY)  ;  presumably 
from  Quebrada  de  Carcamo,  Ruiz  and  Pavon,  as  macrostema,  Madrid  Herb. 


I 


Vol.  XXVJ  MUNZ:  THE  GENUS  FUCHSIA  11 

(photo  F,  POM).  Bio  Bio:  Pailalmeqne,  Padre  Pivian  201  (GH).  Cautin : 
Hotel  Tolhiiaca  on  road  to  Banos  Calientes,  Morrison  and  Fagenknecht  17499 
(GH).  Magallanes :  Port  Famine,  Lee  in  1888  (F,  US)  ;  Eberhard,  Donat  in 
1931  (CAS,  GH,  NY).  ARGENTINA:  Terr,  del  Neuquen :  Puerto  Blest, 
Bruch  in  1898  (NY) . 

Intergrades  have  been  discussed  under  var.  typica. 

(3c)  Fuchsia  magellanica  Lam.  var.  Molinae  Espinosa 
Fuchsia  magellanica  Lam.  var.  Molinae  Espinosa,  Bol.  Mus.  Nac.  Chile,  12:102,  1929. 

Pedicels  and  ovaries  greenish ;  hypanthium,  sepals  and  stamens  and  style 
pink ;  corolla  lilac-pink. 

Type  locality,  CHILE :  Prov.  Llanquihue :  Isla  Chiloe,  near  Castro,  Que- 
brada  de  La  Chacra,  type  coll.  by  Espinosa,  Jan.  31,  1929  (SGO).  Another 
collection  seen  from  Isla  Chiloe  :  F.  Flaminio  Ruiz  (POM) .  Valdivia  :  Corral, 
West  4846  (GH),  Gnnckel  2133  (GH).  Cultivated  at  Valparaiso,  Ooodspeed 
4589  (GH) .  There  is  some  question  as  to  whether  this  is  a  natural  entity  or  a 
cultivated  one,  but  it  seems  to  be  the  former. 

(4)  Fuchsia  coccinea  Soland. 
(Plate  1,  fig.  4) 

Fuchsia  coccinea  Soland.  in  Aitoii,  Hort.  Kew.,  ed.  1,  2:8,  1789. 

F.  coccinea  J.  D.  Hooker,  Bot.  Mag.,  94: pi.  5740, 1868. 

Nahusia  coccinea  Schneevogt,  Icones,  21,  1793. 

STcinnera  coccinea  Moench,  Meth.  Suppl.,  270,  1802. 

Fuchsia  elegans  Salisbury,  Ic.  Stirp.  Ear.,  13,  pi.  7, 1791 ;  said  to  come  from  Brazil. 

F.  pendula  Salisbury,  Prodr.,  279,  1796;  cites  plate  and  page  in  Stirp.  for  elegans. 

F.  pubescens  Cambess.,  in  St.  Hil.,  Fl.  Bras.  Merid.,  2:275,  pi.  134,  1829. 

F.  montana  Cambess.,  I.e.,  pi.  135. 

F.  Glazioviana  Taub.,  Bot.  Jahrb.  15,  Beibl.,  34:16,  1892. 

Shrub,  bushy  and  up  to  1  m.  tall,  or  semiscandent  and  3  or  more  m.  tall ; 
freely  branched,  the  younger  twigs  very  slender,  scarcely  1  mm.  thick,  sub- 
terete,  densely  puberulent,  sometimes  also  pilose,  more  or  less  reddish ;  leaves 
opposite  or  in  3's,  membranaceous,  narrow-ovate,  paler  beneath,  puberulent 
above  and  beneath,  often  also  pilose  beneath  along  main  veins,  rounded  to 
cordate  at  base,  subacuminate  at  apex,  usually  plainly  serrulate,  the  blades 
with  4r-6  main  veins  on  each  side  of  midrib,  15-45  (60)  mm.  long,  10-20  (25) 
mm.  wide ;  petioles  mostly  1-3  mm.  long,  frequently  pilose ;  stipules  lance- 
deltoid,  less  than  1  mm.  long,  often  reflexed,  caducous;  flowers  solitary  in 
upper  axils ;  pedicels  filiform,  puberulent,  sometimes  scattered-pilose,  2-A  cm. 
long;  ovary  oblong;  hypanthium  red,  subfusiform- tubular,  glandular-puber- 
ulent  without,  sometimes  with  few  long  hairs,  pubescent  within,  4—6  mm.  long, 
ca.  3  mm.  wide ;  sepals  red,  glandular-puberulent,  oblong-linear  to  -lanceolate, 
acuminate,  15-20  mm.  long,  connate  at  base  for  3-6  mm. ;  petals  violet  to 
purplish,  obovate,  6-9  mm.  long ;  episepalous  stamens  15-30  mm.  long ;  epi- 
petalous  2-5  mm.  shorter ;  anthers  reddish,  ca.  2  mm.  long ;  style  pubescent  at 


12  CALIFORNIA  ACADEMY  OF  SCIENCES  [Proc.  4th  Ser. 

base,  2-4  cm.  long;  stigma  subentire,  reddish,  1.5  mm.  long;  berry  mostly 
oblong,  about  1  cm.  long. 

Type  locality :  Hooker  (Bot.  Mag.  pi.  5740)  says :  "The  native  country  of  the 
true  coccinea  is  unknown  ;  it  is  probably  Brazilian,  as  it  resembles  the  Brazilian 
more  than  the  Western  or  Southern  American  species ;  Salisbury  says  it  was 
introduced  by  Vandeli  from  Brazil,  whereas  Alton  attributes  its  introduction 
to  Captain  Frith,  from  Chile. 

"The  evidence  of  the  plant  here  figured  being  the  true  and  original  plant  of 
Alton's  'Hortus  Kewensis,'  ed.  1,  rests  on  the  fact  that  named  specimens  of  the 
same  are  preserved  in  the  Banksian  Herbarium  of  the  British  Museum,  and 
in  Sir  J.  Smith's  Herbarium  at  the  Linnean  Society,  all  procured  at  Kew  in 
the  year  of  the  introduction  of  the  plant,  and  at  the  date  of  its  being  described 
by  Alton."  . 

It  takes  some  courage  to  apply  the  name  coccinea  to  the  Brazilian  plant 
commonly  passing  as  puhescens  or  montana,  but  I  see  no  alternative.  These 
are  the  only  Fuchsias  which  I  can  find  that  agree  with  all  that  Hooker  says 
above  and  with  specimens  seen.  ]\Iaterial  seen,  WITHOUT  LOCALITY :  F. 
coccinea,  Herl).  Steud.  (NY),  Meisner  Herh.  (NY),  Hort.  Kew.  M.  Ait.  (G). 
BRAZIL :  So.  Brazil,  Glaziou  17614,  F.  Olazioviana  (fragment  from  Berlin 
and  photograph  at  F,  photograph  at  POM)  ;  without  locality,  F.  puhescens 
Camb.,  St.  Hil.  C  1545,  fragment  from  Mus.  Paris  (F).  Minas  Geraes:  F. 
montana  Camb.,  St.  Hilaire,  fragment  from  Mus.  Paris,  must  be  from  Serra 
da  Caraca.  (F)  ;  Serra  da  Piedade,  Municipio  de  Caete,  Barreto  7160  (F) ,  8803 
(F),  7162  (F),  7158  (F);  Serra  da  Piedade,  Hoehne  6278  (POM),  6281 
(POM).  There  is  every  intergradation  from  being  merely  puberulent  as  in 
the  St.  Hilaire  specimen  of  montana,  through  having  some  longer  hairs  at  the 
base  of  the  midrib,  as  in  St.  Hilaire  collection  of  puhescens,  Hoehne  6278, 
6281,  Barreto  7160,  7162,  8803,  to  being  pilose  on  the  stems :  Barreto  7158, 
garden  specimens  from  Herb.  Steudel,  etc.  The  species  has  been  naturalized 
in  some  places  as  in  Jamaica :  Cinchona,  Nichols  60  (F,  GH) ,  Harris  9130  (F) 
and  Greenhill,  Harris  in  1899  (F). 

(5)  Fuchsia  hybrida  Hort. 

Fuchsia  hyhrida  Hort.  ex.  Vilmorin,  Blumeng.,  ed.  3,  Sieb.  and  Voss,  1 :332,  1894;  Behrens, 

Blattenformen,  pi.  71,  fig.  d,  e,  f,  1900. 
F.  speciosa  Hort.  ex  Bailey,  Cyclop.  Amer.  Hort.,  614-615, 1900. 

Sturdy,  with  twigs  usually  reddish;  leaves  opposite,  broadly  ovate  to 
oblong-ovate,  rounded  to  cordate  at  base,  apically  acute,  strongly  serrulate, 
3-5  cm.  long,  1.5-3  em.  wide ;  petioles  mostly  1-2  cm.  long;  pedicels  commonly 
4r-5  cm.  long;  ovary  ovoid,  6-8  mm.  long,  4-5  mm.  wide;  hypanthium  1-2  or 
more  cm.  long,  4-6  mm.  wide;  sepals  2.5-3  cm.  long,  8-11  mm.  wdde;  petals 
1-2  cm.  long ;  stamens  included  to  exserted ;  style  4-5  or  more  cm.  long ;  stigma 
3-4  mm.  wide. 

Manv  manv  strains  of  the  garden  fuclisias,  whicli  are  almost  universally 


Vol.  XXV]  MUNZ:  THE  GENUS  FUCHSIA  13 

hybrid,  are  described  more  or  less  accurately  by  the  above  and  are  said  by 
Bailey  (Ilortus,  264.  1930)  to  be  "probably  of  hybrid  derivation  from  forms 
of  F.  ynagellanica  and  F.  fulgens." 

(6)  Fuchsia  regia  (Vand.)  Munz,  new  comb. 
QiLclusia  regia  Vand.  ex  Veil.,  Fl.  Flum.,  149, 1825  and  4:pl.  6, 1827. 

Semiscandent  shrub  up  to  6  m.  tall,  freely  branched,  the  ultimate  branchlets 
slender,  1-2  mm.  thick,  flexuous,  angled,  dark  red,  glabrous  to  puberulent  to 
pilose;  leaves  opposite  or  in  3's,  oblong-ovate,  entire  to  remotely  and  incon- 
spicuously denticulate,  coriaceous,  rounded  at  base,  more  or  less  abruptly 
acuminate  at  apex  (this  usually  not  in  same  plane  as  rest  of  blade  and  some- 
what folded  in  pressed  specimens),  glabrous  to  puberulent  or  pilose,  often 
reddish  on  veins ;  blades  mostly  4-10  cm.  long,  2-4  cm.  wide,  with  5-7  main 
veins  on  each  side  of  midrib ;  petioles  5-10  mm.  long ;  stipules  deltoid,  about 
0.5  mm.  long,  deciduous;  flowers  solitary  in  upper  axils;  pedicels  almost  fili- 
form, 2-4.5  cm.  long,  glabrous  to  pubescent ;  ovary  oblong;  hypanthium  deep 
red,  cylindric,  slightly  enlarged  toward  apex,  sometimes  somewhat  fusiform, 
3-15  mm.  long,  3-5  mm.  wide,  glabrous  to  pilose  without  and  within ;  calyx 
red,  glabrous  to  somewhat  pilose,  13-35  mm.  long,  4-lobed,  the  segments  con- 
nate for  2-15  mm.,  the  free  segments  lanceolate,  spreading,  10-25  (30)  mm. 
long,  acuminate;  petals  purplish,  obovate,  obtuse,  10-15  (18)  mm.  long;  epi- 
sepalous  stamens  20-35  (40)  mm.  long,  epipetalous  16-30  (35)  mm.  long; 
anthers  red,  2.5-3.5  mm.  long;  style  somewhat  pubescent  at  base,  3.5-5.5  (7.5) 
em.  long;  stigma  yellowish,  or  reddish,  capitate,  subentire,  2.5-3.5  mm.  long; 
berry  oblong  or  more  spherical,  1.5-2.5  cm.  long. 

The  name  regia  is  based  on  a  plate,  which  shows  large  leaves  and  flowers 
and  a  fusiform  hypanthium,  sepals  connate  a  short  distance.  This  combination 
of  characters  and  the  locality  can  mean  only  one  thing  :  the  species  which  has 
been  called  F.  integrifolia.  It  is  fair  to  admit  that  the  plate  shows  the  leaves 
more  dentate  than  one  would  like. 

Key  to  Varieties  of  Fuchsia  regia 
A.Young  twigs  shaggy-pilose;  hypanthium  3-6  mm.  long;  calyx  (united  plus  free  por- 
tions) 13-21  mm.  long 6d.  var.  alpestris. 

AA.  Young  twigs  not  pilose;  hypanthium  7-15  mm.  long;  calyx  20-35  mm.  long. 

B.  Young  twigs  densely  and  closely  puberulent ;  connate  portion  of  calyx  9-15  mm. 

long 6c.  var.  affinis. 

BB.  Young  tmgs  essentially  glabrous. 

C.  Connate  base  of  calyx  2-5  mm.  long;  free  segments  18-30  mm.  long. 

6a.  var.  typica. 
CC.  Connate  base  of  calyx  9-14  mm.  long  ;  free  segments  12-20  mm.  long. 

6b.  var.  radieans. 

(6a)  Fuchsia  regia  (Vand.)  Munz  var.  typica  Munz,  new  name 

(Platel,  fig.  5) 
Quelusia  regia  Vajstd.  ex  Veil.,  I.e. 

Fuchsia  iniegrifolia  Cambess.,  in  St.  Hilaire,  Fl.  Bras.  Merid.,  2  :273,  1829;  Bot.  Mag.,  68: 

pl.  3948,  1842;  MiCHELi  in  Martins,  Fl,  Bras.  13,  pt.  2:174,  pi.  37,  1875. 


14  CALIFORNIA  ACADEMY  OF  SCIENCES  [Proc.  4th  Ser. 

Glabrous  except  sometimes  slightly  and  minutely  puberulent  about  the 
young  gro^v1:h ;  pedicels  3-4.5  cm.  long;  hj^panthium  7-15  mm.  long;  calyx 
24r-35  mm.  long,  connate  at  base  for  2-5  mm.,  free  portion  18-30  mm.  long. 

Tj-pe  locality,  "ad  Alpes  Pharmacopolitanas  fruticetis,  per  quae  ad  oppi- 
dum  Cunha  iter  fit,"  hence  near  tlie  western  end  of  the  state  of  Rio  de  Janeiro, 
Brazil.  Ranging  through  the  mountains  of  the  states  of  Rio  and  Minas  Geraes. 
Material  seen,  BRAZIL  :  without  locality,  Burchell2196  (GB.) ,  Riedelin  1816 
(GH),  Gardner  375  (NY).  Minas  Geraes :  Ouro  Prieto,  Cochran  29191  (US)  ; 
Districto  Carangola,  trail  to  Areponga,  Mexia  4326  (F,  CAS,  GH,  NY,  US)  ; 
top  of  Serra  da  Gramma,  Mexia  4278  (F,  GH,  NY,  US) ;  Caldas,  Claiissen  in 
1840  (GH),  Regnell  in  1878  (NY,  US)  ;  Serra  do  Cipo,  Barreto  9254  (F)  ; 
Serra  do  Caraga,  Barreto  7157  (F).  Rio  de  Janeiro:  Maceirras,  Mt.  Itatiaya, 
L.  B.  Smith  1476  (GH)  ;  near  Rio  de  Janeiro,  Wilkes  Exped.  in  1838-42  (NY) ; 
Organ  Mts.,  Wilkes  Exped.  (US). 

Since  this  is  the  most  common  form  of  the  species  and  is  in  the  Organ  Mts. 
and  since  the  original  plate  shows  sepals  slightly  connate  and  does  not  show 
hair,  I  have  to  assume  that  this  is  typical  regia. 

(6b)  Fuchsia  regia  (Vand.)  Munz  var.  radicans  (Miers)  Munz,  new  comb. 

F.  radicans  Miers  in  Bot.  Reg.,  27:  Misc.  no.  167,  and  pi.  66, 1841. 
F.  pyri folia  Presl,  Symbol,  bot.,  2  :19,  pi.  65,  1858. 

Glabrous  throughout;  pedicels  2-3  cm.  long;  hypanthium  9-11  mm.  long; 
calyx  2^34  mm.  long,  connate  at  base  for  9-14  mm.  then  free  for  12-20  mm. 

Type  locality,  Organ  Mts.,  Rio  de  Janeiro,  Brazil.  Material  seen,  BRAZIL  : 
Rio  de  Janeiro :  Organ  Mts.,  3Iiers  4453,  Feb.,  1838  (US)  ;  Serra  do  Itatiaia  : 
Retiro,  Dusen  81  (GH,  US).  Parana :  Serra  do  Mar,  Desvio  Ypiranga,  Dusen 
6766  (GH). 

(6c)  Fuchsia  regia  (Vand.)  Munz  var.  affinis  (Camb.)  Munz,  new  comb. 
F.  afflnis  Cambess.,  in  St.  Hilaire,  Fl.  Bras.  Merid.,  2  :274,  1829. 

Young  twigs  densely  and  closely  puberulent  (leaves  maj-  be  somewhat 
pilose  on  under  surface  along  veins);  pedicels  2-4  cm.  long;  hypanthium 
8-10  (13)  mm.  long;  calyx  20-30  mm.  long,  connate  at  base  for  9-15  mm., 
then  free  for  10-20  mm. 

Type  locality,  Serra  da  Boa  Vista,  Rio  de  Janeiro,  Brazil.  Material  seen 
from  the  states  of  Sao  Paulo  and  Parana.  BRAZIL :  without  locality,  St. 
Hilaire  22,  ex  Herb.  Mus.  Paris  (F).  Sao  Paulo :  Alto  da  Serra,  at  about  900 
m.,  L.  B.  Smith  1900  (F,  G}l),Mimz  15415  (F,  NY,  POM,  US)  ;  Est.  Campo 
Grande,  Edicall  1957  (POM).  Parana:  Curitiba,  Hoehne  in  1928  (POIM), 
Dusen  14362  (F,  GH,  NY).  A  collection  from  Horto  Botanico  Cantareira, 
Puttemans  204  (P)  has  the  puberulence  of  affinis  and  the  flower  of  typica. 

(6d)  Fuchsia  regia  (Vand.)  Munz  var.  alpestris  (Gardner)  IMunz,  new  comb. 

F.  alpestris  Gardner,  Bot.  Mag.,  69:pl.  3999,  1843. 
F.  mollis  Krause,  Engl.  Jahrb.,  37:600,  1906. 


Vol.  XXV]  MUNZ:  THE  GENUS  FUCHSIA  15 

Young  twigs  densely  shaggy-pilose ;  pedicels  2.5-4.5  cm.  long;  hypanthium 
3-6  mm.  long,  glandular-pubescent  and  more  or  less  pilose ;  calyx  13-21  mm. 
long,  connate  at  base  for  2-6  mm.,  then  free  for  10-19  mm. 

T.ype  locality.  Organ  Mts.,  Rio  de  Janeiro,  Brazil.  Material  seen ;  without 
locality,  H.  B.  Glasgov,  F.  alpestris,  Sept.,  1850  (NY).  BRAZIL:  Minas 
Geraes :  Serra  do  Itacolomy,  Municipio  de  Ouro,  Barreto  9168  (F) ;  Itacolomi, 
Ouro  Preto,  de  Vasconcellos  in  1901  (POM).  Rio  de  Janeiro:  bet.  Petropolis 
and  Theresopolis,  Princess  Therese  von  Bayer^i  in  Herb.  Monacense  (photos 
F,  GH,  POM)  ;  Organ  Mts.,  Gardner  5706,  type  coll.  aliwstris  (NY),  Nova 
Friburgo,  Ule  4418,  labelled  F.  integrifolia  var.  mollis,  a  name  seeming  not 
to  have  been  published,  specimen  at  Rio  and  Berlin  (photo  F,  POM) .  Parana  : 
Porto  de  Cima,  Dusen  14133  (US).  This  entity  approaches  F.  coccinea  in 
becoming  dentate,  but  the  pilosity  is  more  dense,  the  leaves  usually  larger. 
Barreto  9168  is  quite  like  rcgia  typica,  but  more  toothed. 

Section  2.  EUFUCHSIA  Baillon. 

Eiifuchsia  Baill.,  Hist.  PI.,  6  :467,  1877;  Eaimann  in  Engler  and  Prantl,  Die  nat.  Pflanzen- 

f am. ,111,  7:219,  1893. 
Fuchsia,  as  genus,  L.,  Gen.  PL,  ed.  5,  126,  1754;  Spach,  Nouv.  Ann.  Sci.  Nat.,  (2),  4:177, 

1835 ;  as  section,  Endlicher,  Gen.  PI.,  1193,  1840. 
Ellobium  Lilja,  Linnaea,  15 : 262,  1841. 

Flowers  perfect ;  petals  convolute.  Stamens  erect,  somewhat  exserted  or  not, 
those  opposite  the  petals  shorter.  Fruit  many-seeded.  Leaves  opposite  or 
whorled. 

Type  species,  F.  triphylla  L.  .     • 

Key  to  Species  of  Section  EUFUCHSIA 

A.  Flowers  axillary,  that  is  borne  in  axils  of  ordinary  or  somewhat  reduced  foliage  leaves. 
B.  Hypanthium  3-30  mm.  long. 

C.  Hypanthium  3-5  mm.  long,  shorter  than  the  ovary;  young  twigs  subglabrous. 

Colombia 51.  F.  verrucosa. 

CC.  Hypanthium  12-30  mm.  long,  longer  than  ovary. 

D.  Younger  twigs,  pedicels,  etc.,  essentially  glabrous  (sometimes  somewhat 
pilose  about  nodes)  ;  leaves  mostly  glabrous  except  sometimes  along  veins 
of  under  surface. 
E.  Petioles  5-20  mm.  long;  sepals  scarlet.  S.  Peru  to  Bolivia. 

52.  F.  sanctae-rosae. 
EE.  Petioles  20-35  mm.  long;  sepals  whitish,  Ecuador. .  .  17.  F.  pallescens. 
DD.  Younger  twigs,  pedicels,  etc.,  not  glabrous. 

E.  Hairs  on  twigs  and  pedicels  spreading ;  leaf -blades  mostly  5-12  cm. 
long. 
F.  Stamens  conspicuously  exserted ;  petals  green ;  leaves  with  7-9 
principal  lateral  veins  on  each  side  of  midrib.  Mexico  to  Costa 

Eica 7.  F.  splendens. 

FF.  Stamens  not  exserted;  petals  red ;  lateral  veins  10-15. 
G.  Leaves  opposite,  elliptic  to  obovate.  Ecuador. 

50.  F.  scabriuscula. 
GG.  Leaves  mostly  ternate,  lanceolate  to  lanceovate.  West  Indes. 

23.  F.  triphylla. 


I 
16  CALIFORNIA  ACADEMY  OF  SCIENCES  [Proc.  4th  Ser. 

EE.  Ilairs  on  twigs  more  or  less  appressed  (if  somewhat  spreading,  then 
leaf -blades  usually  less  than  5  cm.  long) ;  principal  lateral  veins 
mostly  3-10  on  each  side  of  midrib. 
F.  Twigs  pubescent;  petioles  mostly  less  than  1  cm.  long:  liypan- 
thium  mostly  less  than  2  cm.  long. 
G.  Sepals  7-12  mm.  long.  South  America. 

H.  Pubescence  brownish ;  leaf -blades  usually  not  more 
than  3  times  as  long  as  wide,  pale  green  beneath,  ser- 
rate. Peru 48.  F.  decussata. 

HH.  Pubescence  grayish;   leaf -blades  usually  4  times  as 
'  long  as  wide,  whitish  beneath,  subentire  to  remotely 

serrulate.  Ecuador 49.  F.  hypoleuca. 

GG.  Sepals  18-22  mm.  long.  West  Indes 22.  F.  Fringsheimii. 

FF.  Twigs  minutely  puberulent ;  petioles  1-3  cm.  long ;  hypanthium 

2-3  cm.  long.  Ecuador - 16.  F.  loxensis. 

BB.  Hypanthium  mostly  30-70  mm.  long. 

C.  Petals  narrow,  2-3  times  as  long  as  wide. 

D.  Young  stems  essentially  glabrous  except  for  the  puberulenee  near  the 

tips ;  style  glabrous  throughout ;  petals  mostly  glabrous. 

E.  Hypanthium   35-45   mm.   long;    sepals   long-acuminate.   Peru   and 

Bolivia. 

F.  The  hypanthium  5-7  mm.  wide  at  base,  usually  villous  within,  not 

much  narrowed  above ;  sepals  with  green  tips ;  principal  lateral 

veins  12-15  on  each  side  of  midrib 10.  F.  deniiculata. 

FF.  The  hypanthium  2-3  mm.  thick  at  base,  then  narrowed;  sepals 
apparently  red  throughout. 
G.  Principal  lateral  veins  7-10  on  each  side  of  midrib ;  hypan- 
thium glabrous  within;   sepals  17-20  mm.  long.  Junin  to 

to  Cuzco,  Peru 11.  -F.  leptopoda. 

GG.  Principal  lateral  veins  13-15  on  each  side  of  midrib  ;  hypan- 
thium villous  within ;   sepals  12-14  mm.  long.  Amazonas, 

Peru 12.  F.  TToijIl-oicskii. 

EE.  Hypanthium  50-60  mm.  long ;  sepals  acute.  Colombia. 

13.  F.  magdalenae. 
DD.  Young  stems  not  glabrous ;  style  more  or  less  pilose  on  lower  portion. 

E.  Hypanthium  7-11  mm.  wide  at  summit;  petals  frequently  with  some 
hair  on  backs ;  stems  puberulent  or  villous ;  fruit  8-10  mm.  thick. 
F.  The  young  stems  with  fine  ashy  appressed  puberulenee ;  pedicels 
15-25   mm.   long;    fruit   8-10   mm.   long,   subglobose.   Central 

Cordillera  of  Colomliia 24.  F.  petiolaris. 

FF.  The  young  stems  ferruginous-villous;  pedicels  25-40  (60)  mm. 
long;   fruit  12-15  mm.  long,  ellipsoid.  Eastern  Cordillera  of 

Colombia 25.  F.  Smithii. 

EE.  Hypanthium  5  mm.  wide  at  suiinnit ;  petals  glabrous;  fruit  less  than 
8  mm.  thick.  Peru. 
F.  Stems,  etc.,  crisp-pubescent ;  petioles  10-15  mm.  long ;  petals 

obtusish.  Dept.  Piura 25.  F.  AspJundii. 

FF.  Stems,  etc.,  minutely  appressed-puberulent;   petioles  4-8  mm. 
long;  petals  subacuniinate.  Dept.  Amazonas..  .27.  F.  Llewelynii. 
CC.  Petals  wider,  less  than  twice  as  long  as  wide. 

D.Young   stems   mostly   glabrous,   except   sometimes  (toward   tips;    style 
glabrous  or  nearly  so  to  its  base. 
E.  Petals  red.  So.  America. 


Vol.  XXV]  MUNZ :  THE  GEXUS  FUCHSIA  '  17 

F.  Hypanthium  25-40  mm.  long;  leaf-blades  mostly  2-10  cm.  long, 
1-3  cm.  wide;  petioles  2-10  (15)  mm.  long.  Peru. 

9.  F.  austromontana. 

FF.  Hypanthium  60-70  mm.  long;  leaf -blades  8-18  cm.  long,  4-7.  (9) 

cm.  wide;  petioles  10-30  mm.  long.  Ecuador.  20.  F.  macrostigma. 

EE.  Petals  green.  Guatemala 8.  F.  cordi folia. 

DD.  Youug  stems  hirtellous  to  pilose ;  style  pilose  or  villous  near  base. 

E.  Twigs  hirtellous  with  short  subappressed  hairs;  petioles  1-5  (8)  mm. 
long  ;  pedicels  1-2  cm.  long. 
F.  Leaf -blades  2-4  cm.  long,  1-2  cm.  wide ;  petioles  3-8  mm.  long. 

Colombia  and  Ecuador 14.  F.  coneacens. 

FF.  Leaf -blades  5-6.5  cm.  long,  2.5-3  cm.  wide;  petioles  quite  lack- 
ing. No.  Peru 15.  F.  rivularis. 

EE.  Twigs  pubescent  to  pilose,  with  spreading  hairs;  petioles  5-20  mm. 
long;  pedicels  15-45  mm.  long.  Ecuador  and  Peru. 
F.  Hypanthium  narrow,  2  mm.  thick  at  base,  5-6  mm.  wide  at 
summit;  sepals  10-12  mm.  long,  somewhat  divergent.  Ecuador. 

18.  F.  Townsendii. 

FF.  Hypanthium  -wider,  3-5  mm.  thick  at  base,  7-11  mm.  wide  at 

summit ;  sepals  15-22  mm.  long,  spreading-reflexed. 

G.  Hypanthium  gradually  ampliate,  hence  widest  at  its  mouth ; 

sepals  3-4  mm.  wide;  petals  longer  than  wide.  Dept.  Apuri- 

mac,  Peru 19.  F.  plafi/petola. 

GG.  Hypanthium  abruptly  ampliate  near  middle  and  slightly 
narrowed  at  summit ;  sepals  5-6  mm.  wide ;  petals  scarcely 
longer  than  wide.  Ecuador  to  Ayabaca,  Peru. 

21.  F.  ayavacensis. 
AA.  Flowers  in  clusters,  making  a  definite  inflorescence  with  bracts  or  leaves  much  reduced 
and  different  from  foliage  leaves. 
B.  Hypanthium  1-3  cm.  long. 

C.  Young  twigs  essentially  glabrous  except  for  youngest  tips. 

D.  Leaf -blades  generally  at  least  3  times  as  long  as  wide;  hypanthium  13-16 
mm.  long;  petioles  2-4  mm.  long.  Colombia  and  N.  Ecuador. 

63.  F.  sessilifolia. 
DD.  Leaf -blades  usually  2-3  times  as  long  as  wide;  hypanthium  usually  16-30 
mm.  long. 
E.  Leaf -blades  mostly  5-8  cm.  long,  1.5-3.5  cm.  wide ;  style  glabrous. 
'  F.  Flowers  in  elongate  raceme-like  inflorescences,  with  more  or  less 

reduced  foliage-leaves.  S.  Peru  and  Bolivia.  52.  F.  sanctae-rosae. 
FF.  Flowers  in  definite  very  short  and  compact  racemes.  North  of 
So.  Peru, 
G.  Hypanthium  24-30  mm.  long;  sepals  9-11  mm.  long;  prin- 
cipal lateral  veins  on  each  side  of  midrib  8-12. 
H.  Sepals  oblong-ovate,  5-6  mm.  wide;  berry  over  1  cm. 
long ;  pedicels  15-25  mm.  long.  Peru. .  .  53.  F.  Osgoodii. 
HH.  Sepals  linear-lanceolate,  2.5-3  mm.  wide;  berry  5-6 
mm.  long ;  pedicels  8-18  mm.  long.  Ecuador. 

55.  F.  Lehmannii. 
GG.  Hypanthium  14-18  mm.  long;  sepals  7  mm.  long;  principal 
lateral  veins  about  15.  Putumayo,  Colombia. 

56.  F.  pniumayensis. 
EE.  Leaf-blades  mostly  8-20  cm.  long,  3-9  cm.  wide,  with  12-18  principal 
lateral  veins  on  each  side  of  midrib;  style  pubescent  near  base. 


18  CALIFOEXIA  ACADEMY  OF  SCIENCES  [Proc.4thSer. 

F.  Petioles  2-8  mm.  long;  pedicels  5-7  mm.  long;  petals  almost  as 

wide  as  long;  leaves  quite  glabrous.  Ecuador.  61.  F.  glaberrima. 

FF.  Petioles  10^0  mm.  long ;  pedicels  mostly  10-20  mm.  long ;  petals 

about  3  times  as  long  as  wide;  leaves  puberulent  along  veins 

underneath. 

G.  Leaf -blades  mostly  obtuse  at  base;  fruit  ca.  6  mm.  long. 

Colombia 54.  F.  Andrei. 

GG.  Leaf -blades  subacuminate  at  base;   fruit  8-10  mm.  long. 

Peru 62.  F.  macrophylla. 

CC.  Young  twigs  definitely  not  glabrous. 

D.  Leaf -blades  mostly  5-8  cm.  wide ;  pedicels  10-30  mm.  long. 

E.  Hypanthium  13-15  mm.  long ;  racemes  mostly  lateral ;  pedicels  10-15 

mm.  long.  Hiiauuco,  Peru 31.  F.  ovalis. 

EE.  Hypanthium  20-30  mm.  long;  racemes  terminal;  pedicels  15-30  mm. 

mm.  long.  Dept.  Cuzco,  Peru M.F.  tincta. 

DD.  Leaf -blades  mostly  1.5-5  cm',  wide;  inflorescence  often  terminal. 

E.  Stems  with  fine  close  appressed  puberulence ;  hypanthium  very  nar- 
row, scarcely  bulbous  at  base,  14r-19  mm.  long,  2.5-4  mm.  wide  at 

mouth.  Colombia  and  Venezuela  to  Ecuador 64.  i^.  sylvatica. 

EE.  Stems  pubescent  to  pilose ;  hypanthium  somewhat  bulbous  at  base. 

F.  Hairs  reddish 37.  2?".  StorUi. 

FF.  Hairs  not  reddish, 

G.  Flowers  in  terminal  pyramidal  panicles;  hypanthium  14-20 

mm.  long.  Colomlna 57.  F.  Harticegii. 

GG.  Flowers  in  racemes. 

H.  Twigs  pubescent ;  hypanthium  15-20  mm.  long.  Santo 

Domingo  and  Haiti 23.  F.  triphylla. 

HH.  Twigs  pilose;  hypanthium  20-25  mm.  long.  Peru. 

I.  Bracts  small,  subulate  or  narrow;   petioles  of  a 

given  wliorl  subequal 59.  F.  asperifolia. 

II.  Bracts   more  like   reduced  leaves ;    petioles  of  a 
whorl  very  unequal,  one  much  exceeding  the  otliers. 

60.  F.  pilosa. 
'     BB.  Hypanthium  3-6.5  cm.  long. 

C.  Leaves  small,  crowded,  12-16  mm.  long ;  stems  with  brown  hair.  Peru. 

32.  F.  confertifoUa. 
CC.  Leaves  larger,  30-200  mm.  long. 

D.  Leaf -blades  narrow,  more  than  4  times  as  long  as  wide,  in  whorls  of  4, 

linear-lanceolate.  Huanuco,  Peru 31.  F.  simplicicaulis. 

DD.  Leaf-blades  wider,  2-3  times  as  long  as  wide,  opposite  or  ternate  (some- 
times quaternate  in  F.  hirtella). 
E,  Petals  (under  a  lens)  puberulent  to  sparsely  villous  on  back. 

F.  Sepals  pubescent  on  inner  or  upper  surface;  fruit  oblong;  leaf- 

blades  rigid.  Colombia  and  Venezuela 28.  F.  venusfa. 

FF.  Sepals  glabrous  on  inner  surface ;  fruit  subglobose ;  leaf -blades 
membranaceous.  Venezuela. 
G.  Young   stems   appressed-puberulent ;    pedicels   20-40   mm. 
lolig;  hypanthium  35-40  mm.  long;  sepals  spreading,  18-20 

mm.  long;  petals  strigulose  on  back 29.  F.  Jaltnii. 

GG.  Young  stems  glabrous  or  somewhat  villous ;  pedicels  15-20 
mm.  long;  hypanthium  40-50  mm.  long;  sepals  divergent, 
14-15  mm.  long;  petals  sparingly  villous  on  back. 

30.  F.  Gehriperi. 


Vol.  XXV]  MUNZ :  THE  GENUS  FUCHSIA  19 

EE.  Petals  glabrous  on  back. 

r.  Young  stems  glabrous ;   leaves  subglabrous  above,  puberulent 
beneath ;  petioles  2-8  mm.  long. 
G.  Leaf -blades  3  times  as  long  as  wide ;  principal  lateral  veins 
ca.  20  on  each  side  of  midrib;  pedicels  25-40  mm.  long. 

Peru 43.  F.  abrupta. 

GG.  Leaf -blades  less  than  3  times  as  long  as  wide ;   principal 
lateral  veins  10-12  on  each  side  of  midrib;  pedicels  7-13 

mm.  long.  Colombia 44.  F.  Cuatrecasasii. 

FF.  Young  stems  not  glabrous  under  a  lens. 
G.  The  young  stems  finely  puberulent. 

H.  Principal  lateral  veins  about  12  on  each  side  of  midrib  ; 
berry  5-6  mm.  long;  pedicels  10-40  mm.  long.  Colom- 
bia  45.  F.  Eillipii. 

HH.  Principal  lateral  veins  14-24  on  each  side  of  midrib ; 
berry  6-12  mm.  long;  pedicels  6-15  mm.  long. 

I.  Leaves  quite  glabrous ;  berry  subglobose,  almost 

1  cm.  long.  Junin,  Peru 39.  F.  Munzii. 

II.  Leaves  not  glabrous  under  a  lens ;  berry  longer 
than  thick. 
J.  Leaf-blades  densely  soft-puberulent,  with  14- 
16  principal  lateral  veins  on  each  side  of  mid- 
rib;  style  pubescent  near  base.  Ecuador  to 

Huanuco,  Peru 42.  2^.  corymbiflora. 

JJ.  Leaf -blades  glabrous  on  casual  inspection,  but 
minutely  strigulose  under  a  lens,  with  18-24 
principal  lateral  veins ;  style  almost  glabrous. 

Libertad,  Peru 30.  F.  Aspiazui. 

GG.  The  young  stems  pubescent  to  villous  (very  sparsely  pubes- 
cent in  F.  fidgens). 
H.  Stems  and  leaves  pilose  or  villous  with  brownish  or 
reddish  hairs ;  pedicels  15-40  mm.  long. 

I.  Sepals  15-20  mm.  long ;   hypanthium  35-45  mm. 

long.  Bolivia 38.  2^.  furfuracea. 

II.  Sepals  8-12  mm.  long.  Peru. 

J.  Hypanthium  20-35  mm.  long;  petioles  10-50 

mm.  long.  Cuzco 34.  F.  tincta. 

JJ.  Hypanthium  40-45   mm.   long ;    petioles   2-5 
mm.  long. 
K.  Leaves   ternate,  broadlj^  elliptic.   Ama- 

zonas 35.  F.  Mathewsii. 

.     KK.  Leaves  opposite,  oblong-lanceolate.  Caja- 

marca 36.  i^.  Fischeri. 

HH.  Stems   and   leaves   strigulose   to   pubescent ;    pedicels 
mostly  5-20  mm.  long. 

I.  Sepals  red,  pubescent  on  backs.  Mostly  So.  Ameri- 
can. 
J.  Sepals     somewhat     divergent ;     hypanthium 
strigulose  without. 
K.  Leaves  membranaceous,  the  apex  in  the 
same  plane  as  the  lilade ;  berry  over  1  cm. 
long.  Central  Colomhia..  .40.  F.  hi rfella. 
^  KK.  Leaves  rigid,  rugose,  the  apex  l)ent  to 


20  CALIFORNIA  ACADEMY  OF  SCIENCES  [Proc.  4th  Ser. 

one  side  in  pressing ;  berry  less  than  1  cm. 
long.  So.  Colombia  and  No.  Ecuador. 

41.  F.  polyaniha. 

J  J.  Sepals  spreading-reflexed ;  hypantliium  -with 

spreading   hairs.   West   Indies   and   Central 

America  to  Argentina 46.  F.  boJiviana. 

II.  Sepals  green,  glabrous  except  for  the  sparse  pubes- 
cence near  the  tips,  divergent-spreading.  Mexico. 

47.  F.  fulgens. 

(7)  Fuchsia  splendens  Zuccarini 

(Plate  1,  fig.  6) 

Fuchsia  splendens  Zucc,  Flora,  1832,  ii,  Beibl.,  102,  1832. 

F.  splendens  in  Curtis,  Bot.  Mag.,  70:pl.  4082,  1844;  Bot  Eeg.,  28:pl.  67,  1842;  Hemsley, 

in  Bot.  Biol.  Centr.  Amer.,  1:460,  1880. 
F.  cordifolia  /3  Hooker,  Icon.  PI.,  5:450,  1842. 
F.  intermedia  IIemsley,  Diag.  PI.  Nov.,  1 :  14, 1878 ;  Bot.  of  Biol.  Centr.  Amer.,  1 :457,  1880. 

Based  on  Hartweg  460  which  I  have  not  seen,  but  the  characters  given  do  not  hold  in  a 

series  of  specimens. 

Shrub  6-25  dm.  tall,  often  pendant,  sometimes  a  small  tree,  relatively  few- 
branched,  the  ultimate  branchlets  usually  pilose  or  pubescent,  greenish  to 
reddish;  leaves  opposite,  ovate  to  ovate-cordate,  paler  beneath  than  above, 
membranaceous,  more  or  less  pubescent  especially  when  young,  acuminate, 
denticulate,  the  blades  3-10  cm.  long,  2-6  cm.  wide,  with  or  without  reddish 
tinge,  principal  lateral  veins  7-9  on  each  side  of  midrib  ;  petioles  often  tinged 
red,  pilose  to  subglabrous,  slender,  2-5  cm.  long;  flowers  solitary  in  axils  or 
more  or  less  crowded  on  short  lateral  branches  with  somewhat  reduced  leaves ; 
pedicels  nodding,  subfiliform,  pubescent,  with  some  gland-tipped  hairs,  3-5 
(8)  cm.  long,  not  much  elongate  in  fruit;  hypanthium  17-20  (40)  mm.  long, 
somewhat  compressed,  the  flattened  side  much  broader  than  the  ovary  and 
gradually  ampliate  upward,  6-8  mm.  wide  at  base,  7-12  mm.  wide  at  summit, 
more  narrow  in  other  dimension,  rose  to  bright  red,  externallj'  pubescent  with 
some  hairs  gland-tipped,  internally  pubescent-villous ;  sepals  4,  green  except 
for  the  sometimes  reddish  base,  ovate-lanceolate,  8-15  mm.  long,  5  mm.  wide 
at  base,  pubescent,  the  tips  scarcely  free  or  free  for  1  mm.;  petals  green,  ovate, 
subacuminate,  6-10  mm.  long,  4-5  mm.  wide ;  episepalous  stamens  exceeding 
the  sepals  10-18  mm.  long,  epipetalous  8-16  mm.  long ;  filaments  pale  yellow, 
anthers  yellow,  2-3  mm.  long;  style  glabrous,  exceeding  the  stamens;  stigma 
conoid,  greenish-yellow ;  berry  linear,  3-3.5  cm.  long,  5  or  more  mm.  thick. 

Type  locality,  "in  imperio  mexicano."  Ranging  from  Chiapas  to  Costa  Rica, 
at  altitudes  of  2000  to  10,000  feet.  Variable  in  pubescence,  some  plants  being 
cinereous  (Matuda  2322,  Steyermark  36056,  Pittier  10501,  Standley  and 
Valerio  43829) ,  others  with  longer  pubescence  (Skuteh  118,  Pittier  44,  Steyer- 
mark 32833).  But  there  seems  to  be  no  geographical  or  otlier  correlation  with 
variability  in  pubescence.  There  is  also,  at  least  in  herbarium  material,  con- 
siderable variation  in  relative  length  and  width  of  hypanthium.  Curtis,  Bot. 

Mag.  4082,  mentioned  the  striking  compression  of  the  hypanthium,  "so  that 

t 


Vol.  XXV]  MUNZ :  THE  GENUS  FUCHSIA  21 

ill  one  direction,  the  diameter  of  the  tube  does  not  appear  to  be  half  what  it  is 
in  another" ;  this  would  account  for  some  of  the  differences  in  herbarium 
specimens.  However,  material  from  Costa  Rica  has  a  long  hypanthium  which 
narrows  appreciably  at  the  summit,  at  least  in  Standley  and  Valerio  43829, 
Pittier  10501,  not  so  much  so  in  Stork  3024.  Curtis  shows  such  a  condition  in 
edge-view,  but  not  in  the  flat  view.  More  material  from  Costa  Rica  may 
warrant  nomenclatorial  segregation. 

Representative  material  of  F.  splendens:  H.  B.  Basil,  May  7,  1849  (NY) ; 
"F.  grandiflora  N."  Sesse,  Mogiilo  et  al  5208  (F).  MEXICO:  F.  splendens 
Zuccar.  ipse,  Hab.  in  Mexico,  Karivinski,  Herbarium  Regium  Monacense 
(photo  F,  GH,  POM).  Chiapas,  without  locality,  Ghieslreght  698  (F,  GH, 
MO)  ;  Mt.  Tacana,  Matuda  2322  (F,  MICH,  NY)  ;  between  San  Cristobal  and 
Tencjapa,  Souviron  and  Erianson  83  (US).  GUATEMALA:  San  Marcos, 
Quebrada  Canjula,  between  Sibinal  and  Canjula,  Volcan  de  Tacana,  Steyer- 
mark  36056  (F) .  Quiche,  Chiul,  Heyde  and  Lux,  under  J.  D.  Smith  2890  (NY, 
GH,  US)  ;  Chimaltenango  :  Chichavac,  Skutch  118  (US) ;  Santa  Elena,  Skutch 
220  (US)  ;  Cerro  de  Tecpam  near  Santa  Clara,  Standley  61000  (F,  NY, 
POM)  ;  Volcan  de  Acatenango,  above  Las  Calderas,  Steyermark  61808  (F). 
Sacatepequez,  Volcan  de  Agua,  J.  Donnell  Smith  2174  (F,  GH,  NY,  US), 
Standley  65133  (F,  POM) ;  Jalapa,  Montaiia  Miramundo,  Steyermark  32833 
(F).  COSTA  RICA:  prov?,  Ojo  de  Agua,  Stork  3024  (F)  ;  Prov.  San  Jose, 
Cerro  de  las  Vueltas,  Pittier  10501  (US),  Standley  and  Valerio  43829  (F, 
US). 

(8)  Fuchsia  cordifolia  Bentham 

(Plate  1,  fig.  7) 
Fuchsia  cordifolia  Bentham,  PI.  Hartweg.,  74,  1841;  Lindl.,  Bot.  Reg.,  27:70,  1841. 

Straggling  shrub  1  m.  or  so  high,  to  a  small  tree,  often  epiphytic,  few- 
branched,  the  younger  branches  somewhat  red,  subglabrous  except  toward  the 
glandular-pub erulent  tips ;  leaves  ovate  to  cordate-ovate,  opposite,  paler  be- 
neath than  above,  puberulent  when  young,  nearly  or  quite  glabrate  in  age, 
more  or  less  abruptly  acuminate,  obscurely  denticulate;  blades  thin,  3-9  cm. 
long,  2-6  cm.  wide,  with  9-11  principal  lateral  veins  on  each  side  of  the  mid- 
rib ;  petioles  2-5  cm.  long ;  flowers  solitary  in  axils  of  upper  often  somewhat 
reduced  leaves;  pedicels  pendant,  subfiliform,  puberulent,  3-8  cm,  long  at 
anthesis,  not  much  longer  in  fruit ;  hypanthium  subcylindric,  dull  red,  puber- 
ulent without,  some  of  the  hairs  gland-tipped,  glabrous  within,  somewhat 
swollen  at  base,  then  slightly  narrowed  and  somewhat  broader  only  near  the 
summit,  3.5-5  (6)  cm.  long,  4-5  mm.  wide  except  at  the  summit,  where  7  mm. 
wide;  sepals  mostly  green  except  at  reddish  outer  base,  ovate-lanceolate, 
puberulent  with  some  hairs  gland-tipped,  acuminate,  11-15  mm.  long,  4-5 
mm.  wide  at  base,  free  tips  1-2  mm.  long;  petals  green  to  olive-green,  broadly 
ovate,  glabrous,  acute  to  subacuminate,  6-8  mm.  long;  episepalous  stamens 
about  as  long  as  sepals,  epipetalous  about  two-thirds  as  long ;  anthers  yellow, 


22  CALIFOEXIA  ACADEMY  OF  SCIENCES  [Proc.  4th  8er. 

2-2.5  mm.  long;  style  glabrous,  slightly  exceeding  the  sepals;  stigma  conic. 
1.5  mm.  long;  fruit  apparently  linear-ellipsoid,  more  than  2  cm.  long. 

Type  locality,  "In  monte  ignivomo  Xetucli  infra  apicem,  altitudine  ped. 
10,000,"  prope  Quezaltenango,  Guatemala.  Tlie  species  is  known  from  eleva- 
tions of  8000-11000  ft.  from  the  mountains  of  Guatemala.  It  is  distinguished 
from  F.  fulgens  by  its  green  petals  and  from  F.  splendens  hy  its  shorter 
stamens,  the  hj^Danthium  being  glabrous  within  and  longer  and  subcylindric. 
Specimens  studied,  GUATEMALA  :  Dept.  San  Marcos,  Volcan  de  Tajumulco, 
between  San  Sebastian  and  Todos  Santos,  Steyermark  36942  (F)  ;  Dept.  de 
Quezaltenango,  Volcan  de  Zunil,  Steyermark  34686  (F);  Cerro  Quemado. 
Kellerman  5953  (US)  ;  Volcan  de  Santa  Maria,  Nelson  3723  (F,  US),  Skntch 
867  (F,  NY),  Standley  67592  (F),  67648  (F)  ;  Xetuch,  Hartiveg  528,  frag- 
ment of  type  number  (F) ,  photo  of  type  no.  (F,  POM)  ;  Dept.  Chimaltenango, 
San  Martin,  Lcivis  936  (F.) 

(9)  Fuchsia  austromontana  Johnston 
(Plate  1,  fig.  8) 

Fuchsia  austromontana  Johnston,  Journ.  Arnold  Arboi-etum,  20:242,  1939. 
F.  serrati folia  Hook.,  Bot.  Mag.,  71  :pl.  4174,  1845  ;  not  of  K.  and  P. 

Loose  bushy  shrubs  up  to  4  m.  tall;  twigs  reddish,  1.5-3  mm.  thick,  glabrous 
except  for  the  pilose  tips ;  leaves  opposite  or  ternate  (or  quaternate  ?) ,  elliptic 
to  elliptic-lanceolate  or  -ovate,  acute  to  obtuse  at  base,  acuminate  at  apex, 
serrulate,  strigulose  (at  least  along  the  veins)  and  light  green  on  upper  sur- 
face, paler  beneath  and  villous  on  veins,  rather  fleshy,  with  8-15  principal 
veins  on  each  side  of  midrib,  the  submarginal  vein  often  inconspicuous ;  blades 
2-10  (12)  cm.  long,  1-3  (4)  cm.  wide,  upper  ones  considerably  reduced; 
petioles  2-10  (15)  mm.  long,  villous;  stipules  lanceolate,  2-3  mm.  long,  decidu- 
ous ;  flowers  in  upper  axils ;  pedicels  stoutish,  1-3.5  cm.  long,  sparsely  strigu- 
lose; ovary  fusiform,  sparsely  strigulose,  8-10  mm.  long,  2-2.5  mm.  thick; 
hypanthium  light  red,  subcA'lindric,  2.5-4  cm.  long,  enlarged  at  base  (3.5-5 
mm.  thick) ,  then  narroAved  for  about  1  cm.  (i.e.  2-3  mm.  thick) ,  then  gradually 
wider  toward  summit  (ca.  7  mm.  wide),  sparsely  villous  without,  villous 
wdthin  lower  portion ;  sepals  red,  lanceolate,  15-17  mm.  long,  4  mm.  wide, 
subacuminate  (bud  subacuminate,  but  scarcely  apiculate),  pubescent  to  vil- 
lous without,  glabrous  within  ;  petals  broadly  obovate,  rounded  at  apex,  1^15 
mm.  long,  11-12  mm.  wide,  red,  sometimes  purple  on  drying;  stamens  16-17 
and  10-11  mm.  long;  anthers  4-5  mm.  long;  .style  glabrous,  2-3  mm.  longer 
than  sepals;  stigma  2-3  mm.  thick;  berry  broadly  ellipsoid,  about  15  mm. 
long,  8  mm.  wide. 

Type  locality,  between  Pillahuata  and  Acanacu,  prov.  Paucartambo,  dept. 
Cuzco,  Peru.  Near  to  F.  dcnticuJata  in  the  broad  hypanthium  and  glabrous 
style  and  to  F.  cordifolia  in  the  broad  hypanthium-base.  It  seems  to  be  rather 
local  in  Paucartambo ;  I  have  seen  the  following  collections  :  PERU :  Paucar- 
tambo :  between  Pillahuata  and  Acanacu,  at  2800  m.,  West  7083,  isotype 


Vol.  XXV]  MUNZ :  THE  GENUS  FUCHSIA  23 

(GH)  ;  Pillahuata,  Cerro  de  Cusilluyoc,  3000-3300  m.  Pennell  14110  (F,  GH, 
US) ;  Cabecera  de  Montana,  eamino  a  Tambomayo,  2000  m..  Vargas  10-7083 
(F)  ;  between  Acliirani  and  Medias-Mayu,  2600  m.,  Vargas  11121  (F)  ;  Acca- 
naco,  4000  m.,  Balls  6708  (US) . 

(10)  Fuchsia  denticulata  Rniz  and  Pavon 

(Plate2,  fig.  9) 

Fuchsia  denticulata  Euiz  and  Pavon,  F1.  Peruv.,  3:87,  pi.  325,  fig.  b,  1802. 

F.  fierrati folia  Euiz  and  Pav6n,  I.e.,  86,  pi.  323,  fig.  a.  Since  tins  name  has  page  priority, 
I  had  intended  to  use  it,  but  must  follow  Art.  56  of  the  International  Eules,  since  Mac- 
bride  took  up  this  name  (Field  Mus.  Nat.  Hist.-Bot.  Ser.,  13,  part  4:553,  1941). 

F.tacsoniiflora  Krause,  Fedde  Eep.  Nov.  Sp.,  1:172,  1905;  based  on  Weberbauer  252, 
which  seems  to  belong  here. 

F.grandiflora  Euiz  ex  Dahlgren,  Field  Mus.  Nat.  Hist.-Bot.  Ser.,  21:212,  1940,  nomen 
nudiim.  Have  seen  photograph  of  Geneva  specimen  (F,  POM). 

Semiscandent  shrub  to  tree,  up  to  12  m.  tall,  the  young  twigs  green  to  red- 
dish, mostly  3-5  mm.  thick,  subterete,  mostly  glabrous,  more  or  less  puberu- 
lent  or  pubescent  toward  tips,  about  younger  nodes,  etc. ;  leaves  opposite  or 
ternate,  elliptic  to  oblong,  rather  fleshy,  subentire  to  somewhat  serrate,  acute 
to  obtuse  at  base,  acute  to  acuminate  at  apex,  darker  above  than  beneath, 
subglabrous  to  puberulent  above,  subglabrous  beneath  or  somewhat  villous 
along  veins,  the  principal  lateral  veins  12-15  on  each  side  of  midrib,  sub- 
marginal  vein  evident;  blades  4-11  cm.  long,  2-4  (5)  cm.  wide;  petioles  5-15 
mm.  long,  1-2  mm.  wide,  glabrous  to  pubescent ;  stipules  lance-deltoid,  ca.  1 
mm.  long,  deciduous;  flowers  solitary  in  axils  of  upper  somewhat  reduced 
leaves;  pedicels  rather  stout,  1.5-4  cm.  long,  usually  minutely  strigulose; 
ovary  subfusif orm,  10-12  mm.  long,  2-3  mm.  wide,  minutely  strigulose ;  hy- 
panthium  tubular,  red,  3.5-4  cm.  long,  5-7  mm.  wide  at  base,  very  little  nar- 
rowed, then  very  gradually  widened  toward  mouth  where  7-10  mm.  wide, 
subglabrous  to  sparsely  villous  without,  usually  densely  villous  within  lower 
portion ;  sepals  red  with  green  tips,  lanceolate,  divaricate,  18-24  mm.  long, 
4-5  mm.  wide,  acute  to  acuminate  (bud  acuminate)  in  the  terminal  3-4  mm. 
but  scarcely  apiculate,  glabrous  to  somewhat  villous  on  backs,  glabrous  on 
inner  surface;  petals  crimson  to  scarlet,  oblong  to  oblong-lanceolate,  acute  to 
obtuse,  15-17  mm.  long,  4-6  mm.  wide,  glabrous ;  stamens  17-20  and  13-16 
mm.  long,  glabrous ;  anthers  white,  4-5  mm.  long ;  style  stout,  glabrous,  equal- 
ling or  exceeding  sepals  by  a  few  mm. ;  stigma  white,  subglobose,  ca.  3  mm. 
thick ;  berry  ellipsoid,  16-19  mm.  long,  ca.  8  mm.  thick,  subglabrous  to  min- 
utely strigulose. 

Type  locality,  Muiia,  dept.  Huanuco,  Peru.  AVith  F.  leptopoda,  etc.  this 
species  forms  a  long-tubed,  axillary-flowered  group  related  to  F.  cordifolia, 
F.  austromontana,  etc.  I  am  unable  to  separate  F.  serratifolia  and  F.  denticu- 
lata, type  material  of  both  of  which  has  been  available.  They  are  characterized 
by  their  stout  twigs,  fleshy,  elliptic  subglabrous  leaves,  tubular  hypanthium, 
non-apiculate  buds,  ellipsoid  berry.  Found  in  Peru  and  Bolivia ;  repreSenta- 


24  CALIFORXIA  ACADEMY  OF  SCIEXCES  [Proc.  4th  Ser. 

tive  material  as  follows:  F.  denticulata,  ex  Herb.  Madrid,  Rtiiz  and  Pavon, 
type  material  (F),  photograph  of  Geneva  material  (F,  POM)  ;  F.  serratifoUa 
ex  Herb.  Madrid,  Ruiz  and  Pavon,  type  material  (F),  photograph  (F,  POM)  ; 
F.  grandiflom,  photograph  of  Geneva  specimen  (F,  POM).  PERU  :  Huaniico, 
15  mi.  se.  of  Huanuco,  3200  m.,  Machride  and  Featherstone  2082  (F,  GH,  US)  ; 
Mito,  2800  m.,  Machride  and  Featherstone  1403  (GH,  F,  US)  ;  west  of  Carpish, 
Stork  and  Horton  9893  (F)  ;  Muiia,  trail  to  Tambo  de  Vaca,  2500  m.,  Machride 
4284  (F,  GH,  US)  ;  Carpis,  between  Acamayo  and  Chinehao,  2800  m.,  Mexia 
04127  (GH)  ;  Lima  to  Oroya,  3200  m.,  Wcherhauer  252,  type  tacsoniiflora  at 
Berlin  (photo  F,  PO:\r)  ;  Rio  Blanco,  Lima  at  3600  m.,  Machride  and  Feather- 
stone 723  (F,  GH)  ;  Quebrada  de  San  Mateo,  Isern  2546  (F).  Ayacucho, 
Prov.  Huanta,  Choimacota  Valley,  2900  m.,  Wcherhauer  7587  (F,  GH,  US). 
Huaneavelica :  Prov.  Tayacaja,  Montifungo,  east  of  Surcubamba,  3000  m., 
Stork  and  Horton  10367  (F)  ;  Salcabamba,  3250  m.,  Stork  and  Horton  10261 
(F).  Cuzco:  Prov.  Paucartambo,  Rio  Tambomayo,  1700-2000  m.,  West  7093 
(GH).  BOLIVIA :  La  Paz :  Nor  Yungas :  Unduavi,  3000  m.,  Eyerdam  25137 
(F) ;  Sur  Yungas  :  Unduavi,  3100  m.,  Buchtien  143  (F,  GH,  1<^Y),2923  (NY, 
US)  ;  Yungas,  Ba^ig  731  (F,  GH,  NY,  US),  Rnshij  1801  (NY)  ;  Combaya, 
Prov.  Larecaja,  3000-3300  m.,  Mandon  623  (GH). 

(11)  Fuchsia  leptopoda  Krause 

(Plate  2,  fig.  10) 

Fuchftia  leptopoda  Krause,  Pedde  Eep.,  Nov.  Sp.,  1:171, 1905. 
F.  siphonantha  Krause,  I.e.,  173. 

Scandent  to  erect  shrub,  up  to  3  m.  tall,  the  twigs  dark,  glabrous  to  strigu- 
lose  toward  tips,  terete,  1-3  mm.  thick;  leaves  opposite  or  ternate,  elliptic- 
lanceolate  or  -oblanceolate,  acute  at  base,  acuminate  at  apex,  serrulate  above, 
light  green  and  glabrous  or  strigulose  on  veins,  paler  beneath  and  glabrous 
except  for  pilose  veins,  the  principal  lateral  veins  mostly  7-10  on  each  side 
of  midrib,  the  submarginal  vein  more  or  less  developed;  blades  4-9  cm.  long, 
1.5-2.5  (3)  cm.  wide;  petioles  slender,  5-15  mm.  long,  somewhat  pubescent; 
stipules  lanceolate,  3  mm.  long,  deciduous ;  flowers  in  axils  of  often  reduced 
foliage  leaves;  pedicels  slender,  subglabrous,  3-5  cm.  long;  ovary  fusiform, 
glabrous  to  puberulent,  5-8  mm.  long ;  hypanthium  dark  red,  tubular,  3.5-4.5 
cm.  long,  somewhat  bulbous  at  base  and  ca.  3  mm.  thick,  then  narrowed  to 
width  of  1.5-2  mm.  for  about  1  cm.,  then  widened  toward  mouth  where  6-7 
mm.  wide,  glabrous  to  sparsely  villous  without,  densely  villous  within  lower 
portion ;  sepals  deep  red,  lance-linear,  17-20  mm.  long,  3.5-4  mm.  wide,  acumi- 
nate (bud  very  slender  and  long-acuminate,  with  tip  4-5  mm.  long),  glabrous 
to  pubescent  on  back,  glabrous  within,  petals  fiery  red,  oblong,  17-18  mm. 
long,  4-7  mm.  wide,  acute,  glabrous  or  sparsely  villous  on  back  near  base ; 
stamens  18-22  and  12-15  mm.  long,  glabrous;  anthers  3-4  mm.  long;  style 
glabrous,  equalling  sepals  or  exceeding  them  by  1  cm.,  stigma  subglobose, 
obscurely  lobed,  2-3  mm.  wide ;  fruit  narrow,  over  1  cm.  long. 


Vol.  XXV]  MUNZ :  THE  GENUS  FUCHSIA  25 

Type  locality,  PERU :  Juniii :  Prov.  Tarma,  between  Huacapistana  and 
Palea.  I  have  seen  the  following  examples :  PERU :  Junin :  Prov.  Tarma : 
between  Huacapistana  and  Palea,  2200-2500  m.,  Weherhauer  1772,  type  at 
Berlin  (photo  at  F,  POM)  ;  Montibus  ab  Huacapistana  ad  orientem  versus, 
2600-2700  m.,  Welerlauer  2178  type  siphona7itha  at  Berlin  (photo  F,  POM) . 
Ayacucho  :  Prov.  De  La  Mar :  between  Tambo  and  the  Apurimac,  Yanamonte, 
2700-2800  m.,  Weherhauer  5641  (F,  GH,  US)  ;  Prov.  Huanta,  Choimacota 
Valley,  2900  m.,  Weherhauer  7587a  (F,  GH).  Cuzeo :  Paucartambo:  Tres 
Cruces,  above  Cosnipata,  Weherhauer  6934  (F,  GH,  US).  In  general,  this 
species  is  very  near  to  F.  denticulata  but  with  a  more  narrow  tube  and  leaves, 
long-pointed  buds,  etc. 

(12)  Fuchsia  Woytkowskii  Macbride 

(Plate2,  fig.  11) 

Fuchsia  WoyfkowsTcii  Macbride,  Field  Mus.  Nat.  Hist.,  Bot.  Ser.,  13:  part  4:566,  1941. 

Shrub  up  to  4  m.  tall,  the  young  twigs  glabrous,  purplish  red,  slender; 
leaves  opposite  or  ternate,  elliptic-lanceolate  to  -ovate,  acute  at  base,  acute  to 
acuminate  at  apex,  subentire,  rigid,  dark  and  glabrous  above,  somewhat  paler 
and  glabrous  beneath  except  for  the  minute  appressed  puberulence  on  the 
veins,  principal  lateral  veins  13-15  on  each  side  of  the  midrib,  submarginal 
vein  evident;  blades  4-8  cm.  long,  1.5-3  cm.  wide;  petioles  3-6  mm.  long; 
stipules  subulate,  2  mm.  long,  deciduous;  flowers  solitary  in  axils  of  upper 
leaves  or  apparently  subterminal  on  short  branches ;  pedicels  purplish,  sub- 
filiform,  nearly  glabrous,  1-2.5  cm.  long;  ovary  fusiform,  5-6  mm.  long,  2  mm. 
wide;  hypanthium  deep  vermilion,  3.5-4.7  cm.  long,  bulbous  at  base,  2-2.5 
mm.  wide,  then  narrowed  for  one-third  its  length  then  gradually  widened  until 
6-7  mm.  wide  at  mouth,  glabrous  without,  pilose  within  the  basal  two-thirds ; 
sepals  vermilion,  spreading,  12-14  mm.  long,  4-5  mm.  wide,  glabrous,  sub- 
ulate for  2-3  mm.  at  apex  (the  tips  apparently  not  connivent  in  the  bud)  ; 
petals  bright  red,  oblong-elliptic,  12-13  mm.  long,  glabrous,  erose-rounded  at 
apex;  stamens  10  and  8  mm.  long;  anthers  2  mm.  long;  style  glabrous,  sur- 
passing sepals  by  2-3  mm. ;  stigma  2  mm.  thick.  , 

Type,  PERU :  Dept.  Amazonas  :  near  Almirante,  at  1900  m.,  F.  WoytkowsH 
(F).  This  one  and  only  specimen  is  rather  inadequate,  but  apparently  does 
not  fit  into  any  of  the  previously  described  species.  It  seems  near  F.  leptopoda, 
but  the  hypanthium  is  more  slender  and  not  glabrous  within,  and  the  sepals 
are  shorter  and  with  free  tips  in  the  bud. 

(13)  Fuchsia  magdalenae  Munz,  new  species 

(Plate2,  fig.  12) 

Shrub;  young  twigs  glabrous,  purplish,  1.5-3  mm.  thick;  leaves  opposite, 
ternate  or  quaternate,  coriaceous,  elliptic,  rounded  at  base,  abruptly  acumi- 
nate at  apex,  inconspicuously  denticulate,  quite  glabrous  except  for  the  ciliate 
margins,  somewhat  paler  beneath  than  above,  principal  lateral  veins  dark. 


26  CALIFOIIXIA  ACADEMY  OF  SCIENCES  [Proc.  4th  Ser. 

8-10,  the  submarginal  vein  more  or  less  distinct;  blades  3-6  cm.  long,  1.5-2.5 
cm.  wide,  the  upper  reduced;  petioles  dark,  glabrous,  5-6  (10)  mm.  long; 
stipules  subulate,  1  mm.  long,  deciduous;  flowers  solitary  in  upper  axils,  sub- 
filiform,  glabrous,  4-5  cm.  long,  dark;  ovarj'  glabrous,  7-11  mm.  long,  2-2.5 
mm.  wide ;  hypanthium  purplish  at  base,  brigliter  red  beyond,  5-6  cm.  long, 
about  4  mm.  wide  at  base,  then  not  constricted  but  tubular  and  gradually 
enlarged  until  8-9  mm.  wide  at  mouth,  glabrous  without  and  within;  sepals 
lance-oblong,  16-19  mm.  long,  4.5-5  mm.  wide,  red,  glabrous,  acute,  not  sub- 
ulate at  apex;  petals  scarlet,  oblong-ovate,  15-18  mm.  long,  7-8  mm.  wide, 
acutish ;  stamens  17  and  13  mm.  long ;  anthers  3.5  mm.  long ;  style  slender, 
glabrous,  exceeding  sepals  by  5-6  mm.;  stigma  4-lobed,  4  mm.  wide;  berry 
ellipsoid,  ea.  17  mm.  long,  8  mm.  wide. 

Frutex;  ramulis  glabris,  subpurpureis,  1.5-3  mm.  diametro ;  foliis  oppositis,  ternatis  vel 
quaternis,  coriaceis,  ellipticis,  basi  rotimdatis,  apice  abrupte  aeumiiiatis,  subglabris;  laminis 
3-6  cm.  longis,  1.5-2.5  cm.  latis;  petiolis  glabris,  5-6  (10)  mm.  loiigis;  stipulis  siibulatis, 
1  mm.  longis,  caducis;  floribus  solitariis  in  axillis  superioribus ;  pedicellis  subfiliformibus, 
glabris,  4—5  cm.  longis;  ovario  glabro,  7-11  mm.  longo;  hypanthio  rubro,  5-6  cm.  longo, 
basi  4  mm.  lato,  tubuloso,  supra  8-9  mm.  lato,  c.xtus  intiisquo  glabro ;  sepalis  lanceolato- 
oblongis,  16—19  mm.  longis,  4.5-5  .mm.  latis,  rubris,  glabris,  acutis;  petalis  coccincis,  oblongo- 
ovatis,  15-18  mm.  longis,  7-8  mm.  latis,  subacutis ;  staminibus  17  vel  13  mm.  longis ;  antheris 
3.5  mm.  longis;  stylo  glabro,  sepala  excedente;  stigmate  4— lobato;  bacca  ellipsoidea,  ca.  17 
mm.  longa,  8  mm.  lata. 

Type :  Colombia  :  Dept.  Magdalena :  above  San  Miguel,  at  edge  of  paramo, 

3000  m.,  July,  1932,  Seifriz  392,  United  States  Nat.  Herb.  no.  1572275.  A 

second  collection  from  the  same  department  is  from  between  Pueblo  Viejo  and 

San  Miguel,  900-1700  m.,  8eifriz  537  (US).  The  proposed  species  is  related 

to  F.  dcnticulata  in  its  tubular  hypanthium ;  it  is  characterized  by  its  long 

hypanthium,  thickish  leaves,  glabrous  condition,  non-acuminate  sepals  and 

northern  range.  /-i.N-rr     i,  •  t-.     xi 

(14)  Fuchsia  canescens  Bentham 

(Plate  2,  fig.  13) 

Fuchsin  canescens  Benth.,  PI.  Hartweg.,  178,  1845. 
F.  eorollata  Benth.,  I.e.,  179.  Type  number  seen. 

F.vulcnnica  Andr^,  Rev.  Hortic,  233  and  268,  1888;  from  Yolcan  de  Azufral,  type  not 
seen,  but  characters  as  given  agree. 

Erect  or  clambering  shrubs  up  to  3  m.  tall,  freely  branched,  tlie  young  twigs 
terete,  dark,  hirtellous  with  more  or  less  appressed  short  hairs;  leaves  opposite, 
ternate  or  quaternate,  crowded,  coriaceous,  often  with  revolute  margins,  ellip- 
tic-ovate, obtuse  or  rounded  at  base,  obtuse  to  acute  at  apex,  serrulate,  dark 
green  and  pubescent  to  glabrescent  above,  paler  beneath  and  pilose  on  veins, 
sometimes  more  generally,  the  principal  lateral  veins  mostly  4-7  on  each  side 
of  the  midrib,  submarginal  vein  often  not  conspicuous ;  leaf-blades  2-4  cm. 
long,  1-2  cm.  wide;  petioles  3-5  (8)  mm.  long,  pubescent;  stipules  lance- 
subulate,  1-1.5  mm.  long,  deciduous;  flowers  axillary ;  pedicels  slender,  pubes- 
cent, 1-2  cm.  long ;  ovary  ellipsoid,  pubescent,  5-6  mm.  long ;  hypanthium  deep 
scarlet,  purplish  at  base,  3.5-4.5  cm.  long,  bulbous  at  base  and  2.5-3  mm.  thick. 


Vol.  XXV]  MUNZ:  THE  GENUS  FUCHSIA  ■  27 

then  abruptly  narrowed  to  1-2  mm.  for  one-third  its  length,  then  rather 
gradually  widened  until  6-7  (9)  mm.  wide  at  summit,  more  or  less  pilose 
without,  pilose  within  the  lower  half  or  almost  throughout ;  sepals  deep  scarlet, 
more  or  less  divergent  to  almost  spreading,  10-16  mm.  long,  4—5  mm.  wide, 
acuminate,  with  subulate  tips  about  1  mm.  long,  pilose  without,  glabrous 
within;  petals  scarlet,  sometimes  drying  purplish,  rhombic-ovate  to  -obovate, 
acute  to  obtuse  or  almost  rounded  at  apex,  from  about  as  long  as  to  exceeding 
sepals,  10-16  mm.  long,  6-12  mm.  wide ;  stamens  10-12  and  7-8  mm.  long ; 
anthers  1.5-2  mm.  long ;  style  pilose  within  much  of  the  hypanthium,  about  as 
long  as  sepals ;  stigma  scarlet,  about  2  mm.  thick ;  fruit  apparently  ellipsoid. 

Type  locality,  ascent  to  Paramo  Guanacas,  near  Popoyan,  Cauca,  Colombia. 
Extending  from  Calda,  Colombia  into  Ecuador.  Kepresentative  material, 
COLOMBIA  :  Caldas :  Paramo  del  Quindio,  3800-4000  m.,  Pennell  and  Hazen 
10058  (GH,  NY).  Antioquia  :  Paramo  de  Ruiz,  2800-3300  m.,  LeJimann  3067 
(US.  Cauca :  Paramo  de  Buena  Vista,  Huila  group,  central  Cordillera,  3100 
m.,  Pittier  1188  (US)  ;  ad  Paramo  Guanacas,  near  Popoyan,  Hartweg  992, 
type  no.  canescens  at  Geneva  (photo  F,  POM,  leaf  F)  ;  in  sylvis  Purace,  3100 
m.,  ad  Paramo  de  Guanacas,  near  Popoyan,  Hartweg  993,  type  no.  corollaia 
at  Berlin  (Photo  F,  POM,  leaves  F).  Nariiio :  Volcan  de  Galera  above  Pasto, 
2600-2800  m.,  Lehmann  5615  5  (F)  ;  Altos  Caballos,  3000  m.,  Triana  3808 
(US).  ECUADOR:  Carchi:  Nudo  de  Boliche,  Voladero,  4000  m.,  Penland 
and  Summers  915  (F,  POM) ;  Canton  Tulcan,  road  Tulcan  to  Pun,  3200  m., 
Mexia  7584  (POM,  US)  ;  Paramo  del  Azufral,  east  of  Angel,  Mexia  7512  (NY, 
US).  Bolivar :  Simiatuz :  Hacienda  Talahua,  3200  m.,  Penland  and  Summers 
623  (F,  POM).  Tungurahua:  Paramo  de  Minza,  Minza  Chica,  3800  m.,  Pew- 
kmd  and  Summers  337  (F,  POM). 

F.  canescens  comes  in  a  broad-petaled  group  which  seems  to  me  to  be  related 
to  the  F.  cordifolia  group  in  its  broad  hypanthium-base  and  in  haA'ing  the 
hypanthium  quite  inflated  in  its  upper  third.  The  species  is  near  F.  ayavacensis 
but  differs  in  the  smaller  thicker  leaf -blades,  shorted  pedicels,  the  petals  nearly 
as  long  as  or  longer  than  the  sepals,  the  latter  being  shorter  and  more  spread- 
ing than  in  ayavacensis.  There  is  some  question  as  to  w^hether  my  treatment  of 
canescens  is  too  inclusive ;  Penland  and  Summers  623,  andPJ'5  and  Mexia  7512 
having  the  petals  rather  round  and  shorter  than  the  sepals ;  Hartweg  992, 
Pittier  1382,  and  Penland  and  Summers  623  having  the  upper  surface  of  the 
leaves  finely  strigose.  But  the  whole  range  of  variation  seems  such  as  not  to 
make  possible  a  taxonomic  differentiation  with  the  amount  of  material  avail- 
able. The  leaves  from  the  type  collections  of  canescens  and  coroUata  are  quite 
alike. 

(15)  Fuchsia  rivularis  Macbride 

Fuchsia  rivularis  Macbride,  Candollea,  8:24,  1940;  Field  Mus.  Nat.  Hist.,  Bot.  Ser.,  13, 
part  4:562, 1941. 

Branchlets  puberulent;  leaves  ternate,  ovate-elliptic,  abruptly  acuminate, 
entire,  glabrous  above,  loosely  pilese  beneath,  the  blades  5-6.5  cm.  long,  2.5-3 


28  CALIFORNIA  ACADEMY  OF  SCIENCES  [Proc.  4th  Ser. 

cm.  wide;  petioles  lacking  or  nearly  so;  flowers  solitary  in  axils  of  somewhat 
reduced  upper  leaves;  pedicels  1  cm.  long;  hypanthium  4.5  cm.  long,  sparsely 
puberulent  without,  pilose  within  lower  portion,  gradually  ampliate ;  sepals 
narrowly  lanceolate,  long-acuminate,  2  cm.  long;  petals  oblong,  about  as  long 
as  sepals;  stamens  shorter. 

Based  on  a  Mathews  collection  from  Chachapoyas,  Amazonas,  Peru  and  said 
to  be  near  F.  ayavacensis  or  F.  hirtella,  but  with  narrower  petals  and  more 
sessile  and  wider  leaves.  I  have  seen  no  material  nor  photographs,  but  except 
for  the  larger  leaves  and  longer  petioles  it  seems  much  like  F.  canescens. 

(16)  Fuchsia  loxensisH.B.K. 

(Plate  2,  fig.  14) 

Fuchsia  loxemis  H.B.K.,  Nov.  Gen.  et  Sp.,  6:106,  pi.  536,  1823. 

F.  umbrosa  Bentii.,  PI.  Ilartw.,  176,  1845  ;  I  have  seen  a  photograph  of  the  type. 

F.  apiculata  Johnson,  Contr.  Gray  Herb.,  75:34,  1925.  Type  seen. 

Shrub,  slender,  stiff  and  bushy  or  more  or  less  scandent  or  leaning,  com- 
monly up  to  2  or  3  m.  tall,  the  younger  twigs  slender,  1-3  mm.  thick,  purplish 
red,  loosely  appressed-puberulent,  with  whitish  or  yellowish  hairs ;  leaves  op- 
posite to  ternate,  sometimes  quaternate,  firm,  elliptic  to  elliptic-oblong, 
rounded  to  acute  at  base,  mostly  acute  at  apex,  subentire  to  remotely  denticu- 
late, shining,  deep  green  and  minutely  appressed-puberulent  to  glabrescent 
above,  paler  and  puberulent  beneath,  also  more  or  less  pilose  beneath  espe- 
cially along  or  near  the  midrib,  principal  lateral  veins  often  reddish  beneath, 
prominent,  about  10  on  each  side  of  the  midrib,  the  submarginal  vein  present ; 
leaf -blades  2.5-6  (7)  cm.  long,  1.2-2  (3)  cm.  wide ;  petioles  slender,  strigulose, 
1-3  cm.  long;  stipules  subulate-deltoid,  1-2  mm.  long,  deciduous:  flowers 
solitary  in  axils ;  pedicels  slender,  strigulose,  and  sometimes  pilose,  1-2.5  cm. 
long ;  ovary  ellipsoid,  strigulose,  3-5  mm.  long ;  hypanthium  deep  red,  20-25 
(30)  mm.  long,  the  base  2-3  mm.  thick,  then  very  slightly  narrowed,  then 
gradually  widened  until  5-6  mm.  wide  at  the  mouth,  shining  and  somewhat 
strigulose  without,  densely  white-pubescent  within  the  lower  half;  sepals 
scarlet,  lance-oblong,  8-10  mm.  long,  3.5-4  mm.  wide,  divergent,  puberulent 
to  sparsely  pilose,  the  subulate  tips  about  1  mm.  long ;  petals  ovate,  dull  red, 
6-9  mm.  long,  5-6  mm.  wide,  obtuse ;  stamens  7-8  and  5-6  mm.  long,  anthers 
2  mm.  long;  style  pubescent  in  lower  part,  red,  exceeding  sepals  by  3.5  mm.; 
stigma  2  mm.  wide ;  fruit  ellipsoid,  8-15  mm.  long,  5-6  mm.  thick. 

Type  locality,  near  Loja  (Loxa),  Ecuador.  The  species  is  apparently  con- 
fined to  the  highlands  of  Ecuador,  growing  at  from  2000-3400  meters.  Repre- 
sentative material  seen,  ECUADOR:  without  definite  locality:  Eastern  Cor- 
dillera. Rimhach  6  (GH,  NY,  US)  ;  in  Andibus  Ecuadorensibus,  Spruce  5203 
(GH,  NY).  Pinchincha:  Hacienda  de  Pinantura  near  Quito,  Hartiveg  983, 
type  no.  umbrosa  at  Geneva  (photo  F,  POM);  near  Tambillo,  Penland  and 
Summers  959  (P,  POM)  ;  Qiiitig-Tesalia  (iMaehachi),  Firmin  666  (F,  GH, 
US).  Leon:  Cotopaxi :  road  Pilaloa  to  Macuchi,  Hmiglit  2955  (POM,  US). 


Vol.  XXV J  MUNZ :  THE  GENUS  FUCHSIA  29 

Tunguraliua  :  west  of  Ambato,  Heinrichs  828  (NY)  ;  south  of  Baiioa,  Penland 
and  Summers  93  (F,  POM).  Bolivar:  Simiatug:  Hacienda  Talahua,  Penland 
and  Summers  520  (F,  POM) .  Azuay  and  Caiiar :  between  Cuenca  and  Huigra, 
Hitchcock  21667,  type  no.  apiculata  (GH,  NY,  US).  Loja :  mts.  near  Loxa, 
Hartweg  733  (NY)  ;  F.  loxensis,  Humboldt  in  "Willdenow  Herbarium  at 
Berlin  (photo,  F,  POM). 

F.  loxensis  seems  near  to  F.  canescens  in  its  leaves,  broad  petals,  etc.,  but 
has  shorter  hypanthia  and  somewhat  narrower  petals;  Spruce  5203,  above 
cited,  is  quite  intermediate.  It  is  near  F.  decussata  in  some  ways,  but  has 
longer  hypanthia,  less  serrate  leaves  and  less  evident  stamens. 

(17)  Fuchsia  pallescens  Diels 

Fuchsia  pallescens  Diels,  Notizbl.  d.  bot.  Gart.  u.  Mus.  zu  Berlin-Dahlem,  14:34,  1938. 

Medium  sized  shrub,  younger  branches  subglabrous ;  leaves  opposite  or 
ternate,  membranaceous,  subovate,  subacuminate  at  both  ends,  callous-  den- 
ticulate, almost  glabrous ;  blades  6-7  cm.  long,  3.5-4  cm.  wide ;  petioles  2-3.5 
cm.  long;  flowers  axillary,  but  arranged  subcorymbosely  near  tips  of  branches ; 
pedicels  8-15  mm.  long ;  ovary  attenuate  at  apex ;  hypanthium  pale  carmine, 

2  cm.  long,  nodose  at  base,  then  constricted,  then  gradually  widened  until  4 
mm.  wide  at  mouth,  glabrous  without;  sepals  white,  lanceolate,  very  acute, 
10-12  mm.  long,  3  mm.  wide  at  base;  petals  purplish,  6-7  mm.  long;  stjde  with 
globose  stigma. 

Type  locality,  ECUADOR:  Prov.  Tunguragua :  east  of  Patate,  montane 
forest  above  Leito  at  2750  meters.  I  have  seen  no  material,  but  the  plant 
would  seem  to  be  near  F.  loxensis  and  distinct  from  it  in  being  more  glabrous, 
having  wider  more  plainly  toothed  leaves,  and  in  its  white  sepals. 

(18)  Fuchsia TownsendiiJohnston 

(Plate  3,  fig.  15) 
Fuchsia  Townsendii  Johnston,  Contr.  Gray  Herb.,  75:33,  1925. 

Shrub,  the  young  twigs  purplish,  densely  spreading-pubescent ;  leaves 
crowded,  ternate,  elliptic-oblanceolate,  membranaceous,  acute  at  base,  acum- 
inate at  apex,  remotely  denticulate,  green  and  sparsely  and  finely  pubescent 
above,  paler  and  densely  canescent-villous  beneath,  the  principal  lateral  veins 
10-12  on  each  side  of  midrib ;  blades  6-13  cm.  long,  2-3  cm.  wide ;  petioles 
pubescent,  5-20  mm.  long;  stipules  subulate,  1-2  mm.  long,  deciduous ;  flowers 
few,  axillary;  pedicels  slender,  pubescent,  1.5-2.5  cm.  long,  bulbous  at  base 
where  2  mm.  thick,  then  narrowed  for  one-third  its  length  to  1.5  mm.,  then 
gradually  widened  to  5-6  mm.  at  mouth,  densely  pubescent  without  and 
within  the  lower  half;  sepals  10-12  mm.  long,  ascending,  lanceolate,  acute, 
4  mm.  wide,  pubescent  on  back,  glabrous  on  upper  or  inner  surface ;  petals 
ovate,  9-10  mm.  long,  8  mm.  wide,  obtuse;  stamens  13  and  9  mm.  long ;  anthers 

3  mm.  long ;-  style  pubescent  in  lower  part,  slender,  not  exceeding  sepals ; 
berry  hairy,  narrowly  ellipsoid,  over  1  cm.  long. 

Type  locality,  Sabiango  Hill,  Ecuador,  tj^pe  collected  by  C.  H.  Townsend, 


30  CALIFOBNIA  ACADEMY  OF  SCIENCES  [Proc.  4th  Ser. 

number  A93  (type  at  US,  fragment  GH).  The  type  collection  is  the  only 
material  I  have  seen.;  it  is  distinct  in  its  large  thin  hairy  leaves,  long-  narrow 
hypanthium  and  short  sepals  and  petals.  F.  Schcrffiana  Andre  (Rev.  Ilortic, 
1888,  233,  268)  may  be  an  earlier  name  for  this  species,  but  since  I  have  not 
seen  certain  material  I  am  not  sure  as  to  width  of  petals. 

(19)  Fuchsia  platypetala  Johnston 

(Plate3,  fig.  16) 
Fuchsia  platypetala  Johnston,  Journ.  Arnold  Arbor.,  20:241,  1939. 

Shrub  2-3  m.  tall,  the  younger  branches  2-4  mm.  thick,  terete,  somewhat 
reddish,  pubescent;  leaves  opposite  or  ternate,  elliptic-lanceolate,  membrana- 
ceous, acute  at  both  ends,  serrulate,  pilose-pubescent  on  both  surfaces  espe- 
cially when  young,  principal  lateral  veins  8-11  on  each  side  of  midrib,  with 
submarginal  vein  present ;  blades  2.5-7.5  cm.  long,  1-2.3  cm.  wide ;  petioles 
slender,  4-15  mm.  long,  pilose;  stipules  subulate-deltoid,  ca.  1  mm.  long; 
flowers  in  upper  axils;  pedicels  slender,  2-3.5  cm.  long,  somewhat  pilose; 
ovary  ellipsoid,  8-10  mm.  long,  pilose;  hj^Danthium  red,  4.5-5  cm.  long, 
bulbous  at  base  where  3-4  mm.  thick,  then  2-3  mm.  wide  for  one-third  its 
length,  then  gradually  widened  until  8-9  mm.  wide  at  mouth,  sparsely  pilose 
externally,  densely  so  within  for  two-thirds  its  length ;  sepals  lanceolate, 
crimson,  15-20  mm.  long,  3—4  mm.  wide,  apiculate  for  2-3  mm.,  sparsely  pilose 
without,  glabrous  within  ;  petals  crimson  but  with  central  white  oblong  blotch, 
obovate,  rounded  or  broadly  obtuse,  14-16  mm.  long,  10-12  mm.  wide ;  stamens 
16-17  and  12-13  mm.  long;  anthers  3  mm.  long;  style  pilose  through  most  of 
its  length,  reddish ;  stigma  slightly  lobed,  3  mm.  thick. 

Type  locality,  Chincheros,  Apurimac,  Peru.  Material  seen,  PERU:  Dept. 
Apurimac :  Chincheros,  at  2930  m..  West  3705  (GH,  type).  Cuzco,  cultivado 
en  los  jardines,  Herrera  1514  (GH).  The  plant  would  seem  in  its  wide-based 
hypanthium  to  be  in  the  denticulata,  austromontana,  leptopoda  group  but 
differs  in  its  hairy  style,  thinner  leaves,  wider  petals,  and  whitish  blotch  on 
each  petal. 

(20)  Fuchsia  macrostigma  P>entham 
Fuchsia  macrostigma  Bentham,  PI.  Hartw.,  129.  Dec,  1844. 

Erect  shrub,  0.5-1.5  m.  tall,  apparently  rather  openly  branched,  the  young 
twigs  green  to  reddish,  herbaceous,  terete  or  angled,  1.5-3  mm.  thick,  glabrous 
to  pubescent ;  leaves  opposite,  membranaceous,  broadly  to  narrowly  elliptic, 
acute  to  obtuse  at  base,  abruptly  acute  to  subacuminate  at  apex,  obscurely 
denticulate,  ciliate,  dark  .green  and  subglabrous  to  puberulent  above,  lighter 
and  subglabrous  to  puberulent  beneath,  purplish  on  veins,  principal  lateral 
veins  12-16  on  eacli  side  of  midrib,  submarginal  vein  developed ;  blades  8-18 
cm.  long,  4-7  (9)  cm.  wide;  petioles  1-3  cm.  long;  stipules  deltoid,  2-3  mm. 
long,  more  or  less  persistent ;  flowers  solitary  in  upper  axils ;  pedicels  coarse. 


Vol.  XXV]  MUNZ:  THE  GENUS  FUCHSIA  31 

ascending,  1-2  cm.  long;  ovary  linear,  6-10  mm.  long,  2-3  mm.  thick;  hypan- 
tliiuni  tubular,  reddish  purple,  especially  on  lower  half,  6-7  cm.  long,  2-3.5 
mm.  thick  at  base,  then  very  slightly  narrowed  for  one-third  its  length  and 
gradually  widened  to  6-8  mm.  at  mouth,  puberulent  to  pilose  without,  gla- 
brous within;  sepals  lance-oblong,  purplish  red  or  lighter,  spreading,  16-23 
mm.  long,  7-8  mm.  wide,  subulate  for  about  2  mm.  at  tip,  pilose  to  puberulent 
without,  glabrous  to  pilose  within ;  petals  orbicular-obovate,  12-17  mm.  long, 
somewhat  wider,  cerise  to  crimson,  apically  rounded ;  stamens  12-13  and 
9-10  mm.  long;  anthers  light,  about  3  mm.  long;  style  stout,  sparsely  pubes- 
cent to  glabrous,  extending  about  to  end  of  sepals ;  stigma  subglobose,  4-lobed, 
green,  about  4  mm.  wade ;  fruit  elongate,  more  than  1  cm.  long. 

Characterized  by  its  large  leaves,  purplish  color  on  stems,  veins,  hypan- 
thium,  etc.,  almost  glabrous  condition,  wide  petals  and  long  cylindrical 
hypanthium,  this  species  seems  near  F.  platypetala,  rividaris,  etc. 

Key  to  Varieties  of  Fuchsia  macrostigma 

A.  Leaves  subglabrous  except  on  veins ;  hypanthium  more  or  less  pilose.  Provinces  CarcM 

to  Pichincha,  Ecuador 20a.  var.  longiflora. 

AA.  Leaves  minutely  puberulent;   hypanthium  densely  close-pubescent.  Provinces  Chim- 
borazo  to  El  Oro,  Ecuador 20b.  var.  typica. 

(20a)  Fuchsia  macrostigma  Benth.  var.  longiflora  (Benth.)  Munz,  new  comb. 

F.  longiflora  Bentham,  PI.  Hartw.,  177,  1845. 

F.  spectabilis  Hook.,  Bot.  Mag.,  74:pl.  4375,  1848. 

Leaves  subglabrous  except  on  veins ;  hypanthium  more  or  less  pilose. 

Type  locality,  Andes  of  Quito,  Ecuador,  toward  Nanegal.  Material  seen, 
ECUADOR  :  Carchi :  Olivos,  3200-3500  m.,  Mexia  7462  (US) .  Bolivar :  Simi- 
atuz  :  Hacienda  Talahua,  2300  m.,  Penland  and  Summers  657  (F,  POM),  614 
(POM).  Pichincha:  Quitensian  Andes,  Couthouy  in  1855  (GH)  ;  Montis 
Pichincha,  Jameson  in  1856  (GH,  NY) .  I  have  not  seen  the  type. 

(20b)  Fuchsia  macrostigma  Benth.  var.  typica  Munz,  new  name 

(Plate  3,  fig.  17) 

F.  macrostigma  Bentham,  PI.  Hartw.,  129,  1844. 

F.  speciahilis  Hook.,  var.  puhens  Johnston,  Contr.  Gray  Herb.,  75  :34,  1925. 

Leaves  minutely  puberulent ;  hypanthium  densely  close-pubescent. 

Type  locality,  "In  montibus  Pacha,"  southern  Ecuador.  Benth.,  I.e.,  113 
says  for  nos.  708-869 :  "in  montibus  circa  Loxa" ;  since  this  species  comes 
between  nos.  733  and  734,  one  would  infer  that  it  came  from  that  region.  I 
have  not  seen  the  type  number,  but  the  description  fits  very  well  the  following 
plants  :  ECUADOR :  Chimborazo  :  Huigra  :  Hacienda  de  Licay,  /.  N.  and  G. 
Rose  22479,  type  no.  of  pubens  (GH,  NY,  US).  Oro :  between  La  Chorita  and 
Portovelo,  ffifc/icocA^  5:Z:Z^8  (GH,  NY,  US). 


32  CALIFOBNIA  ACADEMY  OF  SCIENCES  [Proc.  4th  Ser. 

(21)  Fuchsia  ayavacensis  H.B.K. 
(Plate  3,  fig.  18) 
Fuchsia  ayavacensis  H.B.K.,  Nov.  Gen.  et  Sp.,  6:107,  1823. 
F.  ampliata  Bentham,  PI.  Hartw.,  178,  1845. 
F.  Hitchcockii  Johnston,  Contr.  Gray  Herb.,  75:  33,  1925. 

Branching,  sometimes  scandent  shrub  up  to  3  m.  high,  younger  branches 
hirtellous-pilose,  more  or  less  purplish,  terete,  1.5-3  mm.  thick;  leaves  ternate 
or  quaternate,  rather  crowded,  membranaceous,  elliptic-lanceolate  to  -oblance- 
olate,  or  -ovate,  acute  to  obtuse  at  base,  acute  to  acuminate  at  apex,  remotely 
denticulate,  principal  veins  purplish,  laterals  6-10  on  each  side  of  midrib, 
the  submarginal  present ;  leaf-blades  pilose  or  pubescent  and  dark  green 
above,  paler  and  pilose  beneath  especially  along  veins,  4—9  cm.  long,  1.5-3  cm. 
"vvide ;  petioles  pilose,  slender,  mostly  5-20  mm.  long ;  stipules  subulate-lance- 
olate, 1.5-2  mm.,  long,  deciduous;  flowers  in  upper  axils;  pedicels  slender, 
pubescent,  2.5-4  (5)  cm.  long;  ovary  oblong-fusiform,  hairy,  6-7  mm.  long; 
hypanthium  deep  red,  3.5-5  (6)  cm.  long,  bulbous  at  base  and  3.5-5  mm.  wide, 
then  constricted  for  1-2  cm.  and  2-3  mm.  wide,  then  quickly  widened  until 
7-11  mm.  wide  just  below  the  slightly  constricted  apex,  sparsely  pilose  with- 
out, densely  so  within  most  of  its  length ;  sepals  acuminate,  subulate  for  1-2 
mm.  at  apex,  somewhat  pilose  on  back,  glabrous  above;  petals  scarlet,  rounded- 
obovate,  10-13  (15)  mm.  long,  about  as  wide;  stamens  13-17  and  10-14  mm. 
long ;  anthers  3  mm.  long ;  style  densely  pilose  throughout  its  portion  within 
the  hypanthium,  sparsely  so  on  base  of  exposed  portion,  about  equalling  the 
sepals;  stigma  2-2.5  mm.  thick;  fruit  apparently  ellipsoid. 

Type  locality,  Ayavaca  (Ayabaca),  Piura,  Peru.  There  is  some  question 
as  to  whether  I  have  seen  the  original  material ;  number  3115  of  Humboldt 
in  the  Willdenow  Herbarium  at  Berlin  labeled  Fuchsia  is  interpreted  by 
Lewin  as  ayavacensis.  A  photograph  was  distributed  by  the  Field  Museum  as 
F.  hirta,  which  name  has  never  been  published.  No.  3115  in  the  Bonpland  Her- 
barium at  Paris  is  labeled  "F.  ayavacensis  HBK,  F.  ampliaia  Benth. !"  as  from 
Kucu  Pichincha.  It  agrees  well  with  the  description,  as  does  tj^pe  material  of 
F.  ampliata.  In  the  type  of  F.  Hitchcockii,  the  petals  are  only  half  the  sepals, 
the  hypanthium  is  rather  long,  but  both  these  conditions  are  matched  by  some 
of  the  Jameson  collections.  The  reflexed-spreading  sepals  and  round  petals 
shorter  than  the  sepals,  the  pilose  condition  of  the  largish  leaves  characterize 
F.  ayavacensis.  Representative  material.  ECUADOR :  without  locality,  Jame- 
son (US).  Pichincha:  Canton  Quito:  road  from  Cotocollao,  near  Nono,  2987 
m.,  Mexia  7661  (POM,  US)  ;  Volcan  de  Pichincha,  3500-4000  m.,  Mexia  7654 
(POM,  US)  ;  La  Chorera,  above  Quito,  Balls  7234  (US) ;  Vi(t\\mc\m,  James  on 
73  (NY)  ;  sive  Andium  Quitensium,  Jameson  42  (GH),  30  (NY)  ;  Andes  of 
Quito,  Jameson  (US) ;  in  declivitate  montis  Pichincha,  3000  m.,  Harttveg  988, 
type  no.  ampliata  at  Berlin  (photo  F,  PO^M)  ;  Rucu  Pichincha,  Bonpland 
3115,  Herb.  Mus.  Paris  (F),  Humholdt  3115  at  Berlin  (photo  F,  POM). 
Azuay :  between  Oiia  and  Cuenca,  Hitchcock  21603,  type  no.  Hitchcockii  (GH, 
NY,  US). 


Vol.  XXV]  MUNZ :  THE  GENVS  FUCHSIA  33 

(22)  Fuchsia  Pringsheimii  Urban 

(Plate  4,  fig.  19) 
Fuchsia  Pringsheimii  Urban,  Symb.  Antill.,  1:375, 1898-1900. 

Shrub  1-2  m.  tall,  openly  branched,  the  ultimate  branchlets  purplish,  1-2 
mm.  thick,  fine-pubescent ;  leaves  opposite,  fairly  near  each  other,  not  much 
reduced  upward,  subcoriaceous,  strongly  bicolored,  ovate-elliptic,  to  rhom- 
boid-elliptic, acute  at  base,  acute  to  acuminate  at  apex,  somewhat  remotely 
serrulate,  in  maturity  mostly  subglabrous  except  on  veins  of  both  surfaces, 
principal  lateral  veins  3-4  on  each  side  of  midrib ;  petioles  4-7  mm.  long ; 
blades  1.5-2.5  cm.  long,  0.5-1  cm.  wide ;  flowers  in  axils  of  upper  leaves ;  pedi- 
cels subglabrous,  2-5  cm.  long;  hypanthium  funnelform,  23-26  mm.  long,  red, 
glabrous  without  and  within,  constricted  above  ovary,  then  with  tubular 
portion  about  2  mm.  wide,  10-15  mm.  long,  the  upper  expanded  portion  10-15 
mm.  long,  8-12  mm.  wide ;  sepals  red,  lanceolate,  acuminate,  glabrous,  18-22  ■ 
mm.  long,  7-8  mm.  wide  at  base ;  petals  red,  equalling  or  slightly  exceeding 
sepals,  obovate ;  episepalous  stamens  almost  as  long  as  petals,  anthers  3  mm. 
long;  epipetalous  slightly  shorter,  the  anthers  2.5  mm.  long;  style  glabrous, 
equalling  petals  or  up  to  5  mm.  longer ;  stigma  clavate,  slightly  4-lobed,  2  mm. 
long ;  berry  ellipsoid,  10-14  mm.  long,  half  as  thick. 

Type  locality,  "Valle  nuevo,"  Santo  Domingo,  at  2100  m.,  the  type  Eggers 
2159.  The  species  differs  from  F.  triplxyUa  by  having  the  flowers  in  axils  of 
reduced  leaves  not  in  an  inflorescence,  leaves  more  glabrous,  and  by  sepals 
and  petals  being  longer  in  relation  to  the  hypanthium.  It  and  F.  iripliylla  and 
ayavacensis  seem  to  be  related  in  their  ampliate  hypanthium  and  wide  petals. 
I  have  seen  the  following  specimens  of  Pringsheimii :  HAITI :  Massif  de  la 
Pelle,  Petionville,  Morne  La  Visite,  at  2100  m.,  Ekman  H  1450  (US) .  SANTO 
DOMINGO  :  prope  Constanza  in  Valle  nuevo,  2300  m.,  von  Tuercklieim  3151 
(F,  GH,  NY,  US) .  A  collection  from  Massif  de  la  Pelle,  Marigot,  near  Jardins 
Bois-Pin,  Haiti,  Ekman  H  1260  (US)  has  leaves  of  triphyUa  and  flowers  of 
Pringsheimii. 

(23)  Fuchsia  triphylla  L. 

(Plate  4,  fig.  20) 
Fuchsia  triphylla  L.,  Sp.  PL,  1191,  1753. 

Apparently  semishrubby,  3-5  dm.  tall;  stems  1  to  few,  simple  or  few- 
branched,  the  yoimger  ones  reddish,  densely  pubescent,  2-A  mm.  thick;  leaves 
ternate  or  quaternate  or  opposite,  the  lower  ones  remote,  middle  and  upper 
crowded,  coriaceous,  lanceolate  to  lance-ovate,  a'cute  to  acuminate  at  both 
ends,  subentire  to  serrulate,  often  with  inrolled  margin,  bicolored,  often  red- 
dish on  veins,  fine-pubescent  on  both  surfaces;  petioles  pubescent,  3-10  (20) 
mm.  long;  main  blades  3-8  (10)  cm.  long,  1-3  (4.5)  cm.  wide;  uppermost 
reduced  to  leafy  lanceolate  or  lance-ovate  bracts;  flowers  several,  pendulous, 
in  dense  terminal  racemes;  pedicels  15-20  (30)  mm.  long,  rather  slender, 
pubescent ;  hypanthium  slightly  contracted  then  expanded,  funnelform,  red, 


34  CALIFOENIA  ACADEMY  OF  SCIENCES  [Proc.  4th  Ser. 

pubescent  without,  glabrous  within,  the  tubular  part  1.5-2  mm.  wide,  8-16 
mm.  long,  the  expanded  part  4-7  mm.  wide,  15-20  mm.  long ;  sepals  red,  lance- 
olate, erect,  acuminate,  8-13  mm.  long,  4-5  mm.  wide  at  base,  puberulent; 
petals  red  at  apex,  apparently  lighter  toward  base,  ovate-oblong,  8-10  mm. 
long ;  stamens  as  long  as  or  two-thirds  as  long  as  petals ;  anthers  2-2.5  mm. 
long;  stj'le  glabrous,  scarcely  equal  to  sepals;  stigma  conic,  not  much  lobed, 
1.5  mm.  long ;  berry  subglobose,  at  least  1  cm.  long. 

Tj-pe  locality,  "Habitat  in  America"  according  to  Linnaeus  w^ho  refers  to 
Plumier,  Gen.,  14,  1703,  where  the  genus  Fuchsia  was  founded  with  a  rude 
drawing  (cf.  PI.  16,  figs.  92-95)  Linnaeus  basing  F.  triphylla  on  Plumier. 
Lamarck,  Encyc.  2 :565,  1786,  said  that  Plumier  had  found  the  species  in  the 
Island  of  Santo  Domingo,  in  uncultivated  places,  in  going  from  the  region 
"de  la  Bande  du  Sud  a  celui  qu'on  nomme  le  grand  Cul-de-sac."  Linnaeus 
placed  the  genus  under  Tetrandria  Monogynia,  but  in  the  Gen.  PI.  ed.  2,  53, 
1764,  observed:  "Plumier  in  Generibus  stamina  quator,  in  Historia  vere 
saepius  octo  pingit."  Presumably  then,  the  species  should  have  been  described 
as  having  eight  stamens  and  coming  from  Santo  Domingo.  Of  the  specimens 
listed  below,  there  seem  to  be  two  forms :  one  with  leaves  heavy,  lanceolate, 
1-2  cm.  wide ;  the  other  less  thick,  2-4.5  cm.  wide.  In  the  first  may  be  cited : 
HAITI :  vicinity  of  Mission,  Fonds  Varettes,  at  1000  m.,  Leonard  3681  (NY, 
US),  3701  (NY,  US).  Belonging  to  the  second  group  are :  HAITI :  Massif  de 
la  Selle,  Petionville,  at  1600  m.,  Ekman  H  1170  (US).  SANTO  DOMINGO  : 
Prov.  de  la  Vega,  ad  ripam  fontis  Yaque  at  1200  m..  Padre  Miguel  Fuertes 
1720  (NY)  ;  prope  Jarabacoa,  700  m.,  Fuertes  1619  (NY) ;  Constanza,  Ablot 
in  1919  (US),  von  Tuerckheim  2956  (F,  GH,  NY,  US),  554i  (NY).  Perhaps 
more  material  may  show  that  these  forms  should  be  separated  taxonomically. 

(24)  Fuchsia  petiolaris  H.B.K. 

Fuchsia  petiolaris  H.B.K.,  Nov.  Gen,  et  Sp.,  6:104,  1823. 

Erect  bushy  shrubs  or  somewhat  scandent,  up  to  2  m.  tall ;  younger  twigs 
reddish,  but  covered  with  a  fine  ashy  appressed  puberulence,  terete,  1-3  mm. 
thick  ;  leaves  elliptical  to  elliptic-ovate,  rather  dense,  membranaceous,  touched 
with  purple,  obtuse  to  acute  at  both  ends,  remotely  serrulate,  with  5-7  main 
veins  on  each  side  of  midrib  and  with  submarginal  vein,  minutely  puberulent 
on  both  surfaces,  somewhat  paler  on  lower  surface  than  upper;  blades  mostly 
1.5-5  (8)  cm.  long,  1-2.5  cm.  wide;  petioles  slender,  5-15  mm.  long;  stipules 
subulate,  1-1.5  mm.  long,  fairly  persistent ;  flowers  in  uppermost  axils,  some- 
times almost  racemose;  pedicels  minutely  strigulose,  slender,  1.5-2.5  cm.  long ; 
ovary  strigulose-puberulent,  3-4  mm.  long;  hypanthium  red,  3-3.5  mm.  wide 
at  base,  then  2-3  mm.  wide  for  length  of  1  cm.,  then  rather  abruptly  ampliate 
until  8-11  mm.  wide  just  below  the  slightly  constricted  summit,  3.5-5  cm. 
long,  strigulose-puberulent  without,  somewhat  pilose  within  lower  portion ; 
sepals  deltoid-lanceolate,  16-18  mm.  long,  6-7  mm.  wide,  acuminate,  more  or 
less  subulate  for  terminal  1-2  mm.,  red,  finely  strigulose-puberulent  on  backs, 


Vol.  XXV]  "  MUNZ:  THE  GENUS  FUCHSIA  35 

glabrous  on  upper  surface;  petals  deep  red,  lance-obloug,  15-17  mm.  long, 
4-5  (8)  mm.  wide,  acute,  frequently  strigulose-puberulent  on  backs;  stamens 
12  and  8  mm.  long ;  anthers  2.5-3  mm.  long;  style  sparsely  pilose  in  lower  por- 
tion, equalling  sepals ;  stigma  2  mm.  wide;  berry  almost  globose  when  pressed, 
8-10  mm.  long. 

This  and  the  species  that  follow  differ  from  the  three  previous  ones  by  their 
narrower  petals,  and  resemble  them  in  the  long,  more  or  less  ampliate  hypan- 
thium  and  the  axillary  or  subterminal  flowers. 

Key  to  Varieties  of  Fuchsia  petiolaris 

A.  Leaf -blades  2-3  times  as  long  as  wide;  petals  4-5  mm.  wide 24a.  var.  typica. 

AA.  Leaf -blades  4-5  times  as  long  as  wide;  petals  7-8  mm.  wide 24b.  var.  bolivarensis, 

(24a)  Fuchsia  petiolaris  H.  B.  K.  var.  typica  Munz,  new  name 

(Plate4,  fig.  21) 

F.  petiolaris  H.  B.  K.,  Nov.  Gen.  et  Sp.,  6:104,  1823. 

F.  quinduensis  H.  B.  K.,  I.e.,  105 ;  I  have  seen  a  portion  of  the  type  collection. 
F.  curviflora  Benth.,  PI.  Hartweg.,  177,  1845 ;  type  from  near  Bogota,  and  not  seen  by  me. 
Mutis  1365  compared  with  type  by  Johnston  seems  to  agree  thoroughly  and  was  seen. 

Leaf -blades,  1.5-5  cm.  long,  one  half  to  one  third  as  wide ;  petals  4r-5  mm. 
wide. 

Type  locality,  Santa  Fe  de  Bogota,  Cundimarca,  Colombia.  Eepresentative 
material,  all  from  COLOMBIA:  without  locality,  Mutis  1365  (US),  1366 
(US),  1360  (US).  Cundimarca:  Bogota,  Dawe  93  (US),  Ariste-Joseph  56 
(US) ;  Paramo  de  Cruz  Verde,  near  Bogota,  Pennell  2056  (NY)  ;  Santa  Fe 
de  Bogota,  Humboldt,  Humboldt  Herb,  at  Berlin  (photo  F,  POM)  and  leaf 
from  Bonpland  Herb,  at  Paris  (F)  ;  Rio  San  Cristobal,  near  Bogota,  Pennell 
2379  (GH,  NY,  US).  Caldas:  sw.  of  Ruiz,  Termales,  3400  m.,  Cuatrecasas 
9218-A  (US)  ;  Quindio  La  Linea,  Dryander  1939  (US).  Caldas  or  Tolima? : 
Quindiu,  Humboldt,  Willdenow  Herb,  at  Berlin,  type  no.  quinduensis  (photo 
F,  POM),  fragment  from  Bonpland  Herb,  at  Paris  (F).  Tolima  :  Volcancito, 
Holton  896  (GH,  NY) ;  Ibague,  at  1300  m.,  Andre  K  817  (F,  GH,NY)  ;Rosa- 
lito  near  Paramo  de  Ruiz,  Pennell  3131  (F,  GH,  NY,  US)  ;  along  divide  near 
Quindio  Highway,  3300-3500  m.  Killip  and  Varela  34600  (POM,  US).  Comi- 
saria  del  Putumayo  :  Corregimiento  El  Encano,  Laguna  de  la  "Cocha,"  Garcia 
Barriga  7836  (US)  ;  Paramo  El  Tabano,  3300  m.,  Garcia  Barriga  7843  (US) ; 
Paramo  de  Faliano,  Garcia  B.  no.  4560  (US). 

(24b)  Fuchsia  petiolaris  H.  B.  K.  var.  bolivarensis  Munz,  new  var. 

(Plate  4,  fig.  22) 

Leaf -blades  lanceolate,  5-7.5  cm.  long,  12-14  mm.  wide ;  petals  obovate,  7-8 
mm.  wide.  (Folia  lanceolata,  5-7.5  cm.  longa,  1.2-1.4  cm.  lata;  petalis  obovatis, 
7-8  mm.  latis). 

Type :  Colombia :  Bolivar :  below  Paramo  de  Chaquiro,  Cordilerra  Occi- 
dental, 2800-3100  m.,  11-24-1918,  F.  W.  Pennell  4324  (NY).  Like  var.  tijpica 
in  puberulence,  flower-size,  etc. 


36  CALIFOBNIA  ACADEMY  OF  SCIENCES  [Proc.  4th  See. 

(25)  Fuchsia  Smithii  Munz,  new  species 
(Plate  4,  fig.  23) 

Low  shrub  to  Avoody  vine  up  to  2-3  m.  tall,  with  drooping  branches,  the 
ultimate  ones  reddish  to  purplish,  more  or  less  ferruginous-villous,  terete, 
1.5-2.5  mm.  thick,  older  ones  with  exfoliating  epidermis;  leaves  mostly  ter- 
nate,  sometimes  opposite,  elliptic  to  elliptic-ovate  or  -obovate,  obtuse  to  acute 
at  both  ends,  or  abruptly  acuminate  at  apex,  plainly  remote-serrulate,  pubes- 
cent and  green  above  or  glabrescent  except  on  veins,  paler  beneath  and  gen- 
erally pilose  or  only  so  along  the  veins,  membranaceous,  with  9-11  principal 
veins  on  each  side  of  midrib  and  with  submarginal  vein ;  leaf-blades  2.5-8  cm. 
long,  1-2.5  (3.5)  cm.  wide;  petioles  slender,  villous-pubescent,  5-12  mm.  long; 
stipules  subulate-lanceolate,  1-1.5  mm.  long,  deciduous;  flowers  in  upper 
axils;  pedicels  rather  slender,  pubescent,  2.5—4  (6)  cm.  long;  ovarj^  oblong, 
pubescent,  6-7  mm.  long,  1.5-2  mm.  thick;  hypanthium  tubular,  pinkish  red 
to  purplish  red,  4-5.5  cm.  long,  3-4  mm.  wide  at  base,  then  narrowed  to  2-3 
mm.  for  1.5-2  cm.,  then  rather  abruptly  widened  to  7-9  mm.  until  near  the 
sometimes  slightly  constricted  apex,  loosely  pubescent  to  almost  glabrous 
without,  pilose  within  most  of  its  length ;  sepals  divergent,  lanceolate,  light 
to  deep  red,  acuminate  with  subulate  tips  ca.  1  mm.  long,  subglabrous  to  some- 
what pilose  on  both  surfaces,  13-16  (20)  mm.  long,  5-6  mm.  wide ;  petals  dark 
red  to  lighter,  narrowly  oblong-ovate,  13-18  mm.  long,  5-6  (8)  mm.  wide, 
acute,  often  erose-margined,  usually  pilose  on  back  along  median  line ;  stamens 
11-13  and  9-10  mm.  long,  red;  anthers  cream,  2-3  mm.  long;  style  pilose  on 
lower  portion,  equaling  petals ;  stigma  2.5  mm.  thick ;  berry  dark,  broadly 
ellipsoid,  12-15  mm.  long,  8-10  mm.  wide. 

Frutex  erectus  vel  scandens,  ad  2-3  m. ;  ramulis  ferrugineo-villosis,  1.5-2.5  mm.  diametro ; 
foliis  tematis  vel  oppositis,  ellipticis,  elliptico-ovatis  vel  -obovatis,  obtusis  vel  acutis  vel 
abrupte  acuminatis,  in  superficie  siiperiore  pubescentibus,  inferiore  pilosis,  membranaceis ; 
lamiiiis  2.5-8  cm.  longis,  1-2.5  (3.5)  cm.  latis;  petiolis  tenuibus,  villoso-pubescentibus,  5-12 
mm.  longis;  floribus  axillarbus;  pedicellis  pubescentibus,  2.5-4  (6)  cm.  longis;  ovario 
oblongo,  pubcscente,  6-7  mm.  longo;  hypantliio  tubuloso,  4-5.5  cm.  longo,  basi  3-4  mm. 
lato,  supra  2-3  mm.  lato,  versus  apicem  7-9  mm.  lato,  exteriore  pubescente  ad  glabro, 
intcriore  piloso ;  scpalis  divergentibus,  lanceolatis,  acuminatis,  13-16  (20)  mm.  longis,  5-6 
mm.  latis,  subglabris  vel  pilosis;  petalis  oblongo-ovatis,  13-18  mm.  longis,  5-6  (8)  mm. 
latis,  acutis,  pilosis;  staminibus  11-13  et  9-10  mm.  longis;  antheris  2-3  mm.  longis;  stylo 
piloso,  petalis  aequalibus;  stigmate  2.5  mm.  lato;  bacca  late  ellipsoidea,  12-15  mm.  longa, 
8-10  mm.  lata. 

Type:  Dept.  Santander:  vicinity  of  Vetas,  Colombia  at  3100-3250  m., 
open  rocky  hillsides,  Jan.,  1927,  E.  P.  Killip  mid  Albert  C.  Smith  17300,  at 
Gray  Herbarium;  istoypes  (F,  NY,  US).  The  proposed  species  is  near  F. 
petiolaris  of  the  western  Cordillera  of  Colombia  in  having  pubescence  on  the 
back  of  the  petals  and  in  many  features,  but  differs  in  the  coarser  longer 
pubescence,  larger  leaves,  longer  pedicels,  larger  fruit  and  eastern  range. 
Other  material  representative,  COLOMBIA  :  Santander  del  Norte  :  west  slope 


Vol.  XXV]  MUNZ:  TEE  GENUS  FUCHSIA  37 

of  Paramo  del  Hatico,  Toledo  to  Pamplona,  2800  m.,  KilJip  and  Smith  20714 
(GH,  NY,  US) ;  between  Mutiscua  and  Pamplona,  3400  m.,  Killip  and  Smith 
19734  (GH,  NY,  US)  ;  east  slope  of  Paramo  de  Santiirban,  toward  Mutiscua, 
3600-3900  m.,  Killip  and  Smith  19599  (GH,  NY,  US)  ;  Pica-Pica  Valley  above 
Tapata  north  of  Toledo,  Killip  and  Smith  21121  (GH,  NY).  Santander: 
Paramo  de  Romeral,  3800-4100  m.,  Killip  and  Smith  18577  (GH,  NY,  US)  ; 
Paramo  de  las  Puentes,  above  La  Baja,  3500-3700  m.,  Killip  and  Smith  18211 
(GH,  NY,  US) ;  Paramo  Rico,  Killip  and  Smith  17771  (GH,  NY,  US),  17755 
(F,  GH,  NY,  US)  ;  vicinity  of  Vetas,  Killip  and  Smith  17361  (GH,  NY,  US), 
17335  (GH,  US),  17386  (GH,  US)  ;  edge  of  Paramo  de  Las  Vegas,  Killip  and 
Smith  15734  (F,  GH,  NY,  US),  15610  (GH),  between  California  and  Vetas, 
Killip  and  Smith  17236  (GH,  NY,  US).  Boyaca:  between  Soata  and  Cocuy, 
Paramo  del  Alto  Caiiutal,  Cuatrecasas  1183  (US)  ;  Valle  de  la  Uvita,  Cuatre- 
casas  1146  (US),  1161  (US). 

(26)  Fuchsia  Asplundii  Macbride 
(Plate  4,  fig.  24) 
Fuchsia  Asplundii  Macbride,  Field  Mus.  Nat.  Hist.,  Bot.  Ser.,  13,  pt.  4,  no.  1:548,  1941. 

Shrub ;  young  twigs  apparently  dark  in  color,  densely  crisp-pubescent,  1-2 
mm.  thick ;  leaves  ternate  or  quaternate,  membranaceous,  broadly  elliptic- 
lanceolate,  acute  to  obtuse  at  base,  acute  to  subacuminate  at  apex,  subentire, 
strigose  above,  more  or  less  pilose  beneath  especially  along  veins,  principal 
lateral  veins  8-12  on  each  side  of  midrib,  the  submarginal  vein  well  developed ; 
blades  3-8  cm.  long,  1-2.5  cm.  wide ;  petioles  appressed-pubescent,  1-1.5  cm. 
long  on  main  cauline  leaves,  4-5  mm.  on  reduced  upper  leaves;  stipules  lance- 
subulate,  1  mm.  long,  strigose,  deciduous ;  flowers  solitary  in  axils  of  some  of 
principal  leaves,  or  appearing  subterminal ;  pedicels  slender,  crisp-pubescent, 
1-2.5  cm.  long;  ovary  linear-fusiform,  crisp-pubescent,  6-8  mm.  long;  hypan- 
thium  4-5  cm.  long,  2  mm.  thick  at  base,  then  somewhat  narrowed  for  one- 
third  its  length,  then  gradually  ampliate  until  5  mm.  wide  at  mouth,  crisp 
pubescent  without,  pilose  within  lower  half ;  sepals  divergent,  12-14  mm.  long, 
3  mm.  wide,  subulate  for  about  1  mm.  at  apex,  crisp-pubescent  on  backs, 
glabrous  on  upper  surface ;  petals  oblong-ovate,  10-12  mm.  long,  5-6  mm. 
wide,  obtusish,  glabrous;  stamens  10-11  and  7-8  mm.  long;  anthers  3  mm. 
long ;  style  pilose  on  basal  half,  exserted  beyond  sepals  4-6  mm. ;  stigma  about 
1  mm.  thick ;  fruit  narrow-ellipsoid,  4-angled,  at  least  12  mm.  long,  and  3  mm. 
wide. 

Type  from  PERU  :  Dept.  Piura :  Prov.  Huancabamba  :  above  Palambla,  at 
3000  m.,  April,  1912,  Welerlauer  6054  (F,  GH,  US).  This  species  suggests 
F.  ayavacensis  in  the  almost  terminal  flowers,  but  the  hypanthium  is  narrower, 
as  are  the  petals.  In  its  narrow  petals  and  pubescent  stems  it  resembles  F. 
petiolaris  and  F.  Smithii,  but  the  pubescence  is  crisped,  the  fruit  is  more 
narrow  and  the  range  more  southern. 


38  CALIFORNIA  ACADEMY  OF  SCIENCES  [Proc.  4th  Ser. 

(27)  Fuchsia  Llewelynii  IMacbride 

(Plate  5,  fig.  25) 
Fuchsia  Llewelynii  Macbride,  Field  Mus.  Nat.  Hist.,  Bot.  Ser.,  13,  part  4,  no.  1 :556,  1941. 

Shrub  1  m.  tall;  young  twigs  finely  and  closely  appressed-puberulent ; 
leaves  opposite,  rigid,  broadly  oblong-oblaneeolate  to  -lanceolate,  acute  at 
both  ends,  plainly  serrulate,  minutely  appressed-puberulent  above  and  be- 
neath especially  on  veins,  apparently  paler  beneath,  with  about  11-14  prin- 
cipal lateral  veins  on  each  side  of  midrib  and  with  submarginal  vein ;  blades 
4-9  cm.  long,  1.5-3  cm.  wide ;  petioles  4—8  mm.  long;  stipules  subulate,  2  mm. 
long,  fairly  persistent;  flowers  apparently  racemose;  pedicels  subfiliform, 
strigulose,  3-5  cm.  long;  ovary  fusiform,  strigulose,  5-6  mm.  long;  hypan- 
thium  about  5  cm.  long,  2  mm.  wide  at  base,  then  about  1  mm.  wide  for  1.5  cm., 
then  rather  gradually  ampliate  until  5  mm.  wide  at  summit,  strigulose  with- 
out, pilose  within  the  lower  half ;  sepals  linear-lanceolate,  divergent,  14--17  mm. 
long,  4  mm.  wide  at  base,  subulate  for  1-2  mm.  at  apex,  strigulose  without, 
glabrous  within ;  petals  oblong-lanceolate,  15-16  mm.  long,  4^5  mm.  wide, 
acuminate;  stamens  12-13  and  10-11  mm.  long;  anthers  2-2.5  mm.  long; 
style  densely  pilose  on  basal  portion,  exceeding  sepals  by  few  mm. ;  stigma  1.5 
mm.  thick. 

Tj^e  from  PERU :  Dept.  Amazonas :  La  Ventana  Road,  Chachapoyas  to 
Moyobamba,  Llewelyn  Williams  7594  (F).  The  type  specimen  is-much  broken 
and  rather  unsatisfactory ;  apparently  from  a  plant  with  long  slender  racemes. 
It  is  near  F.  petiolaris  in  the  puberulence  and  long  pedicels,  but  the  leaves 
seem  larger  and  with  more  veins,  and  the  hypanthium  is  more  narrow.  It  is 
near  F.  Asplundii  in  the  narrow  hypanthium  but  has  much  finer  puberulence. 

(28)  Fuchsia  venusta  H.B.K. 

Fuchsia  venusta  H.  B.  K.,  Nov.  Gen.  et  Sp.,  6:105,  1823. 

Shrub  or  shrubby  vine,  the  young  twigs  hirtellous  to  puberulent  or  sub- 
glabrous,  often  dark  red,  2-6  mm,  thick ;  leaves  opposite  or  ternate,  elliptic, 
rounded  to  acute  at  base,  acute  to  acuminate  at  apex,  subentire,  subcoriaceous, 
not  much  paler  beneath  that  above,  subglabrous  to  puberulent  or  pubescent 
on  both  sides,  especially  on  veins,  principal  lateral  veins  mostly  10-15  on  each 
side  of  midrib,  with  evident  submarginal  vein;  blades  4.5-8  (10.5)  cm.  long, 
2-3.5  (4)  cm.  wide ;  petioles  6-10  mm.  long,  puberulent  to  pubescent ;  stipules 
triangular-subulate,  barely  1  mm.  long,  deciduous ;  flowers  in  a  terminal 
pendant  corymbose  raceme  or  in  panicles  scarcely  1  dm.  long;  bracts  elliptic- 
lanceolate,  1-2  cm.  long ;  pedicels  1-3  cm.  long,  almost  filiform,  glabrous  to 
pubescent ;  ovary  oblong,  6-7  mm.  long,  ca.  2  mm.  thick ;  hypanthium  tubular, 
red,  subglabrous  to  glandular-pubescent  without,  villous  within  lower  portion, 
3-4  em.  long,  2.5-3  mm.  wide  at  base,  then  about  2  mm.  wide  for  one-third  its 
length  and  gradually  widened  until  6-7  mm.  broad  at  summit;  sepals  lance- 
olate, acuminate,  the  connivent  tips  subulate  for  3-4  mm.,  sepals  red,  glabrous 


Vol.  XXV]  MVNZ :  THE  GENUS  FUCHSIA  39 

to  pubescent  on  backs,  pubescent  within,  more  or  less  spreading,  15-17  (20) 
mm.  long,  3-4  mm.  wide ;  petals  carmine,  oblong-lanceolate,  abruptly  acute, 
more  or  less  pubescent  on  backs,  15-18  (20)  mm.  long;  stamens  12-13  (15) 
and  9-10  (12)  mm.  long,  glabrous;  anthers  2.5-3  mm.  long;  style  slender, 
densely  villous  through  much  of  its  length,  extending  to  about  tips  of  sepals ; 
fruit  oblong,  up  to  14  mm.  long  and  7  mm.  wide. 

This  species  is  characterized  by  its  short  elliptic  coriaceous  leaves,  terminal 
inflorescence,  short  petioles,  long  pedicels ;  sepals  hairy  on  inner  surface  as 
well  as  outer,  petals  hairy  on  backs.  In  the  more  or  less  ampliate  hypanthium 
and  subterminal  flowers  it  agrees  with  those  immediately  preceding  and 
following  in  this  treatment.  There  are  two  forms  as  to  pubescence  which  I  am 
recognizing  as  varieties. 

Key  to  Varieties  of  Fuchsia  venusta 
A.  Young  twigs,  petioles,  hypanthia  pubescent  with  short  spreading  hairs.  28a.  var.  typica. 
AA,  Young  twigs,  petioles,  hypanthia  glabrous  to  minutely  puberulent  with  more  or  less 
appressed  hairs 28b.  var.  huilensis. 

(28a)  Fuchsia  venusta  H.  B.  K.  var.  typica  Munz,  new  variety 

(Plate  5,  fig.  26) 
F.  venusta  H.  B.  K.,  Nov.  Gen.  et  Sp.,  6:105,  1823. 

Hairs  on  young  growth  and  flowers  spreading. 

Type  locality:  near  Guayavalito,  Colombia.  Ranging  in  the  eastern 
Cordillera  of  Colombia  in  Santander  and  Cundimarca.  Representative  mate- 
rial, COLOMBIA:  Guayavalito  (exact  locality  not  determined),  Humholdt, 
ex  Willdenow  Herb.  7301  (photo  F,  POM),  Bonpland  Herb,  ex  Mus.  Paris 
(fragment  F ) .  Without  locality,  Mutis  1370, 1362, 1367,  and  1369  (all  at  US) ; 
eastern  Cordillera,  Bro.  Ariste-Joseph  A893  (US).  Santander:  Chiquinquira, 
Ariste-Joseph  A  842  (US).  Cundimarca:  Tena,  Dawe  23  (US);  Bogota, 
Schultze  69  (JJ^),  Ariste-Joseph  A229  (US)  ;  Salto  de  Tequendama,  2500  m., 
Cuatrecasas  48  (US)  ;  Cordillere  de  Bogota,  1800-2600,  Triana  3814  (NY, 
US) ;  ad  cataractam  Tequedamam,  Holt  on  894  (GH,  NY)  ;  Paramo  de  San 
Fortunato  et  prope  pagum  Fusagasuga,  Hartweg  995  (NY). 

(28b)  Fuchsia  venusta  H.  B.  K.  var.  huilensis  Munz,  new  variety 

Young  growth,  hypanthia,  etc.,  glabrous  to  minutely  appressed  puberulent. 
(Ramula,  petiola,  et  hypanthia  glabra  vel  minute  striguloso-puberulenta) . 

Type:  from  Huila,  Colombia:  Cordillera  Oriental,  east  of  Neiva,  forest  at 
1800-2300  m.,  Rushy  and  Pennell  874,  (NY)  ;  isotypes  (F,  GH,  US).  Mate- 
rial to  be  referred  here  comes  from  two  widely  separate  regions,  VENE- 
ZUELA: Merida:  Paramo  de  Aricagua,  2700  m.,  Jalin  1029  (GH,  US). 
Tachira:  Paramo  de  Angaraveca,  2400  m.,  J  aim  129  (US).  COLOMBIA: 
Cundimarca:  Dintel  (Facatativa-La  Vega)  2700-2300  m.,  Perez  A.  and 
Cuatrecasas  5291  (US) ;  Une,  in  montibus  ad  orient.  Bogotae,  Holton  895 
(NY).  Tolima :  Libano,  1900-2200  m.,  Pennell  3197  (GH,  NY,  US).  Huila: 


40  CALIFORNIA  ACADEMY  OF  SCIENCES  [Proc.  4th  Ser. 

Cordillera  Oriental,  east  of  Neiva,  Rushy  and  Pennell  641  (GH,  NY,  US)  ; 
"Balsillas"  on  Rio  Balsillas,  2000-2300  m..  Rushy  and  Pennell  918  (NY)  ; 
722  (GH,  NY,  US),  809  (NY,  US). 

(29)  Fuchsia  Jahnii  ]Munz,  new  species 
(Plate  5,  fig.  27) 

Apparently  a  scandent  shrub  with  yoiinger  twigs  very  slender,  1-2  mm. 
thick,  drooping,  somewhat  purplish,  puberulent  with  appressed  hair ;  leaves 
opposite  or  ternate,  elliptic,  rounded  to  acute  at  base,  short-acuminate  at 
apex,  remotely  denticulate,  minutely  pubescent  on  both  surfaces,  darker  green 
above  than  beneath,  membranaceous,  with  about  10-12  principal  veins  on 
each  side  of  the  midrib  and  with  submarginal  vein;  blades  3-8  cm.  long,  1-3 
cm.  wide ;  petioles  6-15  mm.  long,  more  or  less  strigulose ;  stipules  lance- 
subulate,  ca.  2  mm.  long,  deciduous;  flowers  few  in  terminal  corymbose  ra- 
ceme, sometimes  also  axillary;  bracts  1-2  (3)  cm.  long;  pedicels  filiform, 
strigulose,  2-4  cm.  long;  ovary  broadly  fusiform,  strigose,  5-6  mm.  long; 
hypanthium  red,  tubular,  3.5-4  cm.  long,  strigulose  without,  villous  within, 
2.5-3  mm.  wide  at  base,  then  1.5-2  mm.  thick  and  very  gradually  widened 
until  7-8  mm.  wide  toward  the  mouth ;  sepals  red,  lanceolate,  spreading,  18-20 
mm.  long,  3.5-4  mm.  wide,  acuminate  (the  connivent  tips  subulate  for  3^  mm. 
in  bud),  strigulose  without,  glabrous  within;  petals  light  red,  oblong-oblance- 
olate,  18-20  mm.  long,  abruptly  acute,  somewhat  strigulose  on  back  toward 
base;  stamens  16-18  and  12-14  mm.  long,  glabrous;  anthers  2.5-3  mm.  long; 
style  slender,  retrorse-\'illous,  projecting  about  as  far  as  petals;  stigma  sub- 
globose,  slightly  lobed,  2  mm.  wide ;  berry  almost  globose,  strigulose,  about 
12  mm.  long,  10-11  mm.  wide. 

Frutex  scandens;  ramulis  subpurpureis,  strigulosis ;  foliis  oppositis  vel  ternatis,  ellip- 
ticis,  basi  rotundatis  vel  acutis,  apice  breviter  acuminatis,  remote  denticulatis,  minute 
pubescentibus,  membranaceis,  laminis  3-8  cm.  longis,  1-3  cm.  latis;  petiolis  6-15  mm. 
longis,  snbstrigulosis;  floribus  paucis,  subracemosis;  bracteis  1-2  (3)  cm.  longis;  pedicellis 
filif  ormibus,  strigulosis,  2-4  cm.  longis ;  ovario  late  f  usif  ormi,  strigoso,  5-6  mm.  longo ; 
liypantliio  rubro,  tubuloso,  3.5-4  cm.  longo,  externe  striguloso,  interne  villoso,  base  2.5-3 
mm.  late,  inde  1.5—2  mm.  lato,  inde  gradatim  ampliato,  usque  ad  apicem  7-8  mm.  lato; 
sepalis  rubris,  lanceolatis,  divergentibus,  18-20  mm.  longis,  3.5-4  mm.  latis,  acuminatis, 
strigulosis,  cum  apicibus  subulatis  3-4  mm.  longis ;  petalis  roseis,  oblongo-oblanceolatis, 
18-20  mm.  longis,  abrupte  acutis,  basi  et  extrinsecus  substrigulosis;  staminibus  16-18  et 
12-14  mm.  longis;  antheris  2.5-3  mm  .longis;  stylo  tenui,  retrorsovilloso,  petalis  aequalibus 
aequescente;  stigmate  subgloboso,  2  mm.  lato;  bacca  subglobosa,  strigulosa,  11-12  mm. 
longa. 

Type:  Merida,  Venezuela,  Paramo  de  la  Sal,  2800  m.,  Sept.  2,  1921, 
John  506  U.  S.  Nat.  Herb.  1186509 ;  isotype  Gray  Herb.  Another  collection, 
INIerida:  Paramo  de  Pinango,  2600  m.,  Jakn  402  (US).  Apparently  closely 
related  to  F.  venusta  and  characterized  by  a  few  flowers,  strigulose  condition, 
spreading  sepals  with  long  subulate  tips,  hair  on  back  of  petals,  and  big  round 
fruit. 


Vol.  XXV]  MUNZ:  THE  GENUS  FUCHSIA  41 

(30)  Fuchsia  Gehrigeri  Munz,  new  species 

(Plate  5,  fig.  28) 

Subscandent  shrub  3-5  m.  tall,  with  slender,  ultimate,  reddish,  drooping 
branchlets  1-2  mm.  thick,  subglabrous  or  somewhat  villous  about  the  nodes 
and  toward  the  inflorescence,  somewhat  reddish ;  leaves  opposite  or  ternate, 
membranaceous,  elliptic,  acute  to  subacuminate  at  both  ends,  serrulate,  green 
and  minutely  strigulose  above,  pale  beneath  and  pubescent  along  the  veins, 
principal  lateral  veins  7-10  on  each  side  of  midrib,  submarginal  vein  scarcely 
evident;  blades  3-6  cm.  long,  1.5-2.5  (3)  cm.  wide  ;  petioles  slender,  10-18  mm. 
long,  subglabrous  to  pubescent ;  stipules  subulate,  1  mm.  long,  deciduous ; 
flowers  few,  in  terminal  clusters  of  about  2  whorls,  pendant,  dark  red ;  bracts 
1—1.5  cm.  long;  pedicels  slender,  1.5—2  cm.  long,  more  or  less  spreading-villous ; 
ovary  broadly  fusiform,  5  mm.  long,  densely  spreading-villous ;  hypanthium 
tubular,  4—5  cm.  long,  2  mm.  wide  at  base,  then  narrower  for  one-third  its 
length,  and  rather  abruptly  widened  until  7-8  mm.  wide  at  summit,  villous 
without  and  within ;  sepals  lanceolate,  divergent,  14-15  mm.  long,  4  mm.  wide, 
acuminate  (subulate  tips  connivent  and  1  mm.  long  in  bud),  pubescent  on 
back,  glabrous  on  inner  surface  ;  petals  dark  red,  elliptic-oblong,  obtuse-erose, 
ca.  14  mm.  long  and  6  mm.  wide,  sparsely  villous  on  back  near  the  base; 
stamens  9-10  and  7-8  mm.  long ;  anthers  2  mm.  long ;  style  pubescent  on  lower 
portion,  exceeding  sepals  by  3-5  mm.,  quite  thick  toward  the  somewhat  lobed 
stigma  (1.5-2  mm.  wide)  ;  fruit  subglobose,  somewhat  villous. 

Frutex  subscandens,  3-5  m.  altus;  ramulis  tenuibus,  subrubris,  1-2  mm.  diametro,  sub- 
g]abris  vel  subvillosis  in  nodis  et  ad  inflorescentiam ;  f  oliis  oppositis  vel  ternatis,  membran- 
aceis,  ellipticis,  basi  et  apice  acutis  vel  subacuminatis,  serrulatis,  in  superficie  superiore 
viridibus  et  minute  strigulosis,  inf eriore  pallidis  et  in  venis  pubescentibus ;  laminis  3-6  cm. 
longis,  1.5-2.5  (3)  cm.  latis;  petiolis  tenuibus,  10-18  mm.  longis,  subglabris  vel  pubescent- 
ibus; floribus  paucis,  in  infloreseentia  terminali,  subrubris;  bracteis  1-1.5  cm.  longis; 
pedicellis  tenuibus,  1.5-2  cm.  longis,  subvillosis ;  ovario  late  fusif ormi,  5  mm.  longo,  villoso ; 
hypanthio  tubuloso,  4-5  cm.  longo,  basi  2  mm.  lato,  inde  angustiore,  inde  abrupte  ampliato, 
ad  apicem  7-8  mm.  lato,  externe  et  interne  villoso;  sepalis  lanceolatis,  divergentibus,  14-15 
mm.  longis,  4  mm.  latis,  acuminatis,  externe  pubescentibus,  apicibus  subulatis  1  mm. 
longis;  petalis  elliptico-oblongis,  obtuso-erosis,  14  mm,  longis,  6  mm.  latis,  sparse  externe 
villosis ;  staminibus  9-10  et  7-8  mm.  longis ;  antheris  2  mm.  longis ;  stylo  pubescenti,  quam 
sepala  3-5  mm.  longiore;  stigmate  1.5-2  mm.  lato;  fructu  subgloboso,  subvilloso. 

Type:  from  Merida:  Mucuruba,  Venezuela,  2800-3100  m.,  July  14,  1930, 
W.  Gehriger  322,  U.  S.  Nat.  Herb.  1515306;  isotypes  F,  GH,  NY.  This  species 
is  near  F.  JaJinii  in  its  leaves,  few  flowers,  round  fruit,  and  petals  with  hair  on 
the  back,  but  the  pubescence  is  longer  and  more  spreading,  the  pedicels  are 
shorter ;  sepal-tips  shorter ;  flowers  smaller,  and  sepals  are  less  spreading. 

(31)  Fuchsia  simplicicaulis  Ruiz  and  Pa  von 

(Plate5,  fig.  29) 
Fuchsia  simplicicaulis  Euiz  and  Pav6n,  F1.  Peruv.,  3  :89,  pi.  322,  1802. 

Shrub,  apparently  growing  up  through  other  shrubs,  up  to  4.5  m.  tall ; 
stems  slender,  the  ultimate  branchlets  pendant,  scarcely  1  mm.  thick,  glabrous 


42  CALIFORNIA  ACADEMY  OF  SCIENCES  [Pboc.  4th  Ser. 

except  for  the  piiberulent  inflorescence ;  leaves  in  remote  whorls  of  4,  linear- 
lanceolate,  coriaceous,  glabrous,  subentire,  obtuse  to  acute  at  base,  acuminate 
at  apex,  not  strongly  bieolorecl,  with  20  or  more  principal  lateral  veins  on 
each  side  of  midrib,  and  with  submarginal  vein,  8-15  cm.  long,  1.5-2  (2.5)  cm. 
wide;  petioles  2-4  mm.  long;  stipules  subulate,  ca.  1  mm.  long,  deciduous; 
inflorescence  a  simple  or  few-branched  hanging  puberulent  raceme  2-3  dm. 
long,  with  remote  whorls  of  4  submembranaceous  lance-ovate,  acuminate, 
green  bracts  2-4.5  cm.  long  and  6-14  mm.  wide ;  flowers  one  in  each  axil,  bright 
red  with  slightly  purplish  tinge ;  pedicels  4-7  mm.  long,  12  mm.  in  fruit ;  ovary 
puberulent,  ellipsoid,  5-6  mm.  long,  1.5-2  mm.  thick;  hypanthium  tubular, 
4-5.5  cm.  long,  puberulent  without  and  within,  ca.  3  mm.  wide  at  very  base, 
then  1.5  mm.  for  two-fifths  its  length,  then  rather  gradually  ampliate  and 
tubular,  5-6  mm.  wide  at  summit;  sepals  4,  linear-lanceolate,  puberulent, 
acuminate,  16-18  mm.  long,  4  mm.  wide,  somewhat  divergent;  petals  red, 
linear -lanceolate,  acute,  11-12  mm.  long,  2-2.5  mm.  wide;  stamens  ca.  14  and 
10  mm.  long;  anthers  3  mm.  long;  style  pubescent  in  lower  hypanthium, 
slightly  exceeding  petals ;  fruit  oblong,  puberulent,  at  least  8  mm.  long. 

Type  locality,  Muna,  Peru.  Specimen  seen,  PERU :  Huanuco :  Muiia,  at 
2200  m..  May  23-June  4,  1923,  J.  F.  Macbride  4014  (F,  GH,  US)  ;  photo  of 
Ruiz  and  Pavon  specimen  from  Muiia  at  Madrid  (F,  POM).  This  species 
stands  apart  from  all  others  in  its  almost  linear  leaves  and  bracts  in  wliorls 
of  4,  in  its  long  pendant  twigs,  and  in  its  puberulence.  It  agrees  with  F.  petio- 
laris  and  F.  ve^iusta  in  its  subterminal  inflorescence,  slightly  inflated  hypan- 
thium and  narrow  petals. 

(32)  Fuchsia  confertifolia  Fielding  and  Gardner 

(Plate6,  fig.  30) 

Fuchsia  confertifolia  Fielding  and  Gardner,  Sert.  PI.,  pi.  28, 1844. 

F.  dolichantha  Krause,  Fedde,  Eep.,  1:172,  1905;  based  on  Weberbauer  4390  from  the 

Chachapoyas  region  in  which  Mathews  collected,  confertifolia  being  based  on  a  Mathews 

type. 

Erect  shrub  up  to  2  m.  tall,  much  branched,  the  young  twigs  1.5-3  mm. 
thick,  densely  f errugineous-hirsute ;  leaves  opposite  to  ternate  or  quaternate, 
crowded,  spreading-reflexed,  oblong-ovate,  rounded  to  acute  at  base,  acute  at 
apex,  entire  to  sub  denticulate,  glabrous  on  both  surfaces  or  sparsely  hairy 
beneath  on  veins,  principal  lateral  veins  obscure,  3-4  on  each  side  of  midrib  ; 
blades  12-16  mm.  long,  3-4  (6)  mm.  wide;  petioles  1-2  mm.  long;  stipules 
dark,  glabrous,  subulate,  1-1.5  mm.  long,  persistent;  flowers  very  few,  sub- 
terminal,  pendant ;  bracts  about  5  mm.  long,  1.5  mm.  wide ;  pedicels  1-1.5  cm. 
long,  up  to  2  cm.  in  fruit,  villous;  ovary  oblong,  4—5  mm.  long;  hypanthium 
dark  red,  tubular,  4r-5  cm.  long,  slightly  enlarged  at  very  base  and  1.5-2  mm. 
wide,  then  narrowed  for  one-third  its  length,  then  abruptly  widened  and  sub- 
cylindric  until  6-8  mm.  wide  at  and  below  the  summit,  sparsely  villous  with- 
out, more  densely  so  within;  sepals  linear-lanceolate,  divergent,  16-18  mm. 


Vol.  XXV]  MUNZ :  TEE  GENUS  FUCHSIA  43 

long,  acuminate,  scarcely  subulate  at  tips,  sparsely  villous  without,  glabrous 
within ;  petals  red,  elliptic-lanceolate,  acuminate,  glabrous,  12-13  mm.  long, 
3  mm.  wide;  stamens  about  13  and  9  mm.  long;  anthers  2.5  mm.  long;  style 
pilose  near  base,  equal  to  sepals;  stigma  capitate,  slightly  lobed,  2  mm.  wide; 
fruit  ellipsoid,  ca.  1  cm.  long. 

Type  locality,  "Peru."  Specimens  seen,  PERU:  Dept.  Amazonas:  between 
Jumbilla  and  San  Carlos,  3000  m.,  WeherUuer  7153  (F,  GH)  ;  ab  Chacha- 
poyas  at  orientem  versus  Tambo  Ventillas,  Weberhauer  4390,  type  doUchantha 
at  Berlin  (photo  F,  POM). 

(33)  Fuchsia  AspiazuiMacbride 
(Plate  6,  fig.  31) 
Fuchsia  Aspiazui  Macbeide,  Field  Mus.  Nat.  Hist.,  Bot.  Ser.,  19,  part  4,  no.  1  -.547, 1941. 

Shrub  up  to  2  m.  tall,  the  younger  branches  4-6  mm.  thick,  minutely  strigu- 
lose  under  a  lens,  greenish,  terete ;  leaves  opposite,  elliptic-ovate  to  -obovate, 
subentire,  apparently  rather  tleshy,  glabrous  on  casual  inspection,  but  min- 
utely strigulose  under  a  lens,  cuneate  at  base,  rather  abruptly  acuminate  at 
apex,  yellowish  green  and  much  alike  on  both  surfaces,  conspicuously  veined, 
the  main  laterals  18-24  on  each  side  of  the  midrib,  with  evident  submarginal 
vein;  blades  8-20  cm.  long,  4-10  cm.  wide ;  petioles  1-2  (3.5)  cm.  long ;  stipules 
lance-deltoid,  2-3  mm.  long,  sometimes  fused  in  pairs,  deciduous ;  inflorescence 
a  terminal  strigulose  pendant  raceme  with  rather  numerous  flowers ;  bracts 
reflexed,  lanceolate,  1-2  cm.  long;  pedicels  8-12  mm.  long;  ovary  linear,  5-8 
mm.  long,  1  mm.  thick,  glabrous  to  somewhat  strigulose ;  hypanthium  blood- 
red,  tubular,  3-5  cm.  long,  slightly  bulbous  at  base,  then  about  1  mm.  wide 
and  very  gradually  ampliate  toward  the  summit  where  it  is  4-5  mm.  wide, 
practically  glabrous  without,  retrorse-pubescent  within  basal  part;  sepals 
divergent,  red,  lanceolate,  12-15  mm.  long,  3-4  mm.  wide,  acute,  without  free 
tips  in  bud;  petals  oblong,  obtuse,  about  1  cm.  long,  3.5  mm.  wide;  stamens 
about  8  and  6  mm.  long;  anthers  barely  2  mm.  long;  style  quite  glabrous, 
extending  about  to  the  end  of  the  petals;  immature  berry  linear-oblong,  12 
mm.  long,  3.5  mm.  wide. 

Based  on  a  collection  from  valley  of  the  Rio  Mixiollo,  Prov.  Pataz,  Dept. 
Libertad,  PERU,  2300-2500  m.,  Weherhauer  7042  (F,  GH,  US).  Character- 
ized by  its  glabrous  condition,  large  leaves  with  many  veins,  short  pedicels,  and 
glabrous  style,  it  comes  in  the  group  of  species  with  large  leaves,  short  terminal 
inflorescence,  and  narrow  long  hypanthium. 

(34)  Fuchsia  tincta  Johnston 
(Plate  6,  fig.  32) 
Fuchsia  tincta  Johnston,  Journ.  Am.  Arb.,  20:242, 1939. 

Shrub,  1-1.5  m.  tall,  the  young  twigs  more  or  less  broAvnish  red,  1-5  mm. 
thick^,  puberulent  or  finely  pubescent  with  somewhat  brownish  hairs;  leaves 
opposite,  membranaceous,  elliptic-ovate  to  -obovate,  rounded  to  obtuse  at  base, 


44  CALIFORNIA  ACADEMY  OF  SCIENCES  [Proc.  4th  Ser. 

acute  to  subaeuminate  at  apex,  remotely  serrulate,  more  or  less  purplish  espe- 
cially beneath,  finely  pubescent  on  both  sides,  even  somewhat  short-pilose  on 
veins  beneath,  the  principal  lateral  veins  16-18  on  each  side  of  midrib,  the 
submarginal  vein  present;  blades  5-12  (15)  cm.  long,  3-8  (10)  cm.  wide; 
petioles  1-4  (6)  cm.  long;  stipules  subulate,  1  mm.  long,  deciduous;  flowers 
few  to  several  in  open  terminal  finely  pubescent  corymbose  racemes;  bracts 
rather  leafy,  1.5-3  cm.  long ;  pedicels  slender,  pubescent,  1.5-3  cm.  long;  ovary 
linear-fusiform,  pubescent,  4r-5  mm.  long ;  hypanthium  20-25  (35)  mm.  long, 
deep  crimson,  1.5-2  mm.  wide  at  base,  then  narrowed  for  one-third  its  length, 
then  gradually  widened  to  3^  mm.  at  summit,  rather  sparsely  villous  with- 
out, villous  within  lower  two-thirds ;  sepals  erimspn,  divergent,  8-10  mm.  long, 
3.5  mm.  wide  at  base,  subulate  for  0.5-1  mm.  at  apex,  villous  on  backs;  petals 
crimson-scarlet,  oblong-ovate,  7-8  mm.  long,  4-5  mm.  wide,  erose-rounded  to 
obtusely  mucronate  at  apex ;  stamens  about  8  and  6  mm.  long ;  anthers  1.5  miu. 
long;  style  pilose  in  basal  portion,  equaling  sepals;  stigma  1-1.5  mm.  wide; 
berry  ellipsoid-oblong,  8-12  mm.  long,  4.5-6  mm.  wide. 

Type  locality,  PERU :  dept.  Cuzco,  prov.  Paucartambo :  Rio  Tambomayo 
at  1800-2200  m.,  the  type  collection  being  West  7092  (GH) ,  another  collection 
from  the  same  locality:  West  7110  (Gil).  I  have  seen  also  from  Cabecera  de 
montana  de  Paucartambo,  at  2000  m.,  Vargas  73/7110  (F,  POM)  and  "Pilla- 
huata,"  Cerro  de  Cusilluyoc,  2000-2400  m.,  Perinell  13956  in  large  part  (F, 
NY,  US).  The  species  is  near  F.  ovalis  in  its  wide  leaves,  but  the  racemes  are 
terminal,  the  pubescence  shorter,  the  stipules  shorter  and  the  hypanthium 
longer.  When  the  hair  is  not  too  brown,  it  suggests  F.  holiviana,  but  the  sepals 
are  not  reflexed-spreading.  In  its  long-tubed  flowers  it  resembles  F.  furfur- 
acea,  but  the  sepals  are  shorter  and  the  plant  is  less  pilose. 

(35)  Fuchsia  Mathewsii  Macbride 

Fuchsia  Mathewsii  Macbride,  Candollea,  8:24,  1940;  Field  Mus.  Nat.  Hist.,  Bot.  Ser.,  13, 
pt.  4,  no.  1:558,  1941. 

Branches  and  petioles  and  lower  leaf-surface  densely  rusty-pilose ;  leaves 
ternate,  broadly  elliptic,  abruptly  narrowed  at  base,  acute  at  apex,  obscurely 
denticulate  to  subentire,  minutely  pilose  above,  densely  so  beneath  along 
midrib;  blades  1  dm.  long;  petioles  4-5  mm.  long;  inflorescence  a  terminal 
short-bracted  panicle  about  5  cm.  long;  hypanthium  4  cm.  long,  glabrate 
without,  hirsutulose  within  lower  half;  sepals  10  mm.  long;  petals  oblong- 
obovate,  almost  as  long  as  sepals. 

Based  on  a  collection  by  Mathews,  from  Chachapoyas,  Amazonas,  Peru. 
Said  to  be  near  F.  pilosa,  but  differing  in  its  short-petioled  leaves,  denser 
shorter  rusty  pubescence  and  its  larger  flowers.  I  have  seen  no  specimens.  The 
meager  description  suggests  F.  tincta  of  a  more  southern  distribution  and 
with  apparently  longer  petioles  and  shorter  hypanthium. 


Vol.  XXV]  MUNZ :  THE  GENUS  FUCHSIA  45 

(36)  Fuchsia  Fischeri  Macbride 

Fuchsia  Fischer i  Macbride,  Field  Mus.  Nat.  Hist.,  Bot.  Ser.,  13,  part  4,no.  1 :554,  1941. 

Shrub,  suberect,  branched,  the  young  twigs  reddish,  densely  hirsutulous ; 
leaves  opposite,  oblong-lanceolate,  subcuneate  at  base,  acute  at  apex,  subentire 
to  obscurely  callose-denticulate,  densely  soft-pilose  on  both  sides,  veins  prom- 
inent beneath ;  petioles  2-5  mm.  long;  stipules  subulate,  somewhat  persistent ; 
inflorescence  a  terminal  leafy -bracted  corymb  5-6  cm.  long ;  pedicels  1.5  cm. 
long;  ovary  velutinous,  4  mm.  long;  hypanthium  about  4.5  cm.  long,  swollen 
at  base,  then  constricted,  then  ampliate  until  4-5  mm.  wide  at  apex,  somewhat 
villous  without  and  within;  sepals  lanceolate,  acuminate,  10-12  mm.  long; 
petals  narrowly  oblong-lanceolate,  short-aciiminate,  about  9  mm.  long;  sta- 
mens shorter  than  petals;  style  pubescent  in  lower  portion,  more  or  less 
exserted. 

Type  not  seen,  based  on  Welerhauer  4097  from  Hualgayoc,  Chugar,  Caja- 
marca,  Peru.  No  specimens  available  and  description  meager,  but  this  seems  to 
be  a  distinct  entity  near  F.  Mathewsii,  perhaps  with  more  narrow  leaves. 

(37)  Fuchsia  Storkii  Munz,  new  species 

(Plate6,  fig.  33) 

Tree,  or  shrub  up  to  3  m.  tall ;  twigs  reddish,  pubescent  or  pilose  with  red- 
dish hairs ;  leaves  ternate,  narrowly  elliptic  to  elliptic-oblanceolate,  firm, 
obtuse  to  acute  at  base,  acute  at  apex,  remotely  serrulate,  light  green  above, 
pilose  to  subgiabrous  or  pubescent  on  veins  above,  slightly  paler  and  more 
pilose  beneath,  principal  lateral  veins  8-9  on  each  side  of  midrib,  submarginal 
veins  present;  leaf -blades  2-7  cm.  long,  1.3-2  cm.  wide;  petioles  reddish- 
pilose,  4-8  mm.  long;  stipules  lance-subulate,  1.5-2  mm.  long,  glabrous,  fairly 
persistent ;  flowers  few,  in  short  terminal  drooping  glandular-pubescent 
corymbose  racemes;  bracts  lanceolate;  pedicels  stout,  red-pubescent,  2-5  mm. 
long ;  ovary  ellipsoid,  glandular-pubescent  to  red-pilose,  4-5  mm.  long ;  hypan- 
thium dark  red,  about  3  cm.  long,  somewhat  bulbous  and  1.5-2  mm.  wide  at 
base,  then  somewhat  narrowed  and  then  gradually  ampliate  until  5  mm.  wide 
at  apex,  glandular-villous  without,  villous  within  lower  half;  sepals  deep  red, 
lanceolate,  about  12  mm.  long,  4  mm.  wide,  subulate  for  1.5-2  mm.  at  apex, 
glandular-pubescent ;  petals  deep  red,  oblong-lanceolate,  7-9  mm.  long,  3  mm. 
wide,  sharply  acute;  stamens  about  8  and  6  mm.  long;  anthers  2  mm.  long; 
style  pubescent  in  lower  portion,  extending  about  to  sepal-tips ;  fruit  appar- 
ently ellipsoid,  at  least  1  cm.  long. 

Arbor  vel  frutex;  ramulis  rubescentibus,  pubescentibus  vel  pilosis,  cum  pubescentia 
rubida ;  f  oliis  ternatis,  anguste  ellipticis  vel  elliptico-oblanceolatis,  base  obtusis  vel  acutis, 
apice  acutis,  remote  serrulatis,  in  venis  superficiei  superioris  pilosis  vel  pubescentibus  vel 
subglabris,  inferiore  pilosioribus;  laminis  2-7  cm.  longis,  1.3-2  cm.  latis;  petiolis  rubido- 
pilosis,  4-8  mm.  longis;  stipulis  lanceolato  subulatis,  1.5-2  mm.  longis,  glabris,  subpersis- 
tentibus ;  floribus  paucis  in  racemis  brevibus,  terminalibus,  glanduloso-pubescentibus ; 
In-acteis  lanceolatis;  pedicellis  rubido-pubescentibus,  2-5  mm.  longis;   ovario  ellipsoideo, 


46  CALIFORNIA  ACADEMY  OF  SCIENCES  [Proc.  4th  Ser. 

4-5  mm.  longo ;  hypanthio  rubido,  3  cm.  longo,  base  1.5-2  mm.  lato,  inde  angustiore,  inde 
gradatim  ampliato,  apice  5  mm.  lato,  externe  glanduloso-villoso,  interne  baseque  villoso ; 
sepalis  rubris,  lanceolatis,  12  mm.  longis,  4  mm.  latis,  glanduloso-pubescentibus,  apice  sub- 
ulatis;  petalis  rubris,  oblongo-lanceolatis,  7-9  mm.  longis,  3  mm.  latis,  acutis;  staminibus 
8  et  6  mm.  longis;  antheris  2  mm.  longis;  stylo  pubescente,  sepala  aequescente;  bacca 
cllipsoidea,  ca.  1  cm.  longa. 

Type:  Cajamarca,  prov.  Chota,  Peru:  pass  south  of  Conchan,  2500  m., 
Dec.  7,  1938,  Stork  and  Horton  10073,  Field  Museum  Herb.  1052248.  The 
reddish  condition  of  the  rather  long  hairs  and  the  narrow  leaves  are  charac- 
teristic of  this  proposed  species,  which  seems  near  to  F.  FiscJieri,  but  it  has  a 
shorter  hypanthium  and  pedicels  and  red  hairs.  It  is  a  pleasure  to  name  this 
for  Dr.  Harvey  Stork  of  Carleton  College. 

(38)  Fuchsia  furfuracea  Johnston 

(P],ate  6,  fig.  34) 
Fuchsia  furfuracea  Johnston,  Contr.  Gray  Herb.,  75:39, 1925. 

Shrubby,  of  unknown  size,  the  branchlets  dark,  sparsely  villous-hirsute ; 
leaves  opposite,  oblong-ovate,  obtuse  to  rounded  at  base,  acuminate  at  apex, 
denticulate,  brownish-pilose  on  both  surfaces,  especially  on  the  veins  beneath, 
with  11-13  principal  lateral  veins  on  each  side  of  the  midrib  ;  blades  6-9  cm. 
long,  2.5-4  (5)  cm.  wide;  petioles  8-20  mm.  long,  pubescent;  stipules  dark, 
subulate-lanceolate,  2-3  mm.  long;  flowers  few  in  terminal  racemes;  bracts 
pilose,  1-2  cm.  long,  lanceolate,  dentate ;  pedicels  pubescent-pilose,  2-4  cm. 
long;  ovary  elliptic,  villous,  6-8  mm.  long;  hypanthium  red,  tubular,  3.5^.5 
cm.  long,  slightly  enlarged  at  base,  then  constricted  so  as  to  be  scarcely  2  mm. 
wide  for  about  half  its  length,  then  rather  abruptly  widened  to  6-7  mm.  near 
the  cylindrical  summit,  villous  without  and  within;  sepals  red,  somewhat 
spreading,  lanceolate,  15-20  mm.  long,  4  mm.  wide,  villous  on  backs,  long- 
acuminate,  with  free  tips  2  mm.  long  in  bud ;  petals  purplish  red,  oblong-ovate, 
abruptly  acutish,  9-10  mm.  long;  longer  stamens  two-thirds  the  sepals, 
shorter  scarcely  one-half  the  sepals ;  anthers  2  mm.  long ;  style  villous,  slightly 
exceeding  sepals;  stigma  1.5  mm.  thick,  subcapitate,  slightly  4-lobed;  im- 
mature berry  ellipsoid,  barely  1  cm.  long. 

Type  locality,  Yungas,  La  Paz,  Bolivia.  Material  seen,  BOLIVIA :  La  Paz  : 
Yungas,  Bang  674,  type  collection  (F,  GH,  NY,  US)  ;  Unduavi,  at'3000  m.. 
Rushy  2511  (NY),  Characterized  by  its  long  stipules  and  pedicels,  brown 
pubescence,  long  sepal-tips,  it  seems  near  to  F.  Storkii,  F.  hirtella,  etc.,  in  its 
large  leaves,  narrow  hypanthium,  short  terminal  inflorescence.  The  "F. 
peruviana"  of  Johnston's  text  accompanying  the  original  description  is  un- 
doubtedly a  misprint  for  F.  holiviana. 

(39)  Fuchsia  Munzii  Macbride 

(riate7,  fig.  35) 
Fuchsia  Munzii  Macbride,  Field  Mus.  Nat.  Hist.,  Bot.  Ser.,  13,  part  4,  no.  1:559, 1941. 

Shrub,  the  younger  branches  1.5-3  mm.  thick,  very  finely  puberulent,  ap- 
parently somewhat  dark ;  leaves  mostly  opposite,  sometimes  ternate,  broadly 


Vol.  XXV]  MUNZ:  THE  GENUS  FUCHSIA  47 

elliptic-lanceolate  to  -ovate  or  -obovate,  subentire  to  inconspicuously  denticu- 
late, the  principal  lateral  veins  about  20  on  each  side  of  midrib,  blades  sub- 
cuneate  at  base,  acuminate  at  apes,  somewhat  bicolored,  quite  glabrous,  6-15 
cm.  long,  2.5-4.5  cm,  wide;  petioles  4-10  mm.  long;  stipules  subulate-lanceo- 
late, about  1  mm.  long,  caducous;  inflorescence  a  small  terminal  raceme  or 
panicle,  up  to  1  dm.  long,  with  rather  few  flowers,  very  minutely  strigulose 
bracts  linear  to  lanceolate,  5-20  mm.  long;  pedicels  5-15  mm.  long;  ovary 
linear,  4—5  mm.  long,  strigulose;  hypanthium  red,  tubular,  4-5  cm,  long, 
slightly  enlarged  at  base,  then  1  mm.  wide,  then  very  gradually  ampliate  and 
tubular,  4  mm.  wide,  minutely  and  rather  faintly  puberulent  without,  pubes- 
cent within ;  sepals  lanceolate,  somewhat  puberulent,  12  mm.  long,  about  3  mm. 
wide,  acute,  somewhat  spreading;  petals  narrowly  oblong,  11-12  mm.  long, 
acute ;  stamens  9  and  6  mm.  long;  anthers  2  mm.  long ;  style  pubescent  in  lower 
portion,  extending  about  to  sepal-tips ;  stigma  subglobose,  somewhat  4-lobed, 
2  mm.  thick ;  berry  subglobose,  almost  1  cm.  long. 

Type  collection,  PERU:  Dept.  Junin:  Rio  Masamerich,  2100-2200  m., 
Weherhauer  6648  (F,  GH,  US).  This  species  suggests  F.  corymhiflora  from 
which  it  differs  in  its  more  glabrous  leaves  and  rounder  fruit. 

(40)  Fuchsia  hirtella  H.B.K. 

(Plate  7,  fig.  36) 

Fuchsia  hirtella  H.  B.  K.,  Nov.  Gen.  et  Sp.,  6 :  107,  1823. 

F.  caracasensis  Fielding  and  Gardner,  Sert.  PI.,  pi.  29,  1844;  based  on  Linden  368,  sup- 
posed to  be  from  Caracas,  Venezuela,  in  1842.  The  plate  and  the  specimen  at  US  indicate 
synonymy  with  hirtella. 

Fuchsia  miniata  Planch  and  Linden,  PL  des  Serres,  Ser.  I,  pi.  8,  1852-1853;  based  on 
Linden  collection  from  "Nouvelle  Grenade."  In  my  key  would  seem  to  run  to  F.  Killipii 
on  basis  of  puberulent  stems,  but  cannot  be  that,  since  the  young  growth  is  not  canescent, 
so  must  be  hirtella.  I  have  seen  no  specimen  of  Linden,  labelled  miniata. 

Shrub  to  low  tree,  the  younger  branches  apparently  mostly  reddish,  2-4 
mm.  thick,  terete,  mostly  rather  densely  pubescent  with  short  somewhat  ap- 
pressed  and  interwoven  hairs,  older  branches  exfoliating  freely;  leaves  in 
whorls  of  3  or  4,  lance-oblong  to  elliptic-ovate,  rounded  to  cuneate  at  base, 
acute  to  acuminate  at  apex,  remotely  and  indistinctly  denticulate,  strigulose 
and  dark  green  above,  more  densely  pubescent  and  paler  beneath,  membrana- 
ceous, with  10-15  principal  lateral  veins  on  each  side  of  midrib  and  rather 
inconspicuous  submarginal  vein,  the  blades  6-12  (14)  cm.  long,  2.5^  (5)  cm. 
wide;  petioles  densely  pubescent,  5-20  (35)  mm.  long;  stipules  lance-linear, 
1-2  mm.  long,  deciduous;  flowers  in  terminal  pendant  corymbose  racemes 
which  may  be  grouped  in  a  panicle  1-2  dm.  long  and  equally  wide ;  bracts 
lance-ovate,  mostly  1-2  cm.  long,  strigulose ;  pedicels  more  loosely  pubescent, 
slender,  mostly  5-10  mm.  long ;  ovary  oblong-fusiform,  strigulose-pubescent, 
5-7  mm.  long,  1.5-2  mm.  thick;  hypanthium  tubular,  rose-red,  3-3.5  (4,5) 
cm.  long,  2.5-3  mm.  thick  at  base,  then  1.5-2  mm.  for  about  one-third  its 
length  and  very  gradually  widened  until  6-8  mm.  at  summit,  strigulose  with- 


•48  CALIFORXIA  ACADEMY  OF  SCIENCES  [Proc.  4th  Ser. 

out,  villous  witliiu  lower  half ;  sepals  4,  red,  somewhat  divergent,  lanceolate, 
acuminate,  subulate  for  terminal  2  mm.,  12-13  mm.  long,  2.5-3  mm.  wide, 
strigulose  on  backs;  petals  red,  lance-oblong,  glabrous,  acutish,  11-14  mm. 
long ;  stamens  10-14  and  6-10  mm.  long,  glabrous;  anthers  2-3  mm.  long ;  style 
red,  filiform,  almost  or  quite  glabrous,  exserted  4—8  mm.  beyond  sepals; 
stigma  subglobose,  4-lobed,  ca.  2  mm.  thick;  berry  ellipsoid,  puberulent,  14 
mm.  long,  6  mm.  wide. 

Type  locality,  in  mts.  near  Fusagasuga,  Cundimarca,  Colombia.  This  species 
seems  near  to  F.  polyantha  and  F.  holiviana  in  pubescence,  leaves,  inflores- 
cence, hypantliium,  etc.,  but  distinct  in  its  membranaceous  leaves,  ascending 
sepals,  and  long  fruit.  It  ranges  rather  widely  in  Colombia  as  can  be  seen  from 
citation  of  representative  material :  COLOMBIA :  without  locality,  Mutis 
1364  (US) ;  Bonpland  Herharium  1783,  probably  type  material,  it  is  rather 
fragmentary,  a  condition  referred  to  in  the  original  description  (fragment  F) . 
Antioquia :  Tamesis,  near  Medellin,  Toro  965  (NY)  ;  San  Pedro,  Tomds  and 
Daniel  1306  (F).  Caldas :  Pinares,  above  Salento,  2600-2900  m.,  Pennell  9195 
(Gil),  9324  (GH,  NY,  US);  Volcancito,  Holton  893  (NY);  Mediacion- 
Quindio,  Andre  2152  (NY) ;  "Laguneta"  to  "Magana,"  old  Quindio  trail, 
3000-3300  m.,  Pennell  9418  (GH,  NY,  US)  ;  Salento  to  "Laguneta,"  Pennell 
9141  (GH,  NY,  US)  ;  "Laguneta"  to  "Magaiia,"  Pennell  9425  (GH).  Tolima  ; 
"La  Lora"  to  "Cucarronera,"  new  Quindio  trail,  Hazen  9679  (GH).  Cundi- 
marca :  near  Bogota,  Ariste- Joseph  (US) ;  Sibate,  2900-3000  m.,  Pennell  2517 
(GH,  NY,  US)  ;  El  Peiion,  Pennell  2435  (NY)  ;  Cortesito,  pro.  de  Mariquita, 
Triana  3809  his  (US)  ;  San  Miguel,  extremo  W.  de  la  sabana  de  Bogota, 
Ciiatrecasas  6657  (US).  Cauca :  "Paletara"  to  "Calaguala,"  3000-3200  m., 
Pennell  7083  (GH,  NY,  US),  7094  (GH,  NY,  US);  "Canaan,"  Mt.  Purace, 
3100-3300  m.,  Pennell  6669  (GH,  NY,  US).  Narino :  Vulcan  Galera,  near 
Pasto,  Lehmann  5497  (GH,  F,  US)  :  Putumayo :  Corregimiento  El  Encano, 
Laguna  de  la  Cocha,  Paramo  El  Tabano,  Garcia  B.  7822  (US).  There  is  con- 
siderable variation  in  amount  of  hairiness  and  the  more  southern  more  hairj^ 
specimens  may  represent  F.  miniata,  but  at  present  I  am  unable  to  separate  it. 

(41)  Fuchsia  polyantha  Killip,  new  species 
(Plate?,  fig.  37) 

Bush,  up  to  about  1  m.  high;  young  stems  quite  simple,  purplish  red,  min- 
utely and  closely  appressed-puberulent  under  a  lens,  2-4  mm.  thick ;  leaves  not 
crowded,  ternate  or  quaternate,  rigid,  more  or  less  rugose,  elliptic  to  broadly 
oblong-lanceolate,  rounded  to  obtuse  at  base,  acuminate  at  apex  (the  apex 
not  in  the  same  plane  as  the  rest  of  the  blade  and  bent  at  an  angle  in  drying) , 
subentire,  light  green  and  glabrescent  above  except  for  the  appressed  puber- 
ulence  along  the  veins,  about  the  same  color  beneath,  minutely  appressed- 
puberulent  to  almost  glabrous  and  with  prominent  veins  beneath,  principal 
lateral  veins  14-18  on  each  side  of  midrib,  the  submarginal  not  well  formed ; 
blades  6-10  (12)  cm.  long,  2-3.5  cm.  wide;  petioles  purplish,  strigulose,  2-6 


Vol.  XXV]  MUNZ :  THE  GENUS  FUCHSIA  49 

mm.  long;  stipules  dark,  glabrous,  subulate,  1.5-2  mm.  long,  promptly  de- 
ciduous ;  inflorescence  a  compact,  pendulous,  strigulose,  many-flowered  pan- 
icle, 10-15  cm.  long;  bracts  lance-ovate,  1-2  cm.  long;  pedicels  strigulose, 
purplish,  8-12  mm.  long;  ovaries  strigulose,  purple,  fusiform,  6-7  mm.  long; 
liypanthium  purplish-red,  tubular,  3.5  cm.  long,  but  little  enlarged  at  base, 
1.5  mm.  wide,  then  1  mm.  wide  for  one-third  its  length  and  very  gradually 
ampliate  until  5  mm.  wide  at  summit,  minutely  strigulose  without,  pubescent 
within  basal  portion;  sepals  scarlet,  divergent,  linear-lanceolate,  strigailose 
on  backs,  13-15  mm.  long,  2.5-3  mm.  wide  at  base,  subulate  at  apex  for  1.5-2 
mm. ;  petals  crimson,  oblong-oblanceolate,  10-12  mm.  long,  3.5-4  mm.  wide, 
subacuminate,  glabrous;  stamens  11-12  and  7-8  mm.  long;  anthers  2  mm. 
long;  style  quite  glabrous,  equalling  or  slightly  exceeding  sepals;  stigma 
slightly  longer  than  wide,  1-1.5  mm.  thick;  berry  ellipsoid,  8  mm.  long,  4.5-5 
mm.  wide. 

Frutex,  1  m.  altus;  ramulis  subpuri3ureis,  minute  strigulosis,  2-4  mm.  diametro;  foliis 
ternatis  vel  quaternis,  rigidis,  subrugosis,  ellipticis  vel  late  oblongo-lanceolatis,  basi  rotun- 
datis  vel  obtusis,  apice  acuminatis,  subintegris,  subglabris  vel  in  venis  strigulosis ;  laminis 
6-10  (12)  cm.  longis,  2-3.5  cm.  latis;  petiolis  subpurj)ureis,  strigulosis,  2-6  mm.  longis; 
stipulis  glabris,  subulatis,  1.5-2  mm.  longis,  caducis;  inflorescentia  paniculata  floribus 
multis;  bracteis  lanceolato-ovatis,  1-2  cm.  longis;  pedicellis  strigulosis,  subpurpureis,  8-12 
mm.  longis;  ovario  striguloso,  fusiformi,  6-7  mm.  longo;  hypanthio  purpureo-rubro,  tubu- 
loso,  3.5  em.  longo,  basi  1.5  mm.  lato,  inde  1  mm.  lato,  gradatim  ampliato,  apice  5  mm.  lato, 
externe  striguloso,  interne  pubescente;  sepalis  coceineis,  divergentibus,  lineari-lanceolatis, 
strigulosis,  13-15  mm.  longis,  2.5-3  mm.  latis,  apice  subulatis;  petalis  rubris,  oblongo- 
oblanceolatis,  10-12  mm.  longis,  3.5-4  mm.  latis,  subaeuminatis,  glabris;  staminibus  11-12 
vel  7-8  mm.  longis ;  antheris  2  mm.  longis ;  stylo  glabro,  sepalis  aequalibus ;  stigmate  1-1.5 
mm.  lato;  bacca  ellipsoidea,  8  mm.  louga,  4.5-5  mm.  lata. 

Type:  Dept.  Narino :  between  Mayasquer  and  Tambo,  Colombia,  2800  m., 

Aug.  2,  1935,  Mexia  7571,  U.  S.  Nat.  Herb.  1662416.  Another  collection  from 
ECUADOR:  Prov.  Carchi:  between  Moran  and  Olivos,  3200-3500  m.,  July 
10-13,  1935,  Mexia  7474  (US).  The  proposed  species  is  near  to  F.  JiirteJla  in 
puberulence,  divergent  sepals,  inflorescence,  etc.,  but  has  the  leaves  more 
rigid  and  heavily  veined  with  a  sharper  apex,  the  style  less  exserted,  the  fruit 
shorter. 

(42)  Fuchsia  corymbiflora  Ruiz  and  Pavon 
(Plate7,  fig.  38) 

Fuchsia  corymbiflora  Ruiz  and  Pav^n,  F1.  Peruv.,  3:87,  pi.  325,  1802. 

F.  corymhosa  Pritzel,  Ind.  Ic,  1:469,  1866,  apparently  misspelling. 

F.macropetala  Presl,  Eel.  Haenk.,  2:28,  1835;  from  Huanuco,  Peru.  No  specimen  seen, 

but  description  fits  here. 
F.  dependens  Hook.,  Ic.  PL,  l:pl.  65,  1837;  based  on  Jameson  81,  I  am  not  sure  of  having 
seen  it. 
F.  velutina  Johnston,  Contr.  Gray  Herb.,  75:36,  1925. 

Semiscandent  shrub  up  to  4  or  5  m.  tall,  the  branches  rather  few,  simple,  the 
ultimate  branchlets  commonly  3-5  mm.  thick,  densely  canescent-puberulent 
or  -velutinous ;  leaves  opposite,  the  blades  elliptic  to  broadly  oblong-lanceolate, 


50  CALIFORNIA  ACADEMY  OF  SCIENCES  [Proc.  4th  Ser. 

subentire  to  inconspicuously  denticulate,  densely  and  minutely  soft-puber- 
ulent  on  both  surfaces,  rounded  to  obtuse  at  base,  acuminate  at  apex,  6-11  cm. 
long,  3-5  cm.  wide,  the  principal  lateral  veins  1-1—16  on  each  side  of  the  mid- 
rib, with  evident  submarginal  vein ;  petioles  6-14  (18)  mm.  long,  velutinous  ; 
stipules  lance-deltoid,  about  1  mm.  long,  caducous;  flowers  terminal,  few  to 
rather  many,  racemose  to  corymbose,  the  bracts  oblong-linear  to  -lanceolate. 
8-15  (20)  mm.  long ;  pedicels  6-10  (12)  mm.  long,  velutinous ;  ovary  ellipsoid, 
puberulent,  5-6  mm.  long;  hypanthium  tubular,  scarlet,  3.5-6  cm.  long, 
slightly  dilated  beyond  ovary,  then  1.5-2  mm.  wide  and  very  gradually  am- 
pliate  until  5-7  mm.  wide  at  summit,  puberulent  without,  pubescent  within 
much  of  its  length;  sepals  lance-linear,  erect-divergent,  scarlet,  12-15  mm. 
long,  3-3.5  mm.  wide,  acute  to  acuminate,  puberulent  on  backs ;  petals  deeper 
red,  narrow-oblong,  13-16  mm.  long,  about  3  mm.  wide,  acute  to  obtusish ; 
stamens  10-11  and  8-9  mm.  long,  with  reddish  filaments ;  anthers  2  mm.  long; 
style  pubescent  in  lower  part,  reddish,  exceeding  sepals  by  a  few  mm. ;  stigma 
subglobose,  slightly  lobed,  2.5-3  mm.  wide ;  fruit  round-ellipsoid,  densely 
puberulent,  6-10  mm.  long,  4-6  mm.  wide. 

Type  locality,  Chinchao,  Huanuco,  Peru.  As  here  treated  this  species  is 
quite  different  from  the  usual  concept.  I  agree  with  Johnston  (Contr.  Gray 
Herb.  75  :37. 1925)  that  most  material  which  has  been  treated  as  cory ml) (flora, 
including  illustrations,  is  loliviana,  but  I  feel  he  erred  in  saying  that  the' 
former  had  a  glabrous  inner  surface  to  the  hypanthium.  In  fact,  I  do  not  see 
what  he  did  include  in  corymhiflora.  However,  the  Ruiz  and  Pavon  specimen 
now  available  at  Field  Museum  reveals  definitely  what  it  is  and  shows  it  to  be 
a  plant  of  very  minute  soft  pubescence,  rather  short  round-ellipsoid  fruits,  and 
agreeing  with  material  occurring  from  Ecuador  to  north  central  Peru  and 
including  F.  dependens  and  velutina. 

Representative  material  is,  ECUADOR:  In  Andibus  Ecuadoriensibus, 
Spruce  5471  (NY).  Prov.  Carchi:  Guaca  San  Gabriel,  3300  m.,  Balls  7344 
(US)  ;  Canton  Tulcan,  near  Pun,  2987  m.,  Mexia  7595  (POM,  US).  Prov. 
Pichincha :  west  side,  Pichincha,  2700  m.,  Jameson,  possible  type  coll.  depen- 
dens (GH,  NY,  US)  ;  Canton  Quito,  road  Cotocallao  near  Nono,  2987  m., 
Mexia  7658  (POM,  US)  ;  between  Calacati  and  Nono,  Hartweg  986  (NY). 
Prov.  Cotopaxi  ? :  Hacienda  Sumbagua,  3600  m.,  Haught  2942a  (US) .  PERU  : 
Uuiz  and  Pavon,  ex  Herb.  Madrid,  probably  from  Huanuco  (F). Huanuco: 
Yanano,  2300m.,  Machride  3715,  type  no.  velutina  (F,  GH)  ;  road  Mirador  to 
Chanchao,  2300  m.,  Mexia  7765  (GH,  US);  Carpis,  road  Acamayo  to  Chin- 
chao, 2800  m.,  Mexia  04129  (GH). 

(43)  Fuchsia  abrupta  Johnston 
(Plate  7,  fig.  39) 
Fuchsia  abrupta  Johnston,  Contr.  Gray  Herb.,  75 :37, 1925. 

More  or  less  scandent  shrub,  with  glabrous  divaricate  twigs  4-5  mm.  thick ; 
leaves  opposite,  subcoriaceous,  lance-elliptic  to  broadly  oblanceolate,  7-12 


Vol.  XXV]  MUNZ :  THE  GENUS  FUCHSIA  51 

em.  long,  2.5^  cm.  wide,  subentire,  gradually  narrowed  at  base,  acuminate 
at  apex,  dark  green  and  subglabrous  above,  paler  and  puberulent  beneath 
especially  along  the  veins,  principal  lateral  veins  about  20  on  each  side  of 
midrib,  submarginal  vein  evident;  petioles  pubescent,  4-8  mm.  long;  stipules 
lance-deltoid,  2-3  mm.  long,  deciduous;  inflorescence  a  terminal  pendant 
raceme,  1-2  dm.  long ;  bracts  reflexed,  lanceolate,  8-18  mm.  long,  somewhat 
loose-pubescent ;  pedicels  2.5-4  cm.  long,  quite  glabrous ;  ovary  linear,  gla- 
broiLS,  6-8  mm.  long,  1  mm.  wide ;  hypanthium  scarlet,  tubular,  3.5-4.5  cm. 
long,  somewhat  nodose  at  very  base,  then  1  mm.  thick,  then  gradually  ampliate 
until  4-5  mm.  wide  at  summit,  glabrous  without,  'pubescent  within  lower 
portion ;  sepals  scarlet,  lanceolate,  13-15  mm  long,  acuminate,  erect-divergent, 
the  tips  subulate  for  1.5-2  mm.;  petals  oblong,  obtuse,  14-15  mm.  long;  sta- 
mens 10  and  8  mm.  long;  anthers  2  mm.  long;  stigma  subglobose,  slightly 
lobed,  2  mm.  wide ;  fruit  cylindric-oblong,  12-14  mm.  long,  4  mm.  wide. 

Type  collection  from  Cushi,  dept.  Huanuco,  Peru,  at  1600  m.,  Machride 
4541  (F,  GH,  US).  This  is  the  only  collection  I  have  seen;  it  has  glabrous 
stems,  narrow  leaves,  long  pedicels  and  glabrous  hypanthium. 

(44)  Fuchsia  Cuatrecasasii  Munz,  new  species 

(Plate  7,  fig.  40) 
Shrub;  young  twig's  glabrous  except  sometimes  toward  the  inflorescence, 
greenish,  2-4  mm.  thick ;  leaves  opposite,  membranaceous,  not  crowded,  ellip- 
tic-ovate, rounded  to  obtuse  at  base,  acute  to  subacuminate  at  apex,  subentire, 
light  green  and  quite  glabrous  above,  mostly  glabrous  beneath  except  for 
some  exceedingly  minute  appressed  puberulence  along  the  veins,  principal 
lateral  veins  10-12  on  each  side  of  midrib,  submarginal  weakly  developed ; 
blades  4-16  cm.  long,  2-9.5  cm.  wide;  petioles  more  or  less  strigulose,  2-18 
mm.  long;  stipules  deltoid,  more  or  less  connate  and  persistent,  1-1.5  mm. 
long;  flowers  few,  in  short  terminal  racemes;  bracts  lanceolate,  5-15  mm. 
long;  pedicels  slender,  7-13  mm.  long,  almost  glabrous;  ovary  glabrous,  ellip- 
soid, purplish  red,  4—5  mm.  long,  1.5-2.5  mm.  wide ;  hypanthium  bright  red, 
29-40  mm.  long,  2-3  mm.  wide  at  base,  then  slightly  narrowed  for  one-third 
its  length,  then  gradually  ampliate  until  6-8  mm.  wide  at  summit,  glabrous 
to  sparingly  pubescent  without,  densely  pilose  within ;  sepals  scarlet,  diver- 
gent, 10-13  mm.  long,  4-5  mm.  wide,  glabrous,  subulate  at  apex  for  about  1 
mm. ;  petals  scarlet,  narrowly  oblong-ovate,  8-10  mm.  long,  about  half  as  wide, 
somewhat  pointed,  glabrous;  stamens  7-8  and  5-6  mm.  long;  anthers  1  mm. 
long ;  style  sparsely  pubescent,  not  equalling  petals ;  stigma  barely  1  mm. 
thick;  berry  apparently  oblong-ellipsoid. 

Frutex;  ramulis  glabris,  2—4  mm.  diametro;  foliis  oppositis,  membranaceis,  elliptico- 
ovatis,  basi  rotundatis  vel  obtusis,  apice  acutis  vel  subacuminatis,  subintegris,  subglabris, 
laminis  4-16  cm.  longis,  2-9.5  cm.  latis ;  petiolis  strigulosis,  2-18  mm,  longis ;  stipulis  del- 
•  toideis,  connatis,  1-1.5  mm.  longis;  floribus  paucis,  in  racemis,  bracteis  lanceolatis,  5—15 
mm.  longis;  pedicellis  tenuibus,  7-13  mm.  longis;  ovario  subglabro,  ellipsoideo,  4-5  mm. 
longo,   1.5-2.5  mm.  lato;   hypanthio  rubro,  29-40  mm.  longo,  basi   2-3   mm.  lato,  inde 


52  CALIFOBNIA  ACADEMY  OF  SCIENCES  [Proc.  4th  Ser. 

angustiore,  inde  ampliato,  apice  6-8  mm.  lato,  exteriore  subglabro,  interiore  piloso ;  sepalis 
coccineis,  divergentibus,  10-13  mm.  longis,  4-5  mm.  latis,  glabris,  1  mm.  subulatis ;  petalis 
coccineis,  anguste  oblongo-ovatis,  8-10  mm.  longis;  staminibus  7-8  vel  5-6  mm.  longis; 
antheris  1  mm.  longis;  stylo  sparse  pubescente;  stigmate  1  mm,  lato;  bacca  oblongo- 
ellipsoidea. 

Type :  from  Comisaria  del  Caqueta,  Colombia :  Cordillera  Oriental,  verti- 
ente  oriental  Quebrada  del  Rio  Hacha,  bosque  abierto  en  Cajon  de  Pulido, 
1700  m.  alt.,  March  26,  1940,  /.  Cuatrecasas  8738,  U.  S.  Nat.  Herb.  1796401. 
An  additional  collection  from  the  Com.  de  Putumayo,  COLOMBIA,  is  from 
between  El  Sileneio  and  La  Cabaiia  (carretera  de  Sibundoy  a  Urcusique),  at 
2200-2400  m.,  Dec.  31, 1940,  Cuatrecasas  11495  (US).  The  species  seems  quite 
near  F.  abrupt  a  of  Peru,  but  has  shorter  leaves  with  fewer  veins,  shorter 
sepals,  shorter  inflorescence  and  pedicels. 

(45)  Fuchsia  Killipii  Johnston 
(Plates,  fig.  41) 
Fuchsia  Killipii  Johnston,  Contr.  Gray  Herb.,  81:94,  1928. 

Somewhat  scandent  shrub,  up  to  4  m.  tall,  the  younger  branches  brownish, 
sub  terete,  somewhat  puberulent ;  leaves  opposite  or  ternate,  elliptic,  acute  at 
base,  acute  to  acuminate  at  apex,  subentire,  almost  equally  green  on  both 
surfaces,  somewhat  coriaceous,  with  about  12  principal  lateral  veins  on  each 
side  of  the  midrib  and  evident  submarginal  vein,  subglabrous  to  minutely 
strigulose  on  both  surfaces,  the  blades  5-9  cm.  long,  2-3.5  cm.  wide;  petioles 
5-10  mm.  long;  stipules  deltoid,  scarcely  1  mm.  long,  deciduous;  flowers  in 
upper  axils  and  in  terminal  drooping  racemes  or  panicles,  rather  numerous ; 
bracts  elliptic-ovate,  1-2  (3)  cm.  long;  pedicels  subfiliform,  1-2  cm.  long  at 
anthesis,  2-4  cm.  long  in  fruit,  subglabrous;  ovary  round-ellipsoid,  incon- 
spicuously puberulent,  3—4  mm.  long ;  hj^ianthium  red,  tubular,  3-4  cm.  long, 
1.5-2  mm.  wide  at  the  base,  then  narrowed  for  one-third  its  length,  then 
rather  abruptly  widened  until  5—7  mm.  at  summit,  glabrous  without,  retrorse- 
villous  within;  sepals  lanceolate,  divergent,  acuminate,  14-15  mm.  long,  4 
mm.  wide,  somewhat  puberulent  toward  apex  of  outer  surface,  subulate  1-1.5 
mm.  at  tips ;  petals  bright  red,  oblong-oblanceolate,  13-15  mm.  long,  2.5-4  mm. 
wide,  obtuse  to  short-acute,  glabrous;  stamens  12-13  and  9-10  mm.  long, 
glabrous ;  anthers  white,  2  mm.  long ;  style  villous  through  much  of  its  length, 
up  to  5  mm.  longer  than  sepals;  stigma  4-lobed,  subglobose,  1.5  mm.  long; 
berry  globose,  5-6  mm.  thick. 

Type  locality,  Rio  Surata  above  Surata,  Santander,  Colombia.  The  species 
is  distinguished  from  its  closest  relatives  by  having  fewer  veins,  smaller  fruit, 
and  longer  pedicels.  I  have  seen  the  following  collections  from  COLOMBIA  : 
Santander  :  Rio  Surata  valley,  above  Surata,  2000-2300  m.,  Killip  and  Smith 
16695,  type  coll.  (Gil,  NY,  US),  16604  (GH,  US)  ;  near  Charta,  2600  m., 
Killip  and  Smith  18854  (GH,  NY.  US),  17459  (GH,  NY,  US)  ;  near  La  Baja, 
2200-2600  m.,  Killip  and  Smith  16776  (F,  GH,  NY,  US) . 


Vol.  XXV]  MUNZ:  THE  GENUS  FUCHSIA  53 

(46)  Fuchsia  boliviana  Carr. 

Fuclisia  boliviana  Cabr.,  Eev.  Hort.,  1876:150,  with  fig.,  1876. 

Bushy  shrub  or  small  tree  up  to  4.5  (6)  m.  high,  with  rather  open  brauch- 
ing-,  the  branchlets  spreading  or  dependent,  mostly  2-4  (5)  mm.  thick,  terete 
to  angled,  densely  pubescent;  leaves  opposite,  sometimes  alternate  or  ternate, 
elliptic  to  oblong-lanceolate  or  -ovate,  obtuse  to  rounded  at  base,  acute  to 
acuminate  at  apex,  usually  somewhat  denticulate,  soft-pubescent  on  both 
surfaces  to  pilose  or  puberulent  or  glabrescent,  somewhat  paler  beneath,  with 
about  12-24  principal  lateral  veins  on  each  side  of  midrib ;  leaf -blades  5-15 
(20)  cm.  long,  3-6  (10)  cm.  wide;  petioles  1.9-3  (4.5)  cm.  long,  pubescent; 
stipules  subulate-lanceolate,  scarcely  1  mm.  long,  deciduous;  flowers  few  to 
many,  in  terminal  drooping  pubescent  corymbs  or  panicles  which  elongate 
in  fruit;  bracts  7-20  (30)  mm.  long,  3-9  mm.  wide,  usually  lanceolate  and 
reflexed;  pedicels  5-15  mm.  long;  ovary  oblong-linear,  5-7  mm.  long,  1.5-2 
mm.  thick;  hypanthium  tubular,  usually  dark  red,  3-6  cm.  long,  slightly  en- 
larged at  base,  then  1.5-2  mm.  wide  and  very  gradually  ampliate  until  3.5-7 
mm.  at  apex,  more  or  less  pubescent  without  and  within ;  sepals  usually  dark 
red,  lanceolate,  acuminate,  10-20  mm.  long,  2.5-5  mm.  wide,  spreading- 
reflexed  in  anthesis,  more  or  less  pubescent  without ;  petals  dark  to  light  red, 
erect  to  divergent,  oblong,  9-13  mm.  long,  3-4.5  mm.  wide,  acute ;  episepalous 
stamens  equaling  or  slightly  exceeding  petals ;  alternate  three-fourths  as  long ; 
filaments  glabrous,  reddish ;  anthers  light  in  color,  2-3  mm.  long ;  style  slender, 
pubescent  within  hypanthium,  scarcely  reaching  tips  of  sepals ;  stigma  sub- 
globose,  slightly  lobed,  1-2  mm.  thick;  berry  oblong-ellipsoid,  1-2  cm.  long, 
4-6  mm.  thick,  pubescent. 

As  here  understood,  this  species  is  common  and  widespread,  including  most 
of  the  material  once  referred  to  F.  corymbiflora,  from  which  it  can  be  dis- 
tinguished by  its  usually  longer  and  more  spreading  hairs,  spreading  sepals, 
apparently  less  vinelike  habit,  more  apiculate  buds  and  elongate  fruit. 

Key  to  Varieties  of  Fuchsia  'boliviana 
A.  Hypanthium  3-4  cm.  long ;  sepals  10-18  mm.  long.  South  of  Ecuador. 

B.  Plant  pilose  or  pubescent  with  spreading  hairs.  Peru  to  Argentina.  46a.  var.  typica. 

BB.  Plant  minutely  puberulent.  Bolivia 46b.  forma.  Tpuherulenta. 

A  A.  Hypanthium  5-6  cm.  long;  sepals  17-21  mm.  long.  North  of  Ecuador. 

46c.  var.  luxurians. 

(46a)  Fuchsia  boliviana  Carr.  var.  typica  Munz,  new  name 

(Plates,  fig.  42) 

F.  boliviana  Carr.,  Kev.  Hortic,  1876:150,  with  pi.,  1876;  Johnston,  Contr.  Gray  Herb., 

75:37,  1925. 
F.  corymbiflora  of  many  authors,  not  of  Euiz  and  Pavon,  1802. 

Plant  usually  soft-pilose  or  -pubescent  on  all  new  growth ;  hypanthium 
2.5-4  em.  long ;  sepals  mostly  12-18  mm.  long. 

Type  locality,  "sur  les  montagnes  a  la  Bolivie,  a  6000  metres  environ  d'alti- 


54  CALIFORNIA  ACADEMY  OF  SCIENCES  [Proc.  4th  Ser. 

tucle."  Such  an  altitude  is  quite  impossible ;  I  have  seen  specimens  from  as 
high  as  4000  m.;  the  variety  ranges  from  Peru  to  northern  Argentina.  In  the 
southern  part  of  the  range  the  plants  tend  to  be  less  densely  pilose  and 
smaller-flowered.  Representative  material  may  be  cited  as  follows,  PERU : 
Huanuco:  5  km.  SE.  of  Carpish,  2850  m.,  Stork  9921  (F).  Huaneavelica :  at 
2400  m.,  Weherhauer  6565  (F,  GH).  Ayacucho:  Prov.  de  la  Mar,  Tambo, 
Weberbauer  5555  (F,  GH,  US)  ;  Ccarrapa,  between  Huanta  and  Rio  Apuri- 
mac,  2800  m.,  Killip  and  Smith  22280  (NY,  US).  Apurimac :  Abancay : 
Ampuy,  3100  m.,  Stork,  Horton  and  Vargas  10595  (F)  ;  Huancarama,  3200 
m.,  West  3783  (GH).  Cuzco :  Urubamba  Valley,  West  6462  (GH,  US),  Her- 
rera  2094  (US),  1123  (F,  US)  ;  Paucartambo,  Balls  6715  (US);  OUantay- 
tambo.  Cook  and  Gilbert  290  (US).  BOLIVIA:  La  Paz:  Yungas,  Bang  327 
(F,  GH,  NY,  US),  Rusby  1071  (F,  GH,  NY,  US)  ;  Sorata,  2600-2800  m., 
Mandon  622  (F,  GH,  NY,  US).  Santa  Cruz:  Samaipata,  1800  m.,  Steinbach 
3755  (POM);  Comarapa,  Steinbach  8459  (GH,  NY).  Cochabamba :  Prov. 
Chapare:  Incacorral-Aduana,  2700  m.,  Steinbach  9538  (F);  Prov.  Sacaba: 
Cerro  de  Ineachaea,  2500  m.,  Steinba.ch  5763  (F,  GH,  US).  ARGENTINA: 
Jujuy :  San  Lorenzo,  Joergensen  36041  (POM) .  Tueuman  :  Depto  Chicligasta, 
Estancia  Las  Pavas,  Ve7ituri  4657  (F,  US)  ;  Villa  Nougues,  1200  m.,  Munz 
15469  (NY,  POM,  US);  Las  Lenguas  al  Clavillo,  1750  m.,  Munz  15475 
(POM)  ;  Depto.  Famailla:  Quebrada  de  Lules,  Venturi  1295  (F,  CAS,  GH, 
US). 

(46b)  Fuchsia  boliviana  Carr.  forma  puberulenta  Munz,  new  form 

Plant  with  dense  minute  subappressed  puberulence;  hypanthium  3-4  cm. 
long;  sepals  mostly  10-16  mm.  long.  (Planta  cum  puberulencia  minuta  et 
densa :  hypanthio  3-4  cm.  longo.) 

Type:  from  La  Paz,  Bolivia:  Nor  Yungas:  Milluguaya,  1800  m.,  Buchtien 
732,  Field  Mus.  no.  588700;  isotypes  F,  GH,  NY,  US.  Other  material  all  from 
BOLIVIA  :  La  Paz :  Unduavi  Valley,  2000-2600  m.,  Bro.  Julio  478  (US)  ;  Nor 
Yungas:  Coroica  to  La  Paz,  Mexia  04286  (GH) ;  Sur  Yungas:  SirUpaya  near 
Yanaeachi,  2100  m.,  Buchtien  732  (F,  GH)  ;  Colaya,  Mexia  7809  (F,  US). 
Cochabamba :  Prov.  Chapare,  110  km.  north  east  of  Cochabamba,  near  Chi- 
more,  800  m.,  Eyerdam  24740  (F).  This  form  resembles  F.  corynibiflora  in 
puberulence,  though  the  latter  has  it  less  dense,  but  F.  boliviana  has  spreading 
sepals  and  cor^m&i^ora  does  not. 

(46c)  Fuchsia  boliviana  Carr.  var.  luxurians  Johnston 

Fuchsia  boliviana  Carr.  var.  luxurians  Johnston,  Contr.  Gray  Herb.,  75:38,  1925. 

F.  cuspidata  Fawcett  and  Rendle,  Journ.  Bot.,  64:105,  159,  1926,  as  to  tj'pe  cited:  Harris 

5825. 
F.  corynibiflora  of  many  authors:  LiNDL.,  Bot.  Reg.,  26:pl.  70,  1840;  Hook,,  Bot.  Mag., 

69:  pi.  4000,  1843;  not  of  R.  and  P.,  1802. 

Plant  soft-pilose  to  -pubescent;  hypanthium  5-6  cm.  long;  sepals  17-21 
mm.  long. 


Vol.  XXV]  MVNZ:  IRE  GENUS  FUCHSIA  55 

Type  locality,  Colonia  Tovar,  Venezuela.  Representative  material,  JA- 
MAICA :  Cinchona,  Harris  7643  (NY),  Harris  and  Lawrence  C 15438  (NY)  ; 
St.  Helen's  Gap,  St.  Andrew,  3Iaxon  and  Killip  571,  1475  m.  (F,  GH,  NY, 
US).  GUATEMALA  :  Cartago  :  Cartago,  1350  m..  Smith  4804  (US)  ;  Zacate- 
IDequez:  San  Rafael,  2000  m.,  Smith  2176  (US).  San  Marcos:  Tajumulco, 
northwest  slope  of  Volcan  Ta.jmnnlco,  2300  m.,  Steyermark  36510  (F). 
Chimaltenango :  San  Marcos,  Johnson  1231  (F).  EL  SALVADOR:  bought 
in  market  of  San  Salvador,  brought  from  Volcan  de  San  Salvador,  C  alder  on 
2345  (F,  US) .  COLOMBIA  :  Antioquia  :  "Las  Minitas,"  so  of  Caldas,  2650  m., 
Pennell  10970  (GH,  NY);  Medellin,  Toro  721  (NY).  Caldas :  Salento  to 
"Lagunita,"  old  Quindio  trail,  2000  m.,  Pennell  10594  (GH).  Cundimarca : 
Bogota,  2700  m.,  Schultze  135  (US),  Perez  4802  (US).  VENEZUELA: 
Colonia  Tovar,  1800-2000  m.,  Pittier  9252,  type  no.  (GH,  NY,  US),  Allart 
354  (NY,  US),  Ja/iw  439  (US). 

(47)  Fuchsia  fulgens  DeCandolle 
(Plate  8,  fig.  43) 

Fuchsia  fulgens  DC,  Prodr.,  3:39,  1828;  in  Sesse  and  Mo§ino,  Icon.  Fl.  Mex.  ined.,  362; 

LiNDLEY,  Bot.  Eeg.,  24:1, 1838;  Curtis,  Bot.  Mag.,  67:3801, 1840;  Hemsley,  Biol.  Centr. 

Amer.,  Bot.,  1:457,  1880;  Standley,  Contr.  U.  S.  Nat.  Herb.,  23:1078,  1924. 
Ellohium  fulgens  Lilja,  Linnaea,  15:262, 1841. 

Shrub,  commonly  3-12  dm.  tall,  with  tuberous-thickened  roots,  soft  woody 
stems,  few-branched;  young  branches  sparsely  pubescent,  these  and  the  leaves 
tinged  red ;  leaves  broadly  ovate  to  oblong-ovate,  opposite,  paler  beneath  than 
above,  fiue-pubescent  on  both  surfaces,  rounded  to  more  or  less  cordate  at 
base,  abruptly  short-acuminate,  shallowly  and  evenly  serrulate,  the  blades 
thin,  5-17  cm.  long,  3-12  cm.  wide,  with  about  10  principal  lateral  veins  on 
each  side  of  the  midrib;  petioles  1.5-10  cm.  long;  flowers  several,  in  short 
racemes  with  an  axis  5-20  cm.  long;  lower  bracts  leafy  and  ovate,  1-3  cm.  long ; 
upper  reduced,  lanceolate,  0.5  cm.  long,  puberulent ;  pedicels  pendant,  very 
slender,  1-2  (3.5)  em.  long;  hypanthium  5-6.5  cm.  long,  about  2  mm.  wide  at 
base,  then  scarcely  narrowed  and  then  gradually  ampliate  until  5-7  mm.  wide 
near  mouth,  dull  scarlet,  subglabrous  to  puberulent  without,  pubescent  with- 
in ;  sepals  4,  yellowish  to  greenish  with  somewhat  pale  or  yellow  or  red  base, 
ovate-lanceolate,  12-14  mm.  long,  subglabrous  except  for  the  sparse  pubes- 
cence near  the  acuminate  tips;  petals  bright  red,  oblong-ovate,  obtuse,  8  mm. 
long,  6  mm.  wide ;  stamens  about  10  and  7  mm.  long ;  anthers  2-3  mm.  long, 
filaments  whitish  to  reddish ;  style  filiform,  pink,  pubescent  near  base,  slightly 
exceeding  sepals;  stigma  greenish,  subconic,  2  mm.  long;  fruit  ellipsoid,  sub- 
glabrous, dark,  2  or  more  em.  long. 

Type  locality  not  known  ;  the  species  is  Mexican,  is  near  to  such  long-tubed 
species  as  F.  holiviana,  corymhiflora,  etc.,  but  quite  distinct  in  its  green  sepals 
and  leaves  of  cordifolia  type.  It  may  be  an  epiphyte,  may  grow  on  ledges, 
walls,  or  more  open  ground.  Material  seen,  "Fuchsia  cordifolia  N.  No.  195.  t 


56  CALIFOBNIA  ACADEMY  OF  SCIENCES  [Proc.  4th  Ser. 

2.  f.  159  i.  c.  195,"  Sesse,  Mocino  et  al  5211  (F)  ;  photo  of  Geneva  copy  of 
Sesse  and  Mocino  plate  (F)  ;  H.  B.  Basil,  Aug.  1841  (NY).  MEXICO  :  Mex- 
ico City,  Russell  and  Souviron  132  (US).  Mexico:  Temascaltepec  District, 
Rincon,  Hinton  1144  (GH,  US) .  Michoacan  :  near  Tarascon,  Pringle  11988^2, 
at  2000  m.,  (F,  GH,  US) ;  near  Patzcuaro,  Pringle  4123  (F,  GH,  MICH,  NY, 
POM,  US)  ;  Morelia,  Campanario,  2200  m.,  Arsene  in  1911  (US)  ;  Monte  San 
Miguel,  Arsene  6673  (US)  ;  2  miles  north  of  Zitacaura,  Hitchcock  and  Stan- 
ford 7207  (US) ;  Tancitaro,  Leavenworth  303  (F).  Jalisco:  Volcano  Colima, 
2100  m.,  Goldsmith  22  {Gil). 

(48)  Fuchsia  decussata  Ruiz  and  Pa  von 

(Plate  8,  fig.  44) 

Fuchsia  decussata  Euiz  and  Pav6n,  PL  Peruv.,  3  :88,  pi.  323,  fig.  b.,  1802. 

F.scandens  Krause,  Fedde  Eep.,  Nov.  sp.  1:171,  1905;  based  on  Weberbauer  3324  from 

Monzon,  Huanuco,  Peru ;  have  seen  photograph. 
F.  fusca  Krause,  Engl.,  Jahrb.,  37:599,  1906;  based  on  Weberbauer  4975  from  near  Cuzco; 

have  seen  photograph ;  flowers  large  like  Macbride  4414. 
F.  fontinalis  Macbr.,  Candollea,  8:25,  1940.  Eeduced  by  Macbride,  Field  Mus.  Nat.  Hist., 

Bot.,  13,  part  4:552,  1941. 

Shrub,  supported  by  other  vegetation,  up  to  3  ni.  high,  with  long  flexuous 
branches,  the  youngest  reddish,  1-3  mm.  thick,  densely  brown-pubescent  with 
incurved  hairs ;  leaves  ternate  or  opposite,  numerous,  those  of  main  stems 
often  remote  and  supplemented  by  smaller  ones  on  reduced  lateral  branchlets, 
coriaceous,  elliptic  to  elliptic-lanceolate  or  -ovate,  cuneate  at  base,  acute  at 
apex,  serrate  to  serrulate,  dark  green  and  puberulent  to  strigulose  to  glabres- 
cent  above,  paler  and  short-pilose  beneath,  especially  along  veins,  principal 
lateral  veins  6-10  on  each  side  of  midrib,  submarginal  vein  somewhat  devel- 
oped, blades  1.5-5  cm.  long,  0.5-1.5  (2)  cm.  wide;  petioles  pubescent-pilose, 
3-8  (15)  mm.  long;  stipules  dark,  subulate,  1-1.5  mm.  long,  persisting; 
flowers  solitary  in  axils;  pedicels  subfiliform,  pilose,  1.5-3  cm.  long;  ovary 
ellipsoid,  3-4  mm.  long,  more  or  less  pilose;  hypanthium  13-16  (22)  mm.  long, 
red,  2-3  mm.  wide  at  base,  then  very  little  narrowed,  3.5-4.5  mm.  wide  at 
summit,  sparsely  pilose  without,  glabrous  within ;  sepals  divaricate,  lance- 
oblong,  red  with  green  tips,  acute,  7-9  (11)  mm.  long,  2.5  mm.  wide,  sparsely 
pilose  on  backs;  petals  red,  oblong-obovate,  obtusish  to  apiculate,  6-7  (10) 
mm.  long,  3-4  mm.  wide ;  stamens  8-9  and  6-7  mm.  long,  more  or  less  exserted; 
anthers  1.5-2  mm.  long;  style  glabrous,  equaling  or  exceeding  sepals  by  2-3 
(6)  mm. ;  stigma  1.5  mm.  wide;  fruit  red,  oblong,  obtusely  angled,  8-10  mm. 
long,  5-6  mm.  thick. 

Type  locality,  Muila,  dept.  Huanuco,  Peru.  The  species  ranges  at  elevations 
of  2800-3400  m.  from  Hiianuco  to  Cuzco.  It  has  considerable  variation  in 
width  and  aeuteness  of  petals.  It  falls  in  a  group  with  axillary  flowers  and 
short  hypanthium  and  seems  to  be  a  possible  derivative  from  the  cordifolia 
type.  Macbride  (Field  Mus.  Nat.  Hist.  Bot.  Ser.  13,  part  4,  no.  1 :554,  1941) 


Vol.  XXV]  MUNZ :  THE  GENUS  FUCHSIA  57 

maintains  F.  fiisca  as  a  species  on  the  basis  of  the  wider  petals  and  brownish 
pubescence,  but  this  latter  character  seems  to  be  a  general  tendency  in  many 
specimens.  Representative  material  of  F.  decussata  are:  PERU:  without 
definite  locality  given,  Ruiz  and  Pavon,  F.  decussata  sp.  nov.  Herb,  at  Geneva 
(photo  F,  POM),  ex  Herb.  Madrid  (F).  Huanuco:  15  miles  SE  of  Huanuco, 
2900  m.,  Machride  and  Featherstone  2124  (F,  GH,  US),  2081  (F,  GH,  US), 
2107  (F,  GH,  US)  ;  Panao,  Machride  3615  (F,  GH,  US)  ;  Tambo  de  Vaca, 
Machride  4414  (F,  GH)  with  petals  10  mm.  long;  Carpish,  Stork  and  Horton 
9911  (F)  with  very  narrow  petals;  Carpis,  between  Acomayo  and  Chinchos, 
Mexia  04126  (GH),  04128  (GH)  ;  Monzon,  Weherhauer  3324,  type  scandens 
at  Berlin  (photo  F,  POM).  Ayucucho :  prov.  Huanta,  between  Tambo  and 
Apurimac  River,  Weherhauer  5578  (F).  Cuzco :  between  Cuzco  and  Santa 
Anna,  below  Yanamanche,  Weherhauer  4975,  type  fusca  at  Berlin  (photo  F, 
POM). 

(49)  Fuchsia  hypoleuca  Johnston 

(Plate  8,  fig.  45) 
Fuchsia  liypoleuca  Johnston,  Coiitr.  Gray  Herb.,  75:34,  1925. 

Shrub,  1-2  m.  high ;  young  stems  densely  canescent-pubescent  with  more 
or  less  incurved  hairs,  slender,  1.5-3  (4)  mm.  thick;  leaves  opposite  or  ternate, 
rigid,  crowded,  lanceolate,  obtuse  to  acute  at  base,  acute  at  apex,  remotely 
serrulate  in  upper  half,  more  or  less  revolute,  veiny  almost  rugose,  greenish 
and  somewhat  strigulose  above,  pale  and  densely  pilose-pubescent  beneath, 
tending  to  have  interwoven  fungus  mycelium  underneath,  principal  lateral 
veins  6-10  on  each  side  of  midrib,  submarginal  vein  present ;  blades  3-6  (7.5) 
cm.  long,  0.8-1.7  cm.  wide;  petioles  pubescent,  5-10  mm.  long;  stipules  sub- 
ulate, 2-3  mm.  long,  deciduous;  pedicels  few,  solitary  in  axils,  filiform,  pen- 
dulous, pubescent,  1.5-2  cm.  long;  ovary  ellipsoid,  pubescent,  4  mm.  long; 
hypanthium  red,  18-23  mm.  long,  about  2.5  mm.  wide  at  base,  then  somewhat 
narrowed  and  then  gradually  ampliate  until  5  mm.  wide,  pubescent  and 
villous  within  lower  half ;  sepals  divaricate,  scarlet,  lanceolate,  9-12  mm.  long, 
3  mm.  wide,  pubescent  on  backs,  subulate  for  1  mm.  at  apex;  petals  scarlet, 
obovate,  7  mm.  long,  4^5  mm.  wide,  rounded  at  apex ;  stamens  7-8  and  5-6 
mm.  long;  anthers  1.5  mm.  long;  style  pubescent  within  the  hypanthium, 
equalling  the  sepals  or  exceeding  them  by  2-3  mm.;  stigma  green,  1.5  mm. 
wide ;  fruit  apparently  oblong-ellipsoid,  4-angled,  8-10  mm.  long,  3-4  mm. 
wide. 

Type  locality,  between  Loja  and  San  Lucas,  prov.  Loja,  Ecuador.  The 
species  seems  to  me  in  its  floral  position  and  h^^Danthium  shape  to  be  near 
F.  decussata,  but  amply  distinct  in  the  narrow^  crowded  leaves  which  are 
whitish  beneath.  I  have  seen  the  following  specimens,  ECUADOR :  without 
definite  locality,  Jameson  (F,  US),  southern  provinces,  Jameson  (US)  ;  be- 
tween Loja  and  San  Lucas,  2100-2600  m.  Hitchcock  21440,  type  coll.  (GH, 
NY,  US). 


58  CALIFORNIA  ACADEMY  OF  SCIENCES  [Proc.  4th  Rer. 

(50)  Fuchsia  scabriuscula  Bentliam 

(Plate  8,  fig.  46) 
Fuchsia  scabiuscula  Benth.,  PI.  Hartweg.,  177, 1845. 

Spreading  shrub,  7-20  dm.  tall,  the  younger  twigs  sometimes  reddish,  2-4 
mm.  thick,  densely  pilose  with  white  to  somewhat  brownish  hairs ;  leaves 
opposite,  elliptic  to  elliptic-obovate,  rigid  and  very  veiny,  even  rugose, 
rounded  to  obtuse  at  the  base,  acute  to  subacuminate  at  the  apex,  subentire, 
somewhat  strigose  above,  paler  and  pilose  beneath,  especially  on  veins,  with 
10-15  principal  veins  on  each  side  of  midrib  and  with  prominent  submarginal 
vein;  blades  3-10  (13)  cm.  long,  1.5-5  (7)  cm.  wide;  petioles  5-25  mm.  long, 
densely  pilose ;  stipules  subulate,  pubescent,  1.5-3  mm.  long,  quite  persistent ; 
pedicels  solitary,  axillary,  slender,  curved,  pilose,  1-2  cm.  long;  ovary  nar- 
rowly fusiform,  densely  pilose,  5-7  mm.  long;  hypanthium  bright  red,  15-18 
mm.  long,  1.5-2  mm.  wide  at  base,  then  somewhat  narrowed,  then  gradually 
arapliate  until  4.5-5  mm.  wide  at  summit,  densely  pilose  without  and  within 
the  basal  half ;  sepals  bright  red,  divergent,  lance-oblong,  8-9  mm.  long,  ea. 
3  mm.  wide,  subulate  at  apex  for  about  1.5  mm.,  pilose  on  back;  petals  red, 
obovate-oblong,  8-9  mm.  long,  about  0.5  mm.  wide,  rounded  at  apex ;  stamens 
about  8  and  6  mm.  long;  anthers  1.5  mm.  long ;  style  densely  pubescent  within 
hypanthium,  extending  about  to  tips  of  petals ;  stigma  greenish,  about  2  mm. 
Made ;  fruit  oblong,  10-12  mm.  long,  3-4  mm.  wide. 

Tj^pe  locality,  "In  declivities  oecidentali  Andium  Quitensium."  Ecuador, 
This  species  with  F.  verrucosa  seems  to  me  to  be  near  F.  decussata,  but  with 
a  somewhat  more  narrow  base  to  the  hypanthium.  It  is  characterized  by  the 
very  veiny,  rugulose  leaves,  pilose  stems  and  flowers,  and  short  hypanthium. 
It  seems  to  be  confined  to  Ecuador,  from  which  I  have  seen  the  following 
material:  in  Andibus  Ecuadorensibus,  Spruce  5038  (GH,  NY).  Pichincha : 
in  declivitate  oecidentali  Andium  Quitensium,  Hartweg  987,  type  no.,  Geneva 
Herb,  (photo  F,  POM;  leaf  F) ;  Sive  Andium  Quitensium,  2500  m.,  Jameson 
53  (NY),  55  (GH)  ;  between  Baeza  and  Papallacta,  2000  m.,  Mexia  7339  (GH, 
US)  ;  Nanegal  District,  Couthoiiy  in  1855  (GH) ;  west  declivity  of  Pichincha, 
Jameson  7  (NY).  Tungurahua:  Valley  of  Pastaza  River,  between  Baiios  and 
Cashurco,  1300-1800  m.,  Hitchcock  21781  (GH,  NY,  US)  ;  Vulcan  Tungur- 
agua,  LeJimann  4995  (F).  A  collection  from  Huigra,  by  Rose  and  Rose,  no. 
22477  (US)  is  near  this  but  the  leaves  are  thin. 

(51)  Fuchsia  verrucosa  HartAveg 

(Plate8,  fig.  47) 

Fuchsia  verrucosa  Hartweg,  in  Benth.,  PI.  Hartweg.,  178,  1845. 
F.  perbrevis  Johnston,  Contr.  Gray  Herb.,  75:30, 1925. 

Shrub  1-2  m.  high,  young  twigs  subglabrous,  slightly  red,  2-4  mm.  thick ; 
leaves  opposite  or  ternate,  thick  and  firm,  elliptic-ovate  to  ovate-oblong,  acute 
to  acuminate  at  base,  acuminate  at  apex,  subentire  to  somewhat  serrulate, 
quite  glabrous  above,  paler  and  subglabrous  beneath,  or  puberulent  on  the 


Vol.  XXV]  MUNZ:  THE  GENUS  FUCHSIA  59 

veins;  veins  very  prominent  beneath,  the  prineipla  lateral  ones  about  15  on 
each  side  of  the  midrib,  submarginal  prominent ;  blades  4-12  cm.  long,  2.5- 
5.5  cm.  vs^ide ;  petioles  2-10  mm.  long ;  stipules  more  or  less  connate,  reflexed, 
about  2  mm.  long;  flowers  few,  solitary,  axillary;  pedicels  firm,  glabrous, 
1-2.5  cm.  long,  recurved  at  anthesis;  ovary  subglabrous,  10-12  mm.  long, 
about  2  mm.  wide,  4-angled ;  hypanthium  obconic,  3-5  mm.  long,  subglabrous 
without  and  within ;  sepals  8-10  mm.  long,  2-3  mm.  wide,  lance-oblong,  diver- 
gent, acutish,  bright  red;  petals  red,  obovate,  8-9  mm.  long,  4-5  mm.  wide; 
stamens  3-4  mm.  long;  anthers  1.5  mm.  long;  style  glabrous,  7-9  mm.  long; 
stigma  1.5  mm.  thick;  fruit  4-angled,  15-18  mm.  long,  3.5-4  mm.  wide. 

Type  locality,  between  "Paramo  de  San  Fortunato  et  pagum  Fusagasuga, 
prov.  Bogota,"  Colombia.  The  species  seems  to  be  nearest  to  F.  scabriuscula , 
with  which  it  agrees  in  its  thick  veiny  leaves,  but  is  remarkable,  as  has  been 
pointed  out  both  by  Bentham  and  Johnston  in  their  descriptions,  for  its  very 
short  hypnathium.  I  have  seen  the  following  collections,  all  from  COLOM- 
BIA: without  definite  locality,  Muiis  2077  (US),  2047  (US),  1361  (US). 
Dept.  Santander  del  Norte :  road  Pamplona  to  Toledo,  2800-3000  m.,  KilliiD 
and  Smith  19976  (GH,  NY,  US),  19831  (F,  GH,  NY,  US).  Cundimarca :  be- 
tween Paramo  de  San  Fortunato  and  Fusagasuga,  prov.  Bogota,  Hartweg 
991,  type  no.  verrucosa,  Berlin  specimen  (photo  F,  POM),  Geneva  specimen 
(leaf  F)  ;  Flora  Neo-granadina-Bogotana,  Ultra  Abate  (ultra  Ubate  ?), 
Holton  892,  type  perhrevis  (GH).  Huila  :  Comisaria  del  Caqueta  :  Cordillera 
Oriental,  en  Gabinete,  2300-2450  m.,  Cuatrecasas  8465  (US). 

(52)  Fuchsia  sanctae-rosae  0.  Kuntze 

(Plate  9,  fig.  48) 

Fuchsia  sanctae-rosae  O.  Kuntze,  Eev.  Gen.,  Ill,  pt.  2 :98, 1898. 

F.  holiviana  Britton,  Bull.  Torrey  Bot.  Club,  17  :  214,  1890,  not  Carr.,  1876. 

F.  Brittonii  Johnston,  Contr.  Gray  Herb.,  75  :39,  1925,  new  name  for  'boliviana  Britton, 

F.  Weberbeueri  Krause,  Fedde  Rep.  Nov.  Sp.,  1:170,  1905;  based  on  Weberbauer  661,  of 

Avhicli  I  have  seen  a  photograph, 
F.  filipes  RusBY,  Mem.  N.  Y.  Bot.  Gard.,  7:317,  1927;  based  on  White  232,  which  I  have  seen. 

From  suffrutescent  or  almost  herbaceous  and  3-5  dm.  tall,  to  shrubby  and 
2-3  m.  high,  rather  simple  to  many -branched,  erect  to  semidecumbent ;  young 
shoots  mostly  glabrous,  sometimes  somewhat  pilose,  purplish  red,  1.5-4  mm. 
thick;  leaves  mostly  in  whorls  of  3  or  4,  firm,  elliptic-lanceolate  to  almost 
-obovate,  obtuse  to  acute  at  base,  acute  to  acuminate  at  apex,  sub  entire  to 
minutely  serrulate,  deep  green  and  glabrous  above,  lighter  and  glabrous  be- 
neath except  usually  somewhat  pilose  along  midrib,  principal  lateral  veins 
8-12  on  each  side  of  midrib,  submarginal  frequently  not  very  evident;  blades 
3-10  cm.  long,  1.5-4  cm.  wide ;  petioles  5-20  mm.  long,  usually  purplish  red 
and  glabrous,  sometimes  appressed-pubescent ;  stipules  subulate,  1-2  mm. 
long,  sometimes  reflexed,  sometimes  connate,  deciduous ;  flowers  rather  nu- 
merous, but  solitary  in  upper  axils  of  somewhat  reduced  leaves,  hence  more  or 


60  CALIFOENIA  ACADEMY  OF  SCIENCES  [Proc.  4th  Ser. 

less  racemose;  pedicels  very  slender,  1-2.5  cm.  long,  mostly  glabrous;  ovary 
narrow  ellipsoid,  glabrous  to  puberuleut,  4-6  mm.  long ;  hypanthium  deep  to 
bright  red,  13-18  (22)  mm.  long,  2-2.5  mm.  wide  at  base,  then  slightly  nar- 
rowed and  then  gradually  ampliate  until  about  4  mm.  wide  at  summit,  gla- 
brous without,  pilose  within  the  lower  half ;  sepals  scarlet,  lance-oblong,  8-10 
mm.  long,  2.5-3  mm.  wide,  glabrous  or  sometimes  pilose,  acutish ;  petals 
orange-red  to  scarlet,  oblong-obovate,  6-8  mm.  long,  3-6  mm.  wide,  obtuse  to 
acutish,  sometimes  almost  3-toothed;  stamens  7-9  and  5-7  mm.  long;  anthers 
about  1.5  mm.  long;  style  mostly  glabrous,  extending  about  to  the  sepal-tips; 
stigma  red,  about  1  mm.  wide ;  berry  varying  from  globose-ellipsoid  to  oblong- 
ellipsoid,  6-10  mm.  long,  4-5  mm.  wide. 

Type  locality,  Santa  Rosa,  Bolivia  (there  are  several  towns  of  this  name  in 
Bolivia).  I  have  not  seen  the  type.  The  species  is  variable  in  every  way,  but  I 
cannot  find  any  constant  characters  for  division.  It  comes  between  the  race- 
mose and  axillary-flowered  groups  with  short  hypanthium,  so  seems  to  stand 
between  the  species  I  have  just  treated  and  those  like  Osgoodii,  Andrei,  etc., 
which  follow.  It  ranges  at  altitudes  between  2000  and  3000  m.,  in  southern 
Peru  and  Bolivia.  Representative  material,  PERU :  Dept.  Cuzco :  "Pillahu- 
ata,"  Cerro  de  Cusilluyoc,  Pennell  13997  (F,  GH,  NY,  US),  13956  in  part 
(GH);  prov.  Paucartambo,  valley  of  Rio  Tambomayo,  West  7091  (GH)  ;• 
Macchu-Picchu,  Munz  15548  (POM),  West  6414  (GH),  3Iexia  8078  (F,  GH, 
US),  Balls  6802  (US).  Puno  :  Sandia,  Weherhauer  661,  type  Wederhaueri  at 
Berlin  (photo  F,  POM).  BOLIVIA :  Santa  Rosa,  as  serratifolia,  0.  Kuntze  in 
1892  (F,  GH).  La  Paz:  Haciendo  Simaco,  sobre  el  camina  a  Tipuani,  Biich- 
tien  832  (F,  POM),  5508  (GH,  NY,  US)  ;  Pongo,  Pulcheri,  White  232,  type 
no.  filipes  (GH,  US)  ;  Prov.  Larecaja :  near  Ananea :  Machacamarca,  Mandon 
624  (F,  GH,  NY)  ;  Prov.  No.  Yungas,  between  Unduavi  and  Chirca,  Eyerdam 
25388  (F) ;  Yungas,  Bmig  675  (F,  GH,  NY,  US),  Rushij  1812  (F,  NY,  US), 
1813  type  no.  holiviana,  Brittoni  (NY,  US).  Cochabamba :  prov.  Chapare, 
Steinhach  9227  (GH)  ;  Incachaca,  Cdrde^ias  665  (US). 

(53)  Fuchsia  Osgoodii  Macbride 

(Plate9,  fig.  49) 

Fuchsia  Osgoodii  Macbride,  Field  Mug.  Nat.  Hist.,  Botany,  13,  part  4,  no.  1 :559, 1941. 
F.  ovalis  E.  and  P.  var.  aherrans  Macbride,  I.e.,  560. 

Shrub,  up  to  4  m.  tall ;  twigs  dark,  subglabrous,  1-2.5  mm.  thick ;  leaves 
opposite,  firm,  elliptic-lanceolate,  acute  to  acuminate  at  both  ends,  quite  entire, 
deep  green,  glabrous  above,  paler  and  glabrous  beneath  except  minutely 
puberulent  or  short-pilose  on  veins,  with  10-12  principal  lateral  veins  on 
each  side  of  midrib,  submarginal  present;  leaf-blades  5-8  (12)  cm.  long, 
1.5-3  (6)  cm.  wide;  petioles  glabrous,  8-10  mm.  long;  flowers  few,  in  short 
congested  racemes;  bracts  1-2  cm.  long;  pedicels  filiform,  1.5-2.5  cm.  long, 
glabrous ;  ovary  linear-ellipsoid,  ca.  5  mm.  long ;  hypanthium  23-24  mm.  long, 
dark  red,  somewhat  enlarged  at  base,  where  1.5.2  mm.  wide,  then  1  mm.  wide 


Vol.  XXV]  MUNZ :  THE  GENUS  FUCHSIA  61 

for  one-third  its  length,  then  gradually  ampliate  until  4-5  mm.  wide  at  sum- 
mit, glabrous  without,  villous  within  lower  portion  ;  sepals  red,  oblong-ovate, 
10-11  mm.  long,  5-6  mm  wide,  glabrous,  subulate  for  1-1.5  mm.  at  apex; 
petals  red,  oblong,  about  10  mm.  long,  4  mm.  wide,  abruptly  acute;  stamens  8 
and  6  mm.  long ;  anthers  1.6  mm.  long;  style  glabrous,  equalling  petals ;  stigma 
4-lobed,  1.5  mm.  wide;  berry  oblong,  at  least  12  mm.  long  and  4  mm.  wide. 
Type  locality,  PERU :  Dept.  Libertad :  Ucheo,  the  type  Osgood  and  Ander- 
son 47  (F),  From  Cajamarca:  prov.  Cutervo :  Arenales,  Stork  and  Horton 
10155  (F)  comes  the  type  of  ovalis  var.  aherrans,  which  has  some  lower  and 
larger  leaves  than  the  type  of  Osgoodii.  They  are  more  pilose  on  the  veins 
beneath  the  leaves,  but  the  upper  smaller  leaves  have  the  same  puberulence 
as  in  Osgoodii.  F.  Osgoodii  is  near  F.  sanctae-rosae  in  hypanthium,  foliage, 
etc.,  but  has  more  definitely  terminal  inflorescence. 

(54)  Fuchsia  AndreiJohnston 
(Plate9,  fig.  50) 

Fuchsia  Andrei  Johnston,  Contr.  Gray  Herb.,  75  :31,  1925. 

Shrubby,  the  younger  twigs  more  or  less  purplish  red,  fairly  coarse,  sub- 
glabrous  except  toward  the  minutely  pubescent  tips,  3-8  mm.  thick ;  leaves 
opposite  or  ternate,  oblong-elliptic  to  elliptic-ob ovate,  mostly  obtuse  at  base, 
abruptly  acuminate  at  apex,  membranaceous,  subentire,  sparsely  ciliolate, 
green  and  subglabrous  above,  paler  beneath  and  puberulent  on  veins,  with 
about  12  principal  lateral  veins  on  each  side  of  midrib  and  with  evident  sub- 
marginal  vein;  blades  7-15  cm.  long,  3-7  (9)  cm.  wide;  petioles  subglabrous, 
1-2.5  cm.  long;  stipules  deltoid,  ca.  1  mm.  long,  deciduous;  inflorescence  a 
short  lateral  or  terminal  raceme;  bracts  lanceolate,  3-10  mm.  long,  somewhat 
pubescent;  pedicels  slender,  somewhat  pubescent,  10-15  mm.  long;  ovary 
oblong,  subglabrous,  about  5  mm.  long ;  hypanthium  2-3  cm.  long,  somewhat 
enlarged  at  base,  then  1  mm.  wide  for  about  one-third  its  length,  then  gradu- 
ally ampliate  until  4-4.5  mm.  Avide  at  apex,  subglabrous  without,  pilose  with- 
in lower  half ;  sepals  oblong-lanceolate,  7-9  mm.  long,  divergent,  subulate  for 
1.5  mm.  at  tips,  almost  glabrous;  petals  lance-oblong,  9-10  mm.  long,  2.5-3 
mm.  wide,  obtuse ;  stamens  about  8  and  6  mm.  long ;  style  pilose  in  basal  half, 
exceeding  sepals  by  3^  mm. ;  stigma  globose,  4-lobed,  ca.  1  mm.  thick ;  berry 
oblong,  6  mm.  long  and  3  mm.  wide. 

Type  locality,  Rio  de  Huannamaca,  probably  southern  Ecuador.  I  have 
seen  two  collections  :  Rio  de  Huannamaca,  Andre  K820,  type  no.  (F)  ;  Tambo 
de  Savanilla,  Andre  K  818  (F,  GH,  NY).  The  leaves  are  large  and  glabrous, 
the  flowers  in  terminal  racemes. 

(55)  Fuchsia  Lehmannii  Munz,  new  species 
(PlateQ,  fig.  51) 

Soft-wooded  shrub  up  to  2  m.  high,  the  younger  twigs  dark  red,  mostly 
glabrous,  1-3  mm.  thick;  leaves  opposite  or  ternate,  elliptic-lanceolate  or 


62  CALIFORNIA  ACADEMY  OF  SCIENCES  [Proc.  4th  Ser. 

-oblanceolate,  acuminate  or  acute  at  both  ends,  rigid,  subentire,  glabrous,  and 
dark  green  above,  beneath  paler  and  glabrous  except  for  some  short  appressed 
pubescence  on  the  reddish  veins,  the  principal  lateral  veins  8-12  on  each  side 
of  midrib  ;  leaf -blades  mostly  5-10  (12)  cm.  long,  1.5-3  (4)  cm.  wide;  petioles 
glabrous  to  somewhat  strigulose,  5-15  mm.  long;  stipules  lance-subulate,  1 
mm.  or  so  long,  deciduous;  flowers  numerous,  crowded  in  short  lateral  and 
terminal  racemes  arranged  paniculately;  bracts  lance-linear,  reflexed,  some- 
what pubescent,  1-2  cm.  long;  pedicels  slender,  pubescent,  8-18  mm.  long; 
ovary  narrow-fusiform,  more  or  less  strigulose,  3-5  mm.  long ;  hypanthium 
red,  24-30  mm.  long,  1  mm.  wide  at  base,  then  narrowed  for  one-third  its 
length,  then  gradually  ampliate  until  3.5-5  mm.  wide  at  apex,  mostly  glabrous 
without  and  pubescent  within  the  narrow  basal  portion;  sepals  scarlet,  re- 
flexed-spreading,  linear-lanceolate,  glabrous,  9-10  mm.  long,  2.5-3  mm.  wide, 
subulate  for  1-2  mm.  at  apex;  petals  scarlet,  oblong-lanceolate,  about  1  cm. 
long,  2-2.5  mm.  wide,  obtusish ;  stamens  7  and  5.5  mm.  long ;  anthers  1  mm. 
long;  style  glabrous,  slightly  exceeding  petals;  stigma  1-1.5  mm.  wide;  fruit 
ellipsoid,  5-6  mm.  long,  3-4  mm.  wide. 

Frutex,  1-2  m.  altus ;  ramulis  rubris,  subglabris,  1-3  mm.  crassis ;  f  oliis  oppositis  vel 
ternatis,  elliptico-lanceolatis  vel  -oblanceolatis,  acuminatis  vel  acutis,  rigidis,  subintegris,  in 
superficie  superiore  glabris,  inferiore  glabris  vel  in  venis  puberulentis,  laminis  5-10  (12)  ■ 
cm.  longis,  1.5-3  (4)  cm.  latis;  petiolis  glabris  vel  substrigulosis,  5-15  mm.  longis;  floribus 
numerosis,  in  racemis  paniculatis;  bracteis  lanceo-linearibus,  reflexis,  pubescentibus,  1-2 
cm.  longis;  pedicelis  tenuibus,  pubescentibus,  8-18  mm.  longis;  ovario  angusto-fusiforme, 
strigulose,  3-5  mm.  longo ;  hypanthio  rubro,  24-30  mm.  longo,  base  1  mm.  lato,  inde  angus- 
tiore,  inde  ampliato  et  apice  3.5-5  mm.  lato,  externe  subglabro,  interne  pubescente;  sepalis 
coceineis,  reflexo-divergentibus,  lineare-lanceolatis,  glabris,  9-10  mm.  longis,  cum  apicibus 
subulatis  1.5-2  mm,  longis;  petalis  coceineis,  oblongo-lanceolatis,  1  cm.  longis,  2-2.5  mm. 
latis,  oBtusis ;  staminibus  7  et  5.5  mm.  longis ;  antheris  1  mm.  longis ;  stylo  glabro,  petala 
excedentibus ;  stigmate  1-1.5  mm.  lato;  bacca  ellipsoidea,  5-6  mm.  longa,  3-4  mm.  lata. 

Type:  from  Loja,  Ecuador:  east  Andes  of  Sigsig,  1600-1800  m.,  Lehmann 
5498  (equal  Pearce's  235),  type  Field  Museum  550994;  isotypes  GH,  US.  A 
second  collection  also  from  Loja :  Rio  Savanilla,  east  Andes  of  Loja,  800-1300 
m.,  Lehmann  7858  (P,  US) .  The  proposed  species  is  near  F.  Osgoodii  in  leaves 
and  flowers,  but  sepals  are  more  narrow  and  berry  shorter ;  near  F.  Hartwegii 
in  flower-shape,  but  glabrous  and  with  more  pointed  leaves ;  near  F.  Andrei, 
but  with  glabrous  style  and  narrow  leaves. 

(56)  Fuchsia  putumayensis  Munz,  new  species 
(Plate  8,  fig.  52) 

Shrub  or  tree,  younger  twigs  guite  glabrous,  with  some  purplish  red  about 
the  nodes,  2-4  mm.  thick ;  leaves  opposite,  lance-ovate,  rather  membranaceous, 
acute  to  obtuse  at  base,  acute  to  acuminate  at  apex,  subentire,  bright  green 
and  glabrous  above,  lighter  and  glabrous  beneath  except  for  fine  appressed 
puberulence  along  midrib  and  veins,  the  principal  lateral  veins  about  15, 
rather  prominent  beneath,  the  submarginal  inconspicuous;  blades  6-10  cm. 


Vol.  XXV]  MUNZ:  TEE  GENUS  FUCHSIA  63 

long,  2.5-4  cm.  wide ;  petioles  5-12  mm.  long,  purplish  red,  somewhat  strigu- 
lose;  stipules  not  seen;  flowers  several,  in  very  short  compact  terminal  and 
lateral  almost  glabrous  racemes ;  bracts  reflexed,  linear-lanceolate,  3-10  mm. 
long;  pedicels  filiform,  quite  glabrous,  1-2.5  cm.  long;  ovary  linear-fusiform, 
glabrous,  3.5-4  mm.  long,  red ;  hypanthium  bright  red,  14-20  mm.  long,  ca.  1 
mm.  wide  at  base,  then  somewhat  narrowed  for  one-third  its  length,  then 
gradually  ampliate  until  2.5-3.5  mm.  wide  at  summit,  glabrous  without,  pilose 
within  lower  portion ;  sepals  scarlet,  divergent,  7-9  mm.  long,  3  mm.  wide  at 
base,  subulate  at  apex  for  1-1.5  mm.,  glabrous;  petals  scarlet,  oblong,  6  mm. 
long,  about  3  mm.  wide,  rounded-obtuse  at  apex ;  stamens  5  and  3.5  mm,  long ; 
style  glabrous,  barely  equalling  petals;  stigma  ca.  1  mm.  thick;  fniit  oblong- 
ellipsoid,  about  2.5  cm.  long. 

Frutex  vel  arbor ;  ramulis  glabris,  subrubris,  2-4  mm.  diametro ;  f oliis  oppositis,  lanceo- 
lato-ovatis,  membranaceis,  basi  acutis  vel  obtusis,  apice  acutis  vel  acuminatis,  subintegris, 
in  superficie  superiore  glabris,  inferiore  in  venis  strigulosis;  laminis  6-10  cm.  longis,  2.5-4 
cm.  latis ;  petiolis  5-12  mm.  longis,  substrigulosis ;  racemis  brevibus,  compactis,  terminalibus 
et  lateralibus,  subglabris ;  bracteis  reflexis,  lineari-lanceolatis,  3-10  mm.  longis ;  pedicellis 
glabris,  1-2.5  cm.  longis,  ovario  lineare-fusiforme,  glabro,  3.5-4  mm,  longo,  rubro;  hypan- 
thio  rubro,  14^20  mm,  longo,  basi  1  mm,  lato,  inde  angustiore,  inde  ampliato  et  apice 
2.5-3.5  mm.  lato,  externe  glabro,  interne  piloso ;  sepalis  coccineis,  divergentibus,  7-9  mm. 
loiigis,  basi  1  mm.  latis,  glabris ;  petalis  coccineis,  oblongis,  6  mm.  longis,  3  mm.  latis,  apice 
rotundatis  vel  subobtusis;  staminibus  5  et  3.5  mm.  latis;  style  glabro,  petala  non  excedent- 
ibus ;  stigmate  1  mm.  diametro ;  bacca  oblongo-ellipsoidea,  circa  25  mm,  longa. 

Tyije:  Comissaria  del  Putumayo:  Cocoa,  Colombia,  May  23,  1935,  H. 
Garcia  B.  4639,  U.  S.  Nat.  Herb.  1593482.  This  is  rather  a  fragmentary  speci- 
men and  had  almost  finished  flowering,  so  that  the  last  late  flowers  may  be 
somewhat  small  for  the  species.  A  second  specimen,  Plailada  de  Minchoy, 
Cuatrecasas  11436  (US).  It  seems  near  F.  Lehmannii  in  having  lateral  and 
terminal  compact  racemes,  being  glabrous,  having  subulate  sepal-tips  and 
glabrous  style.  But  it  has  more  veins,  too  long  a  berry,  too  long  pedicels  and 
too  short  a  hypanthium, 

(57)  Fuchsia  Hartwegii  Bentham 

(Plate9,  fig,  53) 
Fuchsia  Hartwegii  Bentham,  PI.  Hartw,,  179,  1945. 

Shrub  or  small  tree,  the  younger  branches  slender,  more  or  less  purplish- 
red,  pubescent,  sometimes  also  with  some  longer  hairs ;  leaves  in  3's  or  4's, 
elliptic-oblong,  rounded  to  obtuse  at  base,  obtuse  to  abruptly  acuminate  at 
apex,  subentire  or  inconspicuously  and  remotely  denticulate,  membranaceous, 
darker  green  and  appressed-pilose  above,  paler  and  pilose  beneath  especially 
on  veins,  principal  lateral  veins  14-16  on  each  side  of  midrib,  submarginal 
vein  present;  leaf-blades  mostly  4-10  cm.  long,  2-4  (5.5)  cm.  wide;  petioles 
appressed-pilose,  slender,  5-35  mm.  long;  stipules  subulate-deltoid,  1.5  mm. 
long,  deciduous ;  flowers  numerous  in  terminal  pyramidal  leafy -bracted, 
pendulous  panicle  5-15  cm.  long;  bracts  broadly  elliptic,  8-25  mm.  long; 


64  CALIFOENIA  ACADEMY  OF  SCIENCES  [Proc.  4th  Ser. 

pedicels  slender,  pilose,  5-10  mm.  long;  ovary  ©void,  appressed-pubeseent, 
about  3  mm.  long;  hypanthium  red,  tubular,  sparsely  pubescent  without, 
pilose  within  lower  half,  1.4-2  cm.  long,  1.5  mm.  wide  at  base,  then  somewhat 
narrowed,  then  gradually  widened  until  4—5  mm.  wide  at  apex ;  sepals  orange- 
red  or  scarlet,  linear-lanceolate,  spreading-divaricate,  8-9  mm.  long,  barely 
2  mm.  wide,  acuminate,  with  subulate  tips  less  than  1  mm.  long,  pubescent  on 
backs;  petals  red,  linear-lanceolate,  8-9  mm.  long,  2  mm.  wide,  acute,  gla- 
brous; stamens  10  and  8  mm.  long;  anthers  1.5-2  mm  long;  style  slender, 
glabrous  or  somewhat  pubescent  in  lower  part,  equalling  or  exceeding  sepals 
by  3^  mm ;  stigma  globose,  about  1  mm.  broad ;  fruit  round-ellipsoid,  up  to 
7  mm.  long. 

Type  locality,  Colombia :  "In  sylvis  prope  Pitayo  et  Huambia."  A  species 
of  southern  Colombia,  where  it  grows  at  between  2000  and  3000  m.,  it  seems 
near  F.  Lelimanmi,  F.  ovalis,  etc.,  in  its  general  features:  terminal  inflores- 
cence, short  hypanthium,  etc.,  but  has  its  own  characteristic  pubescence,  leaf- 
size,  pedicel-length,  and  the  like.  Representative  material  is  as  follows, 
COLOMBIA:  Caldas:  Dawe  741  (NY,  US).  Huila:  "Balsillas"  in  Rio  Bal- 
sillas,  Rushy  and  Pennell  789  (NY,  US).  El  Cauca:  near  Pitayo  and  Huam- 
bia,ffarfwegr  994,  type  no.  (fragment  F),  (photo  Berlin  specimen  F,  POM)  ; 
central  Andes  of  Popayan,  Lehmann  5613  (F)  ;  Coconuco,  Killip  6829  (GH, 
NY,  US)  ;  near  Purace,  Lelimaym  3503  (F,  US) ;  San  Isidro,  Purace,  Pennell 
and  Killip  6460  (GH)  ;  Canaan,  Mt.  Purace,  Pennell  and  Killip  6502  (GH)  ; 
del  paramo  a  Purace,  matorrales  en  Chiquin,  Perez  Arheldez  and  Cuatrecasas 

5973  (US). 

(58)  Fuchsia  ovalis  Ruiz  and  Pavon 

(Plate  10,  fig.  54) 

Fuchsia  ovalis  Euiz  and  Pav6n,  F1.  Peruv.,  3 :87,  pi.  324,  fig.  a,  1802. 

F.  polyanthella  Johnston,  Contr.  Gray  Herb.,  75:36,  1925;  I  have  seen  type  material  of 
both  this  and  ovalis. 

Erect  or  somewhat  scandent,  semishrubby,  up  to  1  m.  tall,  rather  sparsely 
branched,  the  younger  branches  3-6  mm.  thick,  more  or  less  dark,  pilose  to 
almost  glabrous ;  leaves  opposite  or  ternate,  membranaceous,  broadly  elliptic 
to  oblong-ovate  or  obovate,  obtuse  to  acute  at  base,  abruptly  acuminate  at 
apex,  entire  to  inconspicuously  denticulate,  darker  green  and  hirsute-strigose 
above,  paler  and  hirsute  beneath  especially  on  veins,  with  about  15-17  prin- 
cipal lateral  veins  on  each  side  of  midrib  and  with  submarginal  vein ;  blades 
mostly  8-15  cm.  long,  5-8  cm.  wide;  petioles  pilose,  2-5  cm.  long;  stipules 
subulate-deltoid,  2-5  mm.  long,  quite  persistent;  flowers  in  mostly  axillary 
racemes  up  to  1  dm.  long ;  bracts  lanceolate,  1-1.5  cm.  long,  pilose ;  pedicels 
1-1.5  cm.  long,  glabrous  to  pilose ;  ovary  ellipsoid,  5-7  mm.  long,  glabrous  to 
sparingly  pilose;  hypanthium  13-15  mm.  long,  scarlet,  1.5-2  mm.  wide  at 
base,  then  narrower  until  gradually  widened  to  summit  where  3.5-4.5  mm. 
wide,  glabrous  to  sparingly  pilose  without,  villous  within  lower  half ;  sepals 
lanceolate,  acute,  somewhat  divergent,  9-11  mm.  long,  acute,  not  subulate, 


1 


Vol.  XXV]  MUNZ :  THE.  GENUS  FUCHSIA  65 

glabrous  to  somewhat  pilose ;  petals  obovate-oblong,  obtusish,  7-8  mm.  long ; 
stamens  5  and  4  mm.  long,  filaments  violet;  anthers  1  mm.  long ;  style  glabrous, 
extending  to  sepal-tips ;  stigma  subglobose,  about  1  mm.  broad ;  berry  oblong, 
red-purple,  somewhat  4-angled,  about  1  cm.  long,  4  mm.  wide. 

Type  locality,  Muna,  Huanuco,  Peru.  The  species  is  characterized  by  its 
large  leaves  which  are  pilose  on  both  surfaces,  by  its  lateral  racemes,  lance- 
olate bracts,  and  hypanthium  15  mm.  long.  I  have  seen  the  following  collec- 
tions, PERU:  the  Ruiz  and  Pavon  collection  from  Madrid  Herb.  (F),  from 
Geneva  Herb,  (photo  F,  POM).  Huanuco:  Huallaga  und  Muiia,  2800-2900 
m.,  Weherbauer  6721  (F)  ;  Muiia,  trail  to  Tambo  de  Vaca,  2600  m.,  Machride 
4290,  type  number  polyanthella  (F,  GH). 

(59)  Fuchsia  asperifolia  Krause 

Fuchsia  asperifolia  Krause,  Fedde  Rep.  Nov.  Sp.,  1 :169, 1905. 

Low  shrub,  up  to  5  dm.  tall,  simple  or  few-branched,  the  young  twigs  dark, 
densely  pilose;  leaves  opposite  or  ternate,  membranaceous,  broadly  elliptic 
to  elliptic-obovate,  acute  at  both  ends,  subentire,  hirsute  on  both  surfaces, 
especially  on  veins  beneath,  with  about  15  principal  lateral  veins  on  each  side 
of  midrib,  submarginal  vein  present;  blades  4-10  cm.  long,  2-5  cm.  wide; 
petioles  hirsute,  2-5  cm.  long ;  stipules  subulate,  deciduous ;  flowers  in  dense 
usually  terminal  hirsute  raceme ;  bracts  apparently  lanceolate ;  pedicels  ap- 
parently scarcely  5  mm.  long;  ovary  about  6  mm.  long,  pilose;  hypanthium 
dark  red,  2-2.5  cm.  long,  slightly  bulbous  at  base,  then  narrowed  and  gradu- 
ally widened  until  5-6  mm.  wide  at  apex,  hirsute  without,  pilose  within; 
sepals  ovate,  acuminate,  7-8  mm.  long ;  petals  scarlet,  narrowly  obovate,  acut- 
ish  at  apex,  8-9  mm.  long;  stamens  included;  style  equaling  sepals;  stigma 
subglobose,  4-lobed ;  fruit  not  seen. 

Type  locality,  Peru :  Amazonas  :  between  Tambos  Bagazan  and  Almirante, 
east  of  Chachapoyas,  at  2700  m. ;  type  Weherhauer  4445  at  Berlin  (photo  F, 
POM) .  Having  seen  only  a  photograph,  I  do  not  have  many  details;  the  species 
seems  near  to  F.  ovalis,  but  with  more  terminal  inflorescence,  is  more  hirsute 
throughout  and  with  shorter  pedicels, 

(60)  Fuchsia  pilosa  Fielding  and  Gardner 

Fuchsia  pilosa  Field,  and  Gardner,  Sert.  PI.,  1,  pi.  27,  1844;  Macbride,  Field  Mus.  Nat. 
Hist.,  Bot.  13,  pt.  4,  no.  1 :561,  1941. 

Shrub,  thinly  white-pilose  on  young  branches,  leaves,  petioles,  hypanthia, 
etc. ;  leaves  ternate,  oblong-lanceolate,  acute  at  base,  acuminate  at  apex,  min- 
utely denticulate,  the  blades  6-9  cm.  long,  1.8-2  cm.  wide,  one  of  each  whorl 
somewhat  larger  than  the  other  two ;  two  of  petioles  at  each  node  5-7  mm. 
long,  the  third  at  least  2  cm.  long ;  flowers  in  terminal  racemes  which  become 
1  or  more  dm.  long;  bracts  like  reduced  leaves  and  1-2.5  cm.  long;  pedicels  a 
few  mm.  long ;  flowers  scarlet,  crowded ;  hypanthium  apparently  about  22 
mm.  long;  sepals  oblong,  8-9  mm.  long,  prominently  subulate  at  apex;  petals 


66  CALIFORNIA  ACADEMY  OF  SCIENCES  [Proc.  4th  Ser. 

obtuse,  wider  than  sepals,  almost  equaling  them ;  style  included ;  fruit  oblong, 
4-angled,  less  than  1  cm.  long. 

Type  locality,  Taulia,  Amazonas,  Peru:  type  Mathews  1482,  which  I  have 
not  seen.  Macbride  separates  this  species  on  the  basis  of  its  unequal  petioles 
and  leaves,  one  at  each  node  being  larger  than  the  other  two.  It  would  seem 
from  the  description  to  be  near  to  F.  asperifoUa  and  the  two  are  perhaps  dis- 
tinct from  each  other  because  of  the  unequal  leaves. 

(61)  Fuchsia  glaberrima  Johnston 
■  (Plate  10,  fig.  55) 
Fuchsia  glaberrima  Johnston,  Contr.  Gray  Herb.,  75:32,  1925. 

Shrub ;  younger  twigs  slightly  reddish,  3-5  mm.  thick,  not  branched,  gla- 
brous except  for  very  minute  scarce  appressed  puberulence  near  tips ;  leaves 
opposite,  firm,  elliptic-oblanceolate,  tinged  reddish  purple,  obtuse  at  base, 
acuminate  to  acute  at  apex,  entire,  quite  glabrous,  with  13-15  principal  lateral 
veins  on  each  side  of  the  midrib  and  with  inconspicuous  marginal  vein ;  blades 
10-20  cm.  long,  4-8  cm.  wide;  petioles  up  to  5  (8)  mm.  long;  stipules  connate, 
often  reflexed,  quite  persistent,  2-2.5  mm.  long ;  inflorescence  a  sliort  terminal 
raceme ;  bracts  lanceolate,  1-3.5  cm.  long ;  pedicels  stout,  5-7  mm.  long,  min- 
utely appressed  puberulent;  ovary  fusiform,  6-9  mm.  long;  hypanthium ' 
bright  red,  2.5-3  cm.  long,  about  2  mm.  wide  at  base,  then  very  little  con- 
tracted, then  widened  until  6-7  mm.  wide  at  apex,  almost  glabrous  without, 
densely  pubescent  throughout  within ;  sepals  scarlet,  obong,  9-11  mm.  ong, 
4-5  mm.  wide,  obtusish ;  petals  scarlet,  broadly  obovate,  9  mm.  long,  rounded 
at  apex;  stamens  5  and  3.5  mm.  long;  anthers  1.5  mm.  long;  style  densely 
pubescent  much  of  its  length,  almost  equaling  sepals ;  stigma  4-lobed,  2.5  mm. 
thick ;  fruit  not  seen. 

Type,  ECUADOR:  Prov.  Tungurahua:  Valley  of  Pastaza  River,  between 
Banos  and  Cashurco,  1300-1800  m.;  type  number  Hitchcock  21750  (GH,  NY, 
US).  Diels,  Bibl.  Bot.  116  :119,  1937,  refers  here  a  Raimondi  collection  from 
Amazonas,  Peru.  This  species  is  like  F.  spectahilis  in  its  broad  petals,  but  has 
a  shorter  hypanthium;  like  F.  asperifoUa  in  general  appearance,  but  more 
glabrous  and  with  more  sessile  leaves. 

(62)  Fuchsia  macrophylla  Johnston 
(Plate  10,  fig.  56) 
Fuchsia  macrophylla  Johnston,  Contr.  Gray.  Herb.,  75 :  35, 1925. 

Shrub  up  to  3  m.  tall,  openly  branched,  the  younger  twigs  3-6  mm.  thick, 
with  some  dark  red  or  purplish  tinge,  subglabrous  to  finely  strigulose-puberu- 
lent  toward  the  tips ;  leaves  opposite,  thick-membranaceous,  elliptic,  gradually 
attenuate  and  subacuminate  at  base,  somcAvhat  abruptly  acuminate  at  apex, 
subentire,  dark  green  above,  paler  beneath,  subglabrous  on  both  surfaces 
except  for  some  minute  puberulence  along  the  veins,  principal  lateral  veins 


Vol.  XXV]  MUNZ :  TEE  GENUS  FUCHSIA  67 

14-18  on  each  side  of  midrib,  submarginal  well  developed;  blades  7-15  (18) 
cm.  long,  3-5  (6.5)  cm.  wide;  petioles  1-4  cm.  long;  stipules  dark,  lance- 
subiilate,  1.5-2  mm.  long,  deciduous ;  inflorescence  mostly  of  short  axillary 
sometimes  terminal  racemes,  often  only  a  few  cm.  long  and  with  few  flowers ; 
bracts  lanceolate  or  wider,  5-15  mm.  long ;  pedicels  slender,  minutely  strigu- 
lose,  5-20  mm.  long;  ovary  ellipsoid,  4—5  mm.  long ;  hypanthium  scarlet,  18-23 
mm.  long,  1.5-2  mm.  wide  at  base,  then  narrowed,  then  gradually  ampliate 
until  4-5  mm.  wide  at  apex,  finely  appressed-puberulent  without,  pilose 
within ;  sepals  divergent,  oblong-lanceolate,  red  with  green  tips,  7-8  mm.  long, 
3  mm.  wide,  appressed-puberulent  on  backs,  apices  subulate  for  0.5  mm. ; 
petals  bright  red,  oblong-oblanceolate,  8-9  mm.  long,  somewhat  mucronate  at 
apex;  stamens  6-8  and  3-5  mm.  long;  anthers  1.5  mm.  long;  style  pubescent 
near  base,  scarcely  extending  to  sepal  tips;  stigma  4-lobed,  1.5  mm.  thick; 
fruit  dark  red,  subglobose  to  somewhat  elongate,  8-10  mm.  long. 

Type  locality.  Hacienda  Schunke,  La  Merced,  Dept.  Junin,  Peru.  This 
species  is  near  F.  Andrei  in  leaf -size,  puberulence,  etc.,  but  has  more  narrow 
and  attenuate-based  leaves,  longer  fruit,  and  more  southern  range.  I  have 
seen,  from  PERU :  Huanuco :  Distrito  Churubamba,  trail  Puente  to  Exito, 
Pampa  Hermosa,  1750  m.,  Mexia  8142  (F,  GH,  NY,  US) .  Junin :  La  Merced, 
Maclride  5616,  type  no.  (F,  GH,  US)  ;  Pichis  Trail,  1600  m.,  Killip  and  Smith 
25436  (F,  GH,  NY)  ;  Eneiias,  Killip  and  Smith  25784  (GH,  NY,  US).  Aya- 
cucho :  Ccarrapa,  between  Huanta  and  Rio  Apurimac,  1500  m.,  Killip  and 
Smith  22353  (F,  GH,  NY),  22336  (NY,  US).  Cuzco:  Prov.  Paucartambo: 
Rio  Tambomayo,  bet.  Tambomayo  and  Consuelo,  1100-1400  m..  West  7113 
(GH). 

(63)  Fuchsia  sessilifolia  Bentham 

(Plate  11,  fig.  57) 
Fuchsia  sessilifolia  Bentham,  PI.  Hartweg.,  176, 1845. 

Slender,  erect  shrubs  to  small  trees,  the  ultimate  branchlets  purplish  red, 
2-5  mm.  thick,  glabrous  except  for  the  minutely  strigulose-puberulent 
younger  parts ;  leaves  rather  remote,  in  whorls  of  3  or  4,  elliptic-lanceolate  to 
narrowly  elliptic-ovate  or  -obovate,  rounded  to  obtuse  at  base,  acuminate  at 
apex,  remotely  serrulate,  rather  coriaceous,  glossy  green  and  quite  glabrous 
above,  as  well  as  minutely  puberulent  on  the  impressed  veins,  glabrous  and 
paler  beneath,  or  puberulent  on  the  veins,  principal  lateral  veins  15-20  on 
each  side  of  midrib,  submarginal  vein  evident ;  blades  8-15  cm.  long,  2.5-4 
(5)  cm.  wide;  petioles  2-4  mm.  long,  strigulose-puberulent ;  stipules  subulate- 
deltoid,  about  1  mm.  long ;  inflorescence  terminal,  drooping,  racemose-panicu- 
late, 1-2  dm.  long,  canescent-strigose,  conspicuously  bracted ;  bracts  mostty 
1.5-2.5  cm.  long,  ovate-elliptic;  pedicels  strigulose,  slender,  3-5  mm.  long^ 
ovary  oblong,  strigulose,  4—6  mm.  long;  hypanthium  scarlet,  13-16  mm.  long, 
1.5  mm.  wide  at  base,  then  very  little  narrowed,  then  gradually  ampliate  until 
4-5  mm.  wide  at  summit,  finely  strigulose-puberulent  without,  pilose  within 


68  CALIF  OMNIA  ACADEMY  OF  SCIENCES  [Proc.  4th  Ser. 

most  of  its  length;  sepals  lanceolate,  acuminate,  divergent,  greenish  red, 
strigulose  on  backs,  8-10  mm.  long,  subulate  for  almost  1  mm.  at  tips;  petals 
scarlet,  oblong,  9-10  mm.  long,  3  mm.  wide,  acutish,  glabrous ;  stamens  ca.  7 
and  5  mm.  long ;  anthers  1  mm.  long ;  style  glabrous,  extending  almost  to  ends 
of  sepals;  stigma  about  1  mm.  thick;  berry  oblong,  10-12  mm.  long,  5-6  mm. 
thick,  somewhat  4-angled. 

Type  locality,  "in  sylvis  Guayan  in  declivitate  occidentali  montis  Pichin- 
cha,"  Ecuador.  The  species  seems  to  me  to  be  near  F.  Hartwegii  in  inflores- 
cence, hypanthium,  etc.,  but  differs  in  the  long,  subsessile,  narrow,  rather 
glabrous,  firm  leaves,  short  pedicels,  conspicuous  inflorescence.  It  grows  in 
southern  Colombia  and  northern  Ecuador  at  altitudes  of  from  2000-3600 
meters.  Representative  material  is,  COLOMBIA.  Antioquia :  Dauro,  Choco, 
near  Medellin,  Toro  1168  (NY).  Caldas:  Buenavista-Quindio,  Andre  in  1876 
(F,  GH,  NY).  Tolima:  Prov.  de  Mariquita;  El  Gallego,  Triana  3813  (US), 
Holton  in  1853  (NY).  Huila:  Cordillera  Central,  east  of  Neiva,  Rushy  and 
Pennell  687  (NY,  US)  ;  Cordillera  Oriental,  vertiente  occidental,  entre  Gabi- 
nete  y  Andalucia,  Cuatrecasas  8586  (US).  Cauca :  Valle  de  Quintero,  near 
Pitaio,  Pittier  1427  (NY,  US)  ;  Canaan,  near  Mt.  Purace,  Pennell  and  Killip 
6674  (GH,  NY,  US);  Paletara  to  Calaguala,  Peniiell  7111  (GH,  NY,  US). 
Nariiio  :  woods  near  Pasto,  Jameson  432  (US).  Putumayo :  above  El  Encavo, 
Laguna  La  Cocha,  Balls  7521  (US).  ECUADOR:  Carchi:  Canton  Tulcan 
near  Pun,  Mexia  7597  (POM,  US).  Pichincha :  sive  Andium  Quitensium, 
Jameson  under  Couthouy  (GH). 

(64)  Fuchsia  sylvatica  Bentham 

(Plate  11,  fig.  58) 

Fuchsia  sylvatica  Bentham,  PI.  Hartweg.,  176,  1845. 

F.  nigricans  Linden,  F1.  de  Series,  Ser.  1,  5  :pl.  481, 1849;  see  discussion  below. 
F.  atrorubra  Johnston,  Contr.  Gray  Herb.,  75 :31, 1925 ;  I  cannot  maintain,  although  sepals 
and  petals  are  somewhat  short. 

Low  shrub,  more  or  less  scandent,  openly  branched,  the  ultimate  twigs  pur- 
plish, minutely  canescent  strigulose,  2-A  mm.  thick ;  leaves  opposite  to  ternate, 
elliptic  to  elliptic-ovate  or  -obovate,  obtuse  to  acute  at  both  ends  or  sub- 
acuminate  at  apex  (  subentire  or  inconspicuously  denticulate,  membranaceous, 
deep  green  and  strigulose  above,  paler  and  more  densely  strigulose  beneatli, 
veins  reddish,  the  principal  lateral  ones  11-15  on  each  side  of  midrib,  sub- 
marginal  present;  leaf -blades  3-10  cm.  long,  1.5-5  (6)  em.  wide;  petioles 
strigulose,  6-20  mm.  long;  stipules  subulate-deltoid,  1-1.5  mm.  long,  some- 
w^hat  persistent ;  flowers  on  terminal  or  lateral  pendant  strigulose  racemes ; 
bracts  ovate  to  oblong  to  suborbicular,  1-2.5  cm.  long ;  pedicels  slender,  strigu- 
lose-canescent,  5-9  mm.  long;  ovary  narrowly  ellipsoid,  4-6  mm.  long;  hypan- 
thium 14-19  mm.  long,  ca.  1.5  mm.  thick  at  base,  gradually  ampliate  until 
2.5-3  mm.  wide  at  summit,  more  or  less  canescent-strigulose  without,  densely 
pilose  within;  sepals  pink  or  pale  red,  divergent,  linear-lanceolate,  6-9  mm. 


Vol.  XXV]  MUNZ :  THE  GENUS  FUCHSIA  69 

long,  acute,  not  subulate  at  apex,  strigulose  on  backs ;  petals  crimson  to  purple 
red,  oblong,  7-9  mm.  long,  acute;  stamens  4-6  and  3-5  mm.  long;  anthers 
1-1.5  mm.  long;  style  glabrous,  scarcely  reaching  sepal-tips ;  stigma  4-lobed, 
2  mm.  wide;  berry  oblong,  12-14  mm.  long,  4.5-6  mm.  thick. 

Type  locality,  "In  sylvis  Guayan  in  declivitate  occidentali  montis  Pichin- 
eha,"  Ecuador.  It  is  near  F.  sessilifolia  in  puberulence  and  short  pedicels,  but 
has  broader  leaves  and  more  pubescent  twigs.  I  have  not  seen  the  type  of 
F.  nigricans  from  Merida,  Venezuela,  but  the  three  Jahn  specimens  seem  to 
agree  with  the  description.  Though  these  come  from  much  farther  north  than 
the  general  range  of  F.  sylvatica,  I  cannot  separate  them  unless  I  see  the 
labels.  Perhaps  these  Venezuela  specimens  have  slightly  looser  pubescence 
and  fruit  about  2  mm.  longer.  Representative  material  of  sylvatica,  VENE- 
ZUELA: Merida:  Paramo  de  Arieagua,  2500  m.,  Jahn  1028  (US)  ;  between 
Bocono  and  Campo  Elias,  2000  m.,  Jahn  14  (US)  ;  selva  arriba  de  Palmira, 
2600  m.,  Jahn  509  (US) .  COLOMBIA  :  Antioquia :  Santa  Elena,  Archer  1195 
(US)  ;  Medellin,  Toro  972  (NY).  Caldas  :  Rio  Santa  Rita,  Salento,  Killip  and 
Hazen  8966  (GH,  NY,  US);  Salento,  Pennell  and  Hazen  10144  (GH,  NY, 
US)  ;  Quindio,  Dawe  769  (NY,  US)  ;  Santa  Elena,  above  Santuario,  Pennell 
103 13,  type  no.  atrorulra  (GH,  NY).  Cundimarca :  Fomeque,  Dawe  555  (US). 
Tolima:  La  Mediacion,  Quindio,  Triana  3511  (US)  ;  Rio  Toche  to  Machin, 
old  Quindio  Trail,  Killip  and  Hazen  9564  (GH).  ECUADOR:  Pichincha: 
w.  slope  Mt.  Pichincha,  Jameson  190  (GH),  Hartweg  984,  Geneva  specimen 
US) ;  between  Cosanga  and  Baeza,  2000  m.,  Mexia  7337  (US)  ;  Rio  Mapoto, 
(photo  F,  POM).  Tungurahua :  Canton  Quito  :  Alaspongo,  Mexia  7703  (POM, 
Penland  and  Summers  282  (F,  POM). 

Section  3.  KIERSCHLEGERIA  (Spach)  Munz,  new  comb. 

Kierschlegeria,  as  genus,  Spach,  Nouv,  Ann.  Sci.  nat.,   (2),  4:176,  1835;   as  subsect., 
Endlicher,  Gen.  PL,  1193, 1840. 

Pedicels  axillary,  pendulous ;  seeds  angled.  Leaves  sparse  or  subverticillate ; 
thickened  petiole-base  persistent  and  becoming  spine-like.  Sepals  reflexed, 
little  shorter  than  hypanthium. 

One  species,  Fuchsia  lycioides  Andr. 

(65)  Fuchsia  lycioides  Andrews 

(Platell,  fig.  59) 

Fuchsia  lycioides  Andr.,  Bot.  Eep.,  II,  pi.  120, 1800;  Bot.  Mag.,  25:1024,  1807. 

F.  rosea  Etnz  and  Pavon,  F1.  Peruv.,  3  :88, 1802. 

F.  spinosa  Presl,  Eel.  Haenk.,  2  :26,  pi.  51,  1835. 

F.  rosea  var.  spinosa  Eeiche,  Anal.  Univ.  Chile,  98:486,  1897;  PL  de  Chile,  2:268,  1898. 

Shrub  up  to  3  m.  tall,  the  older  branches  stout,  grayish,  knobby  with  dwarf 
lateral  branches,  intricately  branched  and  with  some  of  the  twigs  dull-spinose 
at  their  ends,  younger  twigs  more  slender  (2-3  mm.  thick),  fairly  straight, 
reddish,  subglabrous  with  short  sparse  pubescence  especially  toward  tips. 


70  CALIFOBXIA  ACADEMY  OF  SCIENCES  [Proc.  4th  Ser. 

beset  after  the  leaves  are  shed  by  the  persistent  thickened  woody  bases  of  the 
petioles  which  form  heavy  dull  spines  1.5-2  mm.  long;  leaves  alternate,  nu- 
merous, the  blades  lance-ovate  to  ovate,  6-24  mm.  long,  3-15  mm.  wide,  sub- 
entire,  acute  to  obtuse  at  both  ends,  glabrous  except  sometimes  at  tips  and 
margins,  somewhat  paler  beneath;  petioles  slender  above  the  thickened  base, 
glabrous  to  pubescent,  5-15  mm.  long ;  stipules  apparently  fused  with  petiole 
base ;  flowers  axillary,  solitary ;  pedicels  filiform,  5-8  mm.  long,  glabrous  to 
puberulent;  ovary  rounded,  1.5-2  mm.  long;  hypanthium  glabrous  without 
and  within,  red,  subcylindric,  4-7  mm.  long,  2-2.5  mm.  wide ;  sepals  4,  red, 
lanceolate,  acuminate,  spreading-reflexed,  glabrous  or  somewhat  pubescent, 
-4-7  mm.  long ;  petals  purplish,  obcordate,  3^  mm.  long ;  episepalous  stamens 
about  as  long  as  petals,  epipetalous  somewhat  shorter;  style  glabrous,  reddish, 
about  as  long  as  sepals ;  stigma  somewhat  4-lobed,  about  1  mm.  wide ;  berry 
blue-black,  subspheric,  4-6  mm.  thick. 

Type  locality  given  as  "North-west  coast  of  America,"  but  undoubtedly 
Valparaiso,  Chile,  near  w^hieh  most  specimens  of  the  species  have  been  col- 
lected. It  apparently  grows  on  dry  slopes  near  the  sea,  mostly  in  the  provinces 
of  Coquimbo  and  Aconcagua.  Representative  material,  CHILE  :  Fray  Jorge, 
Skottsherg  798  (NY),  3Iu7ioz  P.  no.  B-217  (GH),  Werdermann  1526  (B), 
Berninger  621  (B)  ;  Punta  de  Teatinos,  ^Yest  3921  (GH) ;  Coquimbo,  Gaudi- 
chaud  77  (B,  G).  Santiago:  Santiago,  Germain  in  1856-57  (G)  ;  Concon, 
Poeppig  123  (B).  Aconcagua :  Valparaiso,  Macrae  in  1825  (G),  Bertero  1006 
(G,  GH),  Gaudickaud  248  (F,  G),Rushtj  1800  (F,  NY,  US)  ;  Laguna  Verde, 
near  Valparaiso,  Looser  2290  (POM),  Garaventa  1641  (GH);  Choapa,  near 
Los  Vilos,  Mr.  and  Mrs.  J.  N.  Rose  19280  (NY,  US)  ;  Papudo,  Looser  2291 
(POM). 

Section  4.  SKINNERA  (Forst.)  DeCandolIe 

STcinnera  (FoRST.)  DC,  Prodr.,  3:39,  1828;  Endlicher,  Gen.  PI.,  1183,  1840;  Benth.  and 
Hook.,  Gen  PI.,  1,  pt.  2:791,  1862;  Eaimann  in  Engl,  and  Prantl,  Die  nat.  Pflanzenfam., 
Ill,  7:219,1893. 
as  genus,  Forst.,  Char.  Gen.,  57,  pi.  29,  1776. 

Hypanthium  infundibuliform,  swollen  above  ovary.  Sepals  reflexed.  Petals 
small  and  scale-like  or  wanting.  Filaments  long,  exserted.  Fruit  many-seeded ; 
seeds  very  small,  much  crowded,  in  2  or  more  rows.  Stigma  subcapitate,  some- 
what 4-lobed.  Pedicels  axillary,  pendulous.  Leaves  alternate. 

Type  species,  Fuchsia  excorticata  (Forst.)  L.f. 

Key  to  Species  of  Section  SKINNEEA 
A.  Petals  2-4  mm.  long;  plant  very  woody,  varying  from  a  liane  to  an  erect  shrub  or  tree; 
leaf -blades  whitish  beneath,  1-9  cm.  long 
B.  Trees;  leaf -blades  2-9  cm.  long,  vnt\\  about  10  principal  veins  on  each  side  of 
midrib. 
C.  Leaves  oblong-ovate  to  lance-ovate,  acute  to  acuminate  at  apex,  somewhat 
pilose  beneath  on  midrib.  New  Zealand 66.  F.  excorticata. 


Vol.  XXV]  MUNZ:  THE  GENUS  FUCHSIA  71 

CC.  Leaves  elliptic  to  elliptic-obovate,  rounded  to  obtuse  at  apex,  glabrous  beneath. 

Tahiti 67.  i^.  cyrtandroides. 

BB.  Not  trees;  leaf -blades  1-4  cm.  long,  with  3-9  main  veins. 

C.  Erect  shrub ;  leaves  oblong-ovate,  blades  mostly  2-4  cm.  long. .  .  68.  F.  Colensoi, 

CC.  Liane ;  leaves  suborbicular,  blades  mostly  1-2  cm.  long 69.  F.  perscandens. 

AA.  Petals  lacking ;  plant  small,  procumbent,  wth  very  slender  stems ;  leaf -blades  less  white 
beneath,  0.5-1.8  cm.  long. 
B.  Sepals  acuminate;  style  about  as  long  as  stamens;  hypanthium  tubular-campan- 

ulate,  7-8  mm.  long 70.  F.  procumbens. 

BB.  Sepals  obtuse ;   style  not  exserted  from  hypanthium ;  hypanthium  campanulate, 
6-7  mm.  long 71.  F.  KirTcii. 

(66)  Fuchsia  excorticata  (Forster)  L.f. 

(Plate  11,  fig.  60) 

SMnhera  excorticata  FoRST.,  Char.  Gen.,  58,  pi.  29,  1776. 

Fuchsia  excorticata  (Forst.)  L.  f.,  Suppl.,  217,  1781;  Lindley,  Bot.  Eeg.,  10:857,  1824. 

Wide-spreading  tree  10  m.  tall,  with  papery  exfoliating  light  brown  bark ; 
young  twigs  subgiabrous  to  puberulent,  subterete,  more  or  less  reddish  in 
color;  leaves  alternate,  oblong-ovate  to  lance-ovate,  rounded  to  subeordate  at 
base,  acute  to  acuminate  at  apex,  usually  somewhat  gland-denticulate,  green 
and  almost  glabrous  above,  whitish  and  generally  glabrous  beneath  except 
for  some  pilosity  along  veins  and  margins ;  blades  3-9  cm.  long,  1.5-3  cm. 
wide,  with  about  10  principal  veins  on  each  side  of  midrib  and  with  sub- 
marginal  vein ;  petioles  puberulent,  1-1.5  (4)  cm.  long;  stipules  ovate,  acumi- 
nate, puberulent,  less  than  1  mm.  long,  deciduous;  flowers  solitary,  axillary, 
pendant,  at  first  green,  then  purplish  red ;  pedicels  capillary,  puberulent, 
5-15  mm.  long;  ovary  subgiabrous,  5-6  mm.  long,  about  1  mm.  thick;  hypan- 
thium 4-angled,  8-15  mm.  long,  subgiabrous  without  and  within,  2-5  mm. 
wide  at  very  base  for  1—2  mm.,  then  1.5-2  mm.  wide  and  gradually  ampliate 
until  4-10  mm,  wide  at  summit;  sepals  spreading  to  reflexed,  ovate-lanceolate, 
glabrous,  acuminate,  8-14  mm.  long ;  petals  dark  purple,  erect,  2-4  mm.  long, 
elliptic,  lanceolate,  acutish ;  stamens  erect,  the  episepalous  6-9  mm.  long,  the 
others  somewhat  shorter;  anthers  blue,  1-2  mm.  long;  style  glabrous,  about 
equalling  sepals ;  stigma  subentire,  2  mm.  long,  yellowish ;  berry  ellipsoid, 
about  1  cm.  long,  dark. 

Type  locality  not  given;  species  ranging  through  New  Zealand.  Representa- 
tive material,  NEW  ZEALAND  :  North  Island :  Auckland  :  Waikowhai,  Miss 
Mackie  in  1932  (NY)  ;  Auckland,  Petrie  4110,  in  1898  (NY),  Meehold  9925 
(NY).  Wellington :  Feilding,  Allan  in  1924  (NY),  in  1928  (CAS)  ;  Wainui- 
o-Mata,  Meehold  4619  (BISH)  ;  Khandallah,  Self  in  1941  (POM,  WELT). 
South  Island :  without  definite  locality,  Kirk  (F).  Otago  :  Dunedin  town-belt, 
Colby,  DN,  from  Herh.  D.  Petrie  (GH) ;  cliff  west  of  St.  Clair,  Prue  Collier, 
per  M.  Neal,  168  (BISH)  ;  Nelson,  Kirk  253  (GH),  Marie  Neal  3  (BISH)  ; 
Canterbury :  Kaiturea  Valley,  Bank's  Peninsula,  Cockayne  4111  (NY)  ;  So. 
Canterbury,  edge  of  forest  on  Hunter  Hills,  Anderson  207  (F,  NY) . 


72  CALIFORNIA  ACADEMY  OF  SCIENCES  [Proc.  4th  Ser. 

(67)  Fuchsia  cyrtandroides  Moore 

(Plate  11,  fig.  61) 
Fuchsia  cyrtandroides  Moore,  B.  P.,  Bishop  Museum,  Occasional  Papers,  16:13, 1940. 

Tree  up  to  5  m.  tall,  the  young  twigs  quite  glabrous;  leaves  mostly  alternate, 
broadly  elliptic  to  elliptic-obovate,  green  above,  white  beneath,  glabrous, 
rounded  to  obtuse  at  base  and  apex,  subentire  to  obscurely  denticulate,  with 
about  10  principal  veins  on  each  side  of  midrib  and  with  indefinite  sub- 
marginal  vein;  petioles  8-14  mm.  long;  blades  2-9  cm.  long,  2-5.5  cm.  wide; 
stipules  apparently  ovate,  deciduous;  flowers  solitary,  axillary,  rose-magenta  ; 
pedicels  1.5-2.5  cm.  long ;  ovary  fusiform-clavate,  6-7  mm.  long,  2  mm,  thick ; 
hypanthium  about  1  cm.  long,  tubular-campanulate,  glabrous,  2-2.5  mm.  wide 
at  base  and  gradually  ampliate  to  5-6  mm.  at  summit;  sepals  spreading, 
lanceolate,  subacuminate,  10-14  mm.  long,  4-5  mm.  wide;  petals  erect,  oblong- 
ovate,  4  mm.  long  ? ;  stamens  about  as  long  as  sepals,  the  epipetalous  slightly 
shorter;  style  slightly  exceeding  stamens,  glabrous;  stigma  1.5  mm.  long, 
somewhat  4-lobed;  berry  black,  ellipsoid,  about  1  cm.  long. 

SOCIETY  ISLANDS:  Tahiti:  Orofena,  south  side,  rain  forest,  1500  m.. 
May  14, 1927,  MacDaniels  1315,  type  coll.  (BH,  BISH),  at  1600  m.,  8t.  John 
and  Fosherg  17005  (BISH).  The  species  is  very  near  F.  excorticata  in  its 
arborescent  habit,  large  leaves  which  are  white  beneath,  and  in  the  size  and 
shape  of  the  flowers.  It  differs  from  that  New  Zealand  species,  however,  in 
having  the  leaves  proportionately  wider,  more  obtuse  and  less  pubescent. 

(68)  Fuchsia  Colensoi  Hook.  f. 

(Plate  12,  fig.  62) 

Fuchsia  Colensoi  Hook,  f.,  Handb.  New  Zealand  FL,  728,  1867;  Trans,  and  Proc.  New  Zea- 
land Inst.,  1892 :  25,  pi.  19,  fig.  4,  5, 1893. 

Small  branching  erect  shrub,  with  ultimate  twigs  very  slender,  puberulent ; 
leaves  oblong-ovate,  rounded  to  subcordate  at  base,  obtuse  to  subacuminate  at 
apex,  subentire,  green  and  glabrous  above  except  for  fine  strigulosity  on 
impressed  veins,  white  and  glabrous  beneatli  except  for  some  pilosity  on  main 
veins,  short-ciliate,  principal  lateral  veins  6-9  on  each  side  of  midrib ;  petioles 
very  slender,  5-10  (20)  mm.  long ;  blades  1-4  cm.  long,  0.5-2  cm.  wide;  pedicels 
filiform,  5-10  mm.  long ;  ovary  ellipsoid,  about  5  mm.long,  2-2.5  mm.  wide ; 
hypanthium  8-10  mm.  long,  2-2.5  mm.  wide  at  base,  then  1-1.5  mm.  wide  and 
rapidly  ampliate  until  5-7  mm.,  red,  quite  glabrous;  sepals  greenish  to  reddish, 
spreading-reflexed,  lanceolate,  7-15  mm.  long ;  petals  purple,  2-3  mm.  long ; 
stamens  almost  equaling  sepals  or  about  two-thirds  as  long,  epipetalous ;  style 
exserted  by  2-5  mm.;  stigma  1.5-2  mm.  wide;  berry  oblong,  at  least  8  mm. 
long. 

Type  locality.  North  Island,  New  Zealand.  Material  seen,  NEW  ZEA- 
LAND :  North  Island  :  Papaitonga,  in  Kahikatea  forest,  Oliver  in  1941  (POI\I, 
WELT)  ;  Wammometa,   Wellington,   collector  not  given    (WELT,   photo 


Vol.  XXV]  MUNZ :  THE  GEXUS  FUCHSIA  73 

POM)  ;  Levin,  Wellington,  Oliver  in  1941  (POM,  WELT).  South  Island: 
Kaikoura,  Kirk  (GH)  ;  Pelorus,  Kirk,  (F) ;  Otago :  Port  Chalmers,  Petrie 
4109  (NY).  This  species  is  very  near  to  F.  excorticata  and  not  surely  distinct. 
With  inadequate  material  I  am  following  what  seems  to  be  a  tendency  among 
New  Zealand  botanists,  namely  to  recognize  these  several  Fuchsia  segregates 
as  species. 

(69)  Fuchsia  perscandens  Cockayne  and  Allan 
(Plate  12,  fig.  63) 
Fuchsia  perscandens  Cockayne  and  Allan,  Trans.  Proc.  N.  Z.  Inst.,  57:53, 1927. 

Climbing  shrub,  the  main  stems  few-branched,  the  twigs  appressed-puberu- 
lent,  slender ;  leaves  orbicular-ovate,  subcordate  at  base,  acute  at  apex,  sub- 
entire  to  sinuate-denticulate,  subglabrous  above  except  for  possible  minute 
puberulence  along  the  impressed  midrib,  whiter  and  subglabrous  beneath, 
short-ciliate,  with  about  3-6  main  veins  on  each  side  of  midrib ;  blades  1-2.5 
cm.  long,  almost  as  wide;  petioles  almost  filiform,  1-2  cm.  long;  pedicels  fili- 
form, about  1  cm.  long ;  flower  much  as  in  F.  Colensoi,  the  hypanthium  about 
9  mm.  long,  inflated  at  base,  etc. ;  sepals  about  7  mm.  long ;  petals  2.5  mm.  long; 
berry  dark  purple,  subcylindric,  about  1  cm.  long,  5  mm.  thick. 

Type  locality,  "forests  near  Feilding,"  North  Island,  New  Zealand.  Material 

seen,  NEW    ZEALAND:   North   Island:   Palmerston  North,   Zolro   3471 

(CAS)  ;  Feilding,  forest  margin,  H.  H.  Allan,  Aug.  1924  (NY);  Sinclair 

Head,  Wellington  Harb.,  Bealy  1  and  2  (POM,  WELT).  A  second  sheet  from 

Feilding  also  at  New  York  is  an  intergrade  with  F.  excorticata,  with  longer 

and  more  oblong  leaves  which  are  pilose  beneath  on  the  veins;  it  may  be  a 

hybrid. 

(70)  Fuchsia  procumbens  R.  Cunningham 

(Plate  12,  fig.  64) 
Fuchsia  procumbens  E.  Cunn.,  Ann.  Nat.  Hist.,  3 :31,  1839. 
F.  prostrata  Baill.,  Bull.  Soc.  Linn.,  Par.  1:270,  1880;  apparently  a  name  for  the  above. 

Procumbent,  woody,  with  very  slender  glabrous  stems;  leaves  alternate, 
not  crowded;  petioles  very  slender,  channeled  and  pubescent  above,  1.5-3  cm. 
long ;  stipules  minute,  ovate,  deciduous ;  blades  suborbicular  to  round-ovate, 
subcordate  at  base,  serrulate,  subglabrous  except  on  pubescent  margin  and 
veins  of  lower  surface,  somewhat  paler  beneath  than  above,  1-1.8  cm.  long, 
sometimes  somewhat  wider,  with  about  5  principal  veins  on  each  side  of  mid- 
rib ;  flowers  solitary,  axillary;  pedicels  slender,  subglabrous,  scarcely  1  cm. 
long ;  ovary  3^  mm.  long,  fusif orm-clavate ;  hypanthium  apparently  dark 
red,  tubular-campanulate,  7-8  mm.  long,  constricted  above  ovary,  then 
abruptly  dilated  to  about  3  mm.  wide,  then  more  gradually  until  5  mm.  wide 
at  summit,  glabrous  without  and  within ;  sepals  reflexed,  green,  lance-deltoid, 
acuminate,  6  mm.  long,  2.5  mm.  wide;  petals  lacking;  stamens  slightly  un- 
equal, erect,  ca.  4  and  5  mm.  long;  anthers  1  mm.  or  slightly  longer;  style 
glabrous,  about  as  long  as  stamens,  purplish;  stigma  1.5-2  mm.  thick;  fruit 
not  seen. 


74  CALIFOEXIA  ACADEMY  OF  SCIENCES  [Proc.  4th  Ser. 

Type  locality,  North  Island,  New  Zealand,  "around  the  village  of  ]\Iatauri 
on  the  east  coast  opposite  the  Cavallos  Isles."  Material  seen,  NEW  ZEA- 
LAND:  North  Island,  garden  specimen,  Khandallah,  Sntherland  in  1941 
(POM).  South  Island:  Canterbury:  Christchurch,  Laing's  Garden,  Meehold 
25160  (BISH)  ;  Timaru,  park  gardens,  Anderson  35  (NY). 

(71)  Fuchsia  Kirkii  Hooker  f . 

Fuchsia  Kirkii  Hook.  f.  ex  T.  Kirk,  Trans.  New  Zealand  Inst.,  1 :92,  1868;  Icon.  PL,  11 :66, 
pi.  1083,  1871. 

"With  prostrate  habit  and  slender  stems  of  F.  procumhens ;  petioles  1-1.5 
cm.  long,  blades  5-10  mm.  long,  orbicular-cordate;  pedicels  ca.  5  mm.  long; 
ovary  ca.  3  mm.  long ;  hypanthium  campanulate,  6-7  mm.  long,  5-6  mm.  wide 
at  summit ;  sepals  reflexed,  oblong,  obtuse,  5  mm.  long,  2.5  mm.  wide ;  petals 
0;  stamens  3-4  mm.  long;  style  included  in  hypanthium  ;  stigma  less  than  0.5 
mm.  wide. 

Type  locality.  Great  Barrier  Island,  New  Zealand.  Specimens  seen,  NEW 
ZEALAND  :  North  Island :  Tryphena  Bay,  T.  Kirk  (F,  GH)  ;  Port  Tryphena, 
Kirk  528  (GH).  Apparently  distinct  from  F.  procumhens  in  having  a  shorter 
wider  hypanthium,  sepals  more  oblong  and  obtuse ;  style  shorter  and  stigma 
smaller. 

Section  5.  HEMSLE YELLA  Munz,  new  section 

Nectary  and  hypanthium-base  fused ;  petals  lacking  or  nearly  so.  Hypan- 
thium funnelform  to  cylindro-funnelform,  enlarged  at  base.  Sepals  more  or 
less  connate  at  base.  Leaves  tending  to  be  deciduous  at  anthesis. 

Nectarium  basem  hypanthii  adnatum;  petalis  fere  nullis;  hypanthio  infundibuliforme 
vel  cylindro-infundibuliforme,  base  ampliato;  sepalis  base  fere  connatis. 

Type  species.  Fuchsia  apetala  Ruiz  and  Pavon.  This  section  is  dedicated  to 
W.  B.  Hemsley  who  published  a  paper  on  the  apetalous  Fuchsias  of  South 
America  in  the  Journ.  of  Bot.  14  :69-70,  1876. 

Key  to  Species  of  Section  HEMSLEYELLA 

A.  Hypanthium  less  than  2  cm.  long;  leaves  with  13-16  principal  lateral  veins  on  each  side 

of  midrib 83.  F.  cestroides. 

AA.  Hypanthium  more  than  2.5  cm.  long;  leaves  with  5-10  principal  lateral  veins  on  each 
side  of  midrib. 
B.  Hypanthium  at  least  4-5  times  as  long  as  the  sepals  (including  the  connate  base  of 
the  latter). 

C.  Leaves  well  developed  at  anthesis;  petioles  mostly  3-7  cm.  long;  pedicels  3-5 

cm.  long;  hypanthium  10-14  cm.  long 81.  F.  Garlepiriana. 

CC.  Leaves  mostly  deciduous  at  anthesis;  petioles  mostly  1-3  em.  long;  pedicels 

0.5-2.5  cm.  long;  hypanthium  6-12  cm.  long 82.  F.  maciantha. 

BB.  Hypanthium  usually  1.5-3  times  as  long  as  sepals  (including  connate  base). 
C.  Hypanthium  glabrous  without. 

D.  Leaves  alternate,  lance-ovate,  the  blades  mostly  2.5-4  times  as  long  as 
wide;  hypanthium  8-12  mm.  wide;  sepals  greenish 73.  F.  tuherosa. 

I 


Vol.  XXV]  MUNZ :  TEE  GENUS  FUCHSIA  75 

DD.  Leaves  mostly  subopposite,  elliptic-  or  oblong-ovate,  the  blades  not  more 
than  twice  as  long  as  wide ;  hypanthium  5-8  mm.  wide ;  sepals  red. 

E.  Petals  2  mm.  long ;  sepals  8-12  mm.  long.  Mexico 72.  F.  decidua. 

EE.  Petals  none;  sepals  18-22  mm.  long.  South  America. 

F.  L  eaves  largely  deciduous  at  an  thesis ;  stamens  exceeding  sepals ; 
style  1-1.5  cm.  longer  than  sepals.  Peru  to  Bolivia. 

74.  F.  juntasensis. 

FE.  Leaves  well  developed  during  anthesis ;   stamens  shorter  than 
sepals ;  style  scarcely  extending  beyond  sepals.  Venezuela. 

75.  F.  memhranacea. 
CC.  Hypanthium  not  glabrous  under  a  lens. 

D.  Leaves  lance-ovate ;    twigs   glabrous ;   hypanthium  sparsely  and   finely 

glandular-pubescent 76.  F.  salicifolia. 

DD.  Leaves  elliptic-  to  oblong-ovate ;  twigs  not  glabrous. 

E.  Leaves  subopposite ;  pubescence  very  fine,  short,  velvety ;  hypanthium 

5-8  mm.  wide.  Largely  Bolivian 77.  F.  tunariensis. 

EE.  Leaves  alternate;  pubescence  longer;  hypanthium  7-11  mm.  wide. 

F.  Stamens  longer  than  sepals;  pedicels  1.5-2.5  cm.  long;  inflores- 
cence short-pubescent.  Venezuela  to  northern  Peru. 

78.  F.  apetala. 

FF.  Stamens  shorter  than  sepals ;  pedicels  1-1.5  cm.  long ;  inflores- 
cence shaggy-pilose.  Central  Peru  to  Bolivia. 

G.  Leaves  deciduous  at  anthesis;  hairs  white,  0.5-1  mm.  long. 

79.  F.  liirsuta. 
GG.  Leaves  present  at  anthesis;  hairs  brown,  1-1.5  mm.  long. 

80.  F.  unduavensis. 

(72)  Fuchsia  decidua  Standley 
(Platel2,  fig.  65)      . 
Fuchsia  decidua  Standley,  Pub.  Field  Mus.  Nat.  Hist.,  Bot.  4:248,  1929. 

Shrub  with  long  snbterete  or  somewhat  4-angled.  branches,  the  j^ounger  ones 
3-9  mm.  thick,  brownish,  glabrous,  with  internodes  1-3  cm.  long;  leaves  oppo- 
site, unknown,  deciduous  at  time  of  flowering ;  inflorescence  short  racemose 
lateral  panicles  2-6  cm.  long,  with  glabrous,  somewhat  glaucous  very  slender 
branches;  pedicels  slender,  recurved,  3-8  mm.  long;  hypanthium  2-3.5  cm. 
long,  almost  2  mm.  wide  at  base,  then  narrowed  into  an  elongate  tube  which 
gradually  flares  upward  until  5-6  mm.  wide  at  mouth,  somewhat  glaucous, 
light  vermilion,  glabrous  without  and  within ;  sepals  vermilion,  oblong,  ob- 
tuse, 8-12  mm.  long,  4  mm.  wide ;  petals  suborbicular,  2  mm.  long ;  stamens 
about  9  and  6  mm.  long;  filaments  thickish;  anthers  1.5-1.8  mm.  long;  style 
glabrous,  about  equal  to  sepals;  stigma  barely  1  mm.  long;  berry  not  seen, 
but  probably  longer  than  thick. 

Type  locality,  MEXICO  :  Jalisco :  Real  Alto,  La  Bufa,  Sierra  Madre  Occi- 
dental, at  2500  m.;  Mexia  1601  being  the  type  no.  (F,  GH,  CAS,  MICH,  NY, 
US).  A  second  collection  of  apparently  the  same  thing  comes  from  the  state 
of  Guerrero;  Sierra  Madre  near  Chilpanengo  (Chilpancingo?),  Nelson  2195a 
(US)  ;  none  of  the  flowers  is  fully  matured,  hence  shorter  than  in  the  type 
collection.  Not  knowing  the  leaves,  I  find  it  difficult  to  determine  a  close  rela- 


76  CALIFOENIA  ACADEMY  OF  SCIENCES  [Proc.  4th  Ser. 

tionship  for  this  species  in  the  section  Eufiuchsia,  but  it  seems  to  be  near  to  the 
apetalous  species  of  Hemsleyella  in  its  greatly  reduced  petals.  Some  of  these 
species  have  also  lateral  reduced  inflorescences. 

(73)  Fuchsia  tuberosa  Krause 

Fuchsia  Ulcerosa  Krause,  Fedde  Rep.,  1 :170,  1905. 

Shrub  up  to  1  m.  tall,  epiphytic  or  terrestrial;  roots  bearing  clusters  of 
roundish  tubers;  stems  few-branched,  reddish,  glabrous,  the  branchlets  2-5 
mm.  thick;  leaves  may  shed  at  time  of  flowering,  alternate,  lance-ovate  to 
lanceolate  or  ovate,  subglabrous,  round  or  acute,  sometimes  cordate  at  base, 
curved-acuminate  at  apex,  entire  to  denticulate,  paler  beneath  than  above, 
with  6-8  principal  lateral  veins  on  each  side  of  midrib  ;  blades  5-10  cm.  long, 
2-3.5  cm.  wide;  petioles  slender,  1-2  (3)  cm.  long;  stipules  deltoid,  about  0.4 
mm.  long,  deciduous ;  flowers  in  upper  axils,  sometimes  crowded  terminally ; 
pedicels  slender,  2-6  cm.  long,  glabrous  or  nearly  so;  ovary  cylindric,  glabrous, 
8-12  mm.  long,  1-2  mm.  thick ;  hypanthium  red,  3.5-8  cm.  long,  3-^  mm.  wide 
at  base,  then  narrowed  to  1.5-2  mm.  for  less  than  1  cm.,  then  abruptly  widened 
to  8-12  mm.  for  the  rest  of  its  length,  glabrous  without,  villous  within  at  base ; 
sepals  green,  erect-divergent,  lance-deltoid,  12-20  mm.  long,  5-8  mm.  wide, 
acute,  connate  for  one-fourth  their  length ;  petals  0;  episepalous  stamens  about 
as  long  as  sepals,  alternate  ones  three-fourths  as  long,  yellow ;  anthers  3-4  mm. 
long,  yellow;  style  slender,  pubescent  near  base,  exserted  beyond  sepals  for 
1-5  cm.;  stigma  elongate-capitate,  2-3  mm.  long;  fruit  glabrous,  2-2.3  cm. 
long. 

Key  to  Varieties  of  Fuchsia  tuberosa 

A.  Hypanthium  3.5-5  cm.  long 73a.  var.  typica. 

AA.  Hypanthium  6-8  cm.  long 73b.  var.  inflata. 

(73a)  Fuchsia  tuberosa  Krause,  var.  typica  Munz,  new  name 

(Plate  12,  fig.  66) 

Fuchsia  tuierosa  Krause,  Fedde  Rep.,  1 :170, 1905. 

Fuchsia  chloroloha  Johnston,  Journ.  Arnold  Arboretum,  20:243, 1939. 

Hypanthium  3.5-5  cm.  long. 

Type  locality,  between  Sandia  and  Cuyocuyo,  Puno,  Peru,  at  2700-2800  m., 
Dept.  Cuzco:  "Pillahuata,"  Cerro  de  Cusilluyoc,  2200-2400  m.,  Pennell 
based  on  Weherhauer  875  at  Berlin  (photo  F,  POM).  Material  seen,  PERU : 
13973,  type  no.  chloroloha  (F,  GH)  ;  Marcapata  Valley,  1200  m.,  Herrera  1166 
(US)  ;  between  Achirani  and  Medias-Mayu,  2700  m.,  Vargas  11116  (F)  ;  Rio 
Tambomayo,  from  Pillahuata  bridge  to  head  of  Tambomayo  Grade,  1600- 
2300  m.,  West  7094  (GH);  between  Tanamayo  and  Tambomayo,  2000  m., 
Vargas  70/7094  (F).  The  Krause  description  does  not  mention  green  sepals, 
as  does  Johnston's,  but  the  region,  the  leaf-  and  flower-shape,  glabrous  hypan- 
thium, etc.,  certainly  make  ticberosa  and  chloroloha  the  same. 


Vol.  XXV]  MUNZ :  THE  GENUS  FUCHSIA  77 

(73b)  Fuchsia  tuberosa  Krause  var.  inflata  (Schulze-Menz)  Munz,  new  comb. 
Fuchsia  inflata  Schulze-Menz,  Notizbl.  Bot.  Gart.,  Berlin,  15 :  136, 1940. 

Hypanthium  6-8  cm.  long. 

Type  from  Cuzeo :  prov  Paucartambo,  between  the  tambos  Tres  Cruces  and 
Tambomayo,  above  Cosiiipata,  at  3100  m.,  April,  1914,  Weherhauer  6935  (F, 
GH,  POM,  US).  This  does  not  seem  specifically  distinct,  and  more  material 
may  show  that  it  is  not  worth  any  recognition. 

(74)  Fuchsia  juntasensis  0.  Kuntze 
(Plate  12,  fig.  67) 

Fuchsia  juntasensis  O.  Kuntze,  Kev.  Gen.  3,  part  2 :97, 1898. 

F.  Mattoana  Krause,  Engl.  Jahrb.,  37:599,  1906. 

F.  Steinhachii  Johnston,  Contr.  Gray  Herb.,  75:38,  1925. 

Vinelike,  usually  epiphytic  shrub,  5  or  more  dm.  tall,  few-branched,  the 
younger  twigs  usually  purplish,  subglabrous  or  sparsely  pubescent,  the  older 
ones  with  exfoliating  outer  tissues ;  leaves  mostly  subopposite,  tending  to  fall 
during  anthesis;  the  blades  elliptic-  to  oblong-ovate,  acute  to  obtuse  at  base, 
acuminate  at  apex  ,subentire,  4-8  cm.  long,  1.5-3.5  cm.  wide,  sparsely  short- 
pubescent  to  giabrescent  above  and  beneath,  paler  beneath  than  above,  prin- 
cipal lateral  veins  6-8  on  each  side  of  midrib ;  petioles  sparsely  pubescent, 
5-12  mm.  long,  ca.  1  mm.  wide;  stipules  subulate,  1-1.5  mm.  long,  caducous; 
flowers  few,  crowded  in  terminal  racemose  clusters,  often  on  short  lateral 
twigs ;  pedicels  slender,  subglabrous,  1.5-5  cm.  long ;  ovary  oblong-fusiform, 
puberulent  to  subglabrous,  5-10  mm.  long,  2-2.5  mm.  wide ;  hypanthium  rose 
to  flesh-colored,  tubular-funnelf orm,  2.5-4.5  cm.  long,  2-2.5  mm.  AVide  at  base, 
then  slightly  narrowed  for  one-third  its  length,  then  gradually  ampliate  until 
5-7  mm.  wide  at  summit,  glabrous  without,  pilose  within  near  base ;  sepals  4, 
reddish,  ovate-lanceolate,  18-22  mm.  long,  5-6  mm.  wide,  glabrous,  sub- 
acuminate,  connate  for  2-3  mm.  at  base ;  petals  none ;  episepalous  stamens 
20-26  mm.  long,  alternate  ones  16-20  mm.  long ;  anthers  cream,  3-4  mm.  long; 
style  pubescent  in  lower  portion,  extending  1-1.5  cm.  beyong  the. sepals; 
stigma  green,  oblong-globose,  2-3  mm.  long,  scarcely  lobed ;  berry  ellipsoid, 
about  12  mm.  long. 

Type  locality,  east  side  of  Cordillera,  at  3000  m.,  between  Cochabamba  and 
Rio  Juntas,  Bolivia.  I  have  not  seen  the  type  of  F.  juntasensis  so  labeled  and 
have  had  some  misgiving  about  taking  up  this  name  for  this  concept,  especially 
since  Kuntze's  description  sounds  as  if  the  calyx  were  short,  giving  sepals  as 
12  mm.  long,  and  even  if  one  adds  3-4  mm.  for  the  connate  base,  which  Kuntze 
may  well  not  have  included  as  part  of  the  sepals,  the  length  is  still  rather 
inadequate.  But  all  other  features  :  glabrous  flowers,  ovate,  acuminate  leaves, 
shape  of  hypanthium,  would  point  to  this  concept.  Of  F.  Mattoana  I  have 
seen  only  the  photograph;  between  it  and  F.  Steinhachii  I  can  see  no  difference 
except  the  somewhat  shorter  hypanthium   (25-28  mm.)   given  by  Krau^ 


LISRAR 


78  CALIFORNIA  ACADEMY  OF  SCIENCES  [Proc.  4th  Ser. 

However,  a  series  of  specimens  seems  to  break  down  the  validity  of  this  char- 
acter. For  the  species  which  I  am  calling  F.  jtintasensis  I  have  seen  the  follow- 
ing collections:  without  locality  Herb.  0.  Kuntze  (NY).  PERU:  between 
Cuzco  and  Santa  Ana,  3400  m.,  Weherhmier  4976,  type  of  Mattoana  at  Berlin 
(photo  F,  POM).  BOLIVIA  :  Depto.  Cochabamba  :  Ineachara,  2500  m.,  Stein- 
hach  5038,  type  Steinhachii  (NY)  ;  Quebrada  de  Corani,  prov.  Chapare,  2400 
m.,  Steinlach  9862  (F,  GH,  NY).  The  species  differs  from  F.  apetala  in  its 
glabrous  flowers  and  more  southern  range. 

(75)  Fuchsia  membranacea  Hemsley 
Fuchsia  membranacea  Hemsley,  Journ.  Bot.,  14:70,  1876. 

Shrub,  glabrous  throughout  except  ofr  the  minute  pubescence  on  the  upper 
surface  of  the  young  leaves  which  may  persist  along  the  veins ;  branches  few, 
younger  ones  purplish,  2.5-3.5  mm.  thick,  older  ones  exfoliating ;  leaves  sub- 
opposite,  well  developed  at  anthesis,  the  blades  membranaceous,  oblong-ovate, 
remotely  denticulate,  paler  beneath,  rounded  at  base,  short-acuminate  at  apex, 
3-5.5  cm.  long,  1.5-3.5  cm.  wide,  with  6-7  principal  lateral  veins  on  each  side 
of  midrib ;  petioles  puberulent,  6-15  mm.  long,  scarcely  1  mm.  wide ;  stipules 
subulate-deltoid,  about  0.5  mm.  long;  flowers  axillary ;  pedicels  slender,  2.5-4 
cm.  long;  ovary  oblong,  5-7  mm.  long,  2  mm.  thick;  hypanthium  tubular- 
funnelform,  4-5  cm.  long,  slightly  enlarged  and  2.5  mm.  wide  at  base,  then 
1.5-2  mm.  wide  for  about  1  cm.,  then  gradually  widened  in  upper  half  until 
7-9  mm.  broad,  glabrous  without,  pubescent  within  near  base ;  sepals  diver- 
gent, ovate-lanceolate,  18-22  mm.  long,  6-7  mm.  wide,  subacuminate,  very 
little  connate  at  base ;  petals  0;  episepalous  stamens  14-20  mm.  long,  alternate 
ones  9-15  mm.  long ;  anthers  2-3  mm.  ling ;  style  pubescent  in  lower  portion, 
scarcely  extended  beyond  sepals;  stigma  oblong-capitate,  2-3  mm.  long ;  berry 
short-ellipsoid,  about  1  cm.  long. 

Type  locality,  Caracas,  Venezuela;  type  based  on  Linden  372,  of  which 
number  I  have  seen  fragment  (F)  and  photo  (F,  POM).  Other  material  seen, 
VENEZUELA  :  state  of  Trujillo  :  Paramo  de  la  Cristalina,  2900  m.,  J  aim  12 
(US).  Merida:  Paramo  de  Timotes,  3000  m.,  J  aim  508  (GH,  US).  Like  F. 
juntasensis,  the  species  has  glabrous  flowers,  but  it  is  leafy  at  anthesis  and  has 
shorter  stamens. 

(76)  Fuchsia  salicifolia  Hemsley 

Fuchsia  salicifolia  Hemsley,  Journ.  Bot.,  14:70, 1876. 

Epiphytic  shrub,  mostly  glabrous  except  about  the  flowers ;  branchlets  few, 
purplish  with  persistent  epidermis ;  leaves  fully  developed  during  anthesis, 
alternate,  lanceolate,  obtuse  at  base,  long-acuminate  at  apex,  remotely  den- 
ticulate, thickish,  with  about  10  principal  lateral  veins  on  each  side  of  midrib, 
the  blades  6-10  cm.  long,  2-2.6  cm.  wide  ;  petioles  ca.  1  cm.  long,  1.5  mm.  wide ; 
stipules  not  seen;  flowers  few,  axillary;  pedicels  2-2.5  cm.  long,  slender, 


Vol.  XXV]  MUNZ:  THE  GENUS  FUCHSIA  79 

pubescent,  with  some  gland-tipped  hairs;  ovary  linear-oblong,  1  cm.  long, 
2-2.5  mm.  wide;  hypanthium  tubular-funnelform,  4-5.5  cm.  long,  sparsely 
glandular-pubescent  without,  2.5  mm  .thick  at  base,  then  narrower  for  about 
1  cm.,  then  gradually  widened  to  7  mm. ;  sepals  lanceolate,  acuminate,  spread- 
ing, ca.  3  cm.  long,  1  cm.  wide,  connate  at  base  for  2-3  mm. ;  petals  0 ;  epi- 
sepalous  stamens  about  27  mm.  long ;  alternate  ones  20  mm.  long ;  anthers  3-4 
mm.  long;  style  exserted;  stigma  ''clavate";  berry  about  2  cm.  long. 

Type  locality,  Sandillani ;  I  am  not  sure  whether  this  is  in  southern  Peru  or 
in  Bolivia.  The  only  specimen  I  have  seen  which  I  can  refer  here  is  from 
Unduavi,  North  Yungas,  Bolivia  at  2450  m..  Bushy  1803  (NY)  ;  it  is  a  speci- 
men with  1  leaf  and  1  flower.  The  species  seems  distinct  in  its  narrow  leaf, 
subglabrous  condition  and  long  sepals.  In  hypanthium-shape  it  seems  near 
F.  apetala. 

(77)  Fuchsia  tunariensis  0.  Kuntze 

(Plate  13,  fig.  68) 
Fuchsia  tunariensis  O.  Kuntze,  Eev.  Gen.  PI.,  3,  part  2 :98, 1898. 

Epiphytic  or  prostrate  to  scandent  shrub  with  tuberous  roots ;  stems  up  to 
8  dm.  long,  few-branched,  the  young  growth  pallid  with  densely  velvet  pubes- 
cence ;  twigs  often  knotty  and  irregular,  purplish,  densely  pubescent ;  leaves 
largely  subopposite,  tending  to  be  deciduous  at  anthesis ;  blades  elliptic-  to 
oblong-ovate,  4-15  cm.  long,  2-6  cm.  wide,  membranaceous,  densely  fine- 
pubescent,  velvety  when  young,  entire,  rounded  to  obtuse  at  base,  subacumi- 
nate  at  apex;  petioles  5-20  mm.  long,  densely  pubescent,  about  1  mm.  wide; 
stipules  subulate,  gland-like,  scarcely  1  mm.  long,  deciduous ;  flowers  few, 
solitary  in  uppermost  axils ;  pedicels  densely  pubescent,  1-1.5  cm.  long ;  ovary 
densely  short-pubescent,  oblong,  5-8  mm.  long ;  hypanthium  red  to  pinkish, 
tubular-funnelform,  4.5-5.5  cm.  long,  densely  short-pubescent  without,  pilose 
within  lower  portion,  2.5-3  mm.  wide  at  base,  then  1.5-2  mm.  wide  for  about 
1.5  cm.,  then  gradually  ampliate  until  5-8  mm.  wide  at  mouth;  sepals  red, 
lance-ovate,  acute,  18-22  mm.  long,  connate  for  about  3  mm.  at  base ;  petals 
0;  episepalous  stamens  13-18  mm.  long,  alternate  ones  10-13  mm.  long; 
anthers  about  3  mm.  long ;  style  pilose  at  base,  scarcely  exserted  or  as  much  as 
1  cm.  longer  than  sepals ;  stigma  ca.  2  mm.  long,  subglobose ;  fruit  hairy, 
ellipsoid,  almost  2  cm.  long. 

Type  locality,  "Tunarigebirge,"  Cochabamba,  Bolivia,  at  3000  m.  The  type 
in  Herb.  Otto  Kuntze  (NY)  is  badly  broken,  the  solitary  flower  has  pedicel, 
ovary  and  lower  half  of  hypanthium,  but  the  pubescence  is  very  character- 
istic. In  fact,  it  is  the  close,  fine,  dense  pubescence  which  distinguishes  this 
species,  of  which  I  have  seen  the  following  representation:  PERU:  Cuzco, 
prov.  Paucartambo,  below  "Pillahuata,"  2300  m..  West  7106  (GH)  with 
rather  large  sepals,  but  typical  pubescence.  BOLIVIA  :  Cochabamba  :  Tunari, 
Kuntze  in  1892,  type  (NY)  ;  Ayopaya,  Sailapata,  3500  m.,  Cardenas  3393 
(US),  3369  (US).  Pongo,  Tate  188  (NY). 


80  CALIFORNIA  ACADEMY  OF  SCIENCES  [Peoc.  4th  Ser. 

(78)  Fuchsia  apetala  Ruiz  and  Pavon 
(Platel3,  fig.  69) 
Fuchsia  apetala  Ruiz  and  Pav6n,  F1.  Peruv.,  3 :89,  pi.  322,  fig.  b,  1802. 

F.  msig7iis  Hemsley,  Journ.  Bot.,  14:69,  1^76;  three  collections  were  cited  from  Ecuador, 
2  by  Jameson  and  1  by  Spruce.  The  last  whicli  cannot  be  taken  as  the  type,  is  the  only  one 
I  have  seen,  but  I  am  unable  to  separate  insignis  from  apetala. 

Vine-like  shrub,  often  epiphytic,  up  to  about  1  m.  high,  the  younger 
branches  reddish,  more  or  less  pubescent,  with  some  gland-tipped  hairs,  rather 
coarse,  commonly  3-5  mm.  thick,  with  exfoliating  epidermis  and  frequently 
M'ith  adventitious  roots;  leaves  largely  deciduous  at  anthesis,  alternate,  the 
blades  elliptic-ovate,  subentire,  rounded  to  subcordate  at  base,  acute  to 
acuminate  at  apex,  somewhat  coriaceous,  finely  pubescent  to  glabrescent 
above,  villous  beneath,  especially  beneath  on  midrib  and  veins,  principal 
lateral  veins  6-8  on  each  side  of  midrib ;  leaf -blades  5-9  cm.  long,  3-4.5  cm. 
wide;  petioles  1.5-2.5  cm.  long,  somewliat  villous,  1.5-2  mm.  wide;  stipules 
subulate,  1.5-2  mm.  long,  caducous;  flowers  few,  crowded  near  apex  of  branch- 
lets;  pedicels  slender,  1.5-2.5  cm.  long,  villous,  with  some  gland-tippecl  hairs; 
ovary  cylindrical,  8-15  mm.  long,  villous,  2-A  mm.  thick ;  hypanthium  orange- 
scarlet,  4-5.5  cm.  long,  tubular-funnelform,  3-5  mm.  wide  at  bulbous  base, 
then  abruptly  narrowed  to  1.5-3  mm.  for  about  1  cm.,  then  gradually  ampliate 
until  7-11  mm.  wide,  sparsely  glandular-villous  without,  villous  within  lower 
portion;  sepals  reddish-orange,  more  or  less  spreading,  lance-ovate,  22-25 
mm.  long,  8-10  mm.  wide,  acute  to  obtusish,  pubescent  without,  connate  for 
2-3  mm.  at  base;  petals  0;  episepalous  stamens  25-30  mm.  long,  alternate 
18-20  mm.  long;  anthers  green,  3-4  mm.  long;  style  pubescent  at  base,  ex- 
serted  1-2  cm.  beyond  the  sepals ;  stigma  subglobose,  2-3  mm.  long,  usually 
scarcely  lobed ;  fruit  oblong,  red. 

Type  locality  ''Habitat  copiose  in  Huassahuassi  et  Muna,"  Peru.  At  Field 
]\Iuseum  are  fragments  of  what  I  here  consider  to  be  apetala  mixed  with  F. 
niacrantha  Hook.,  collected  by  Ruiz  and  Pavon ;  my  judgment  as  to  which  is 
to  be  taken  as  apetala  is  based  on  the  plate  by  Ruiz  and  Pavon.  I  prefer,  since 
this  Ruiz  and  Pavon  material  in  its  present  condition  contains  both  entities 
and  since  I  believe  these  are  separable  on  the  basis  of  geographical  and 
morphological  characters,  to  keep  the  name  apetala  for  the  Ruiz  and  Pavon 
plant  that  was  illustrated,  rather  than  to  treat  all  as  one  thing  as  does  Mac- 
bride  (Field  Mus.  Nat.  Hist.,  Bot.  13  :part  4,  no.  1 :546,  1941).  I  would  refer 
the  following  material  to  apetala,  VENEZUELA :  Merida :  Paramo  de  Aric- 
gua,  2000  m.,  Jahn  1030  (US) ;  Paramo  de  San  Jose,  2400  m.,  Jalin  969  (US)  ; 
Paramo  de  Mucuchachi,  1800  m.,  Jahn  981  (US).  ECUADOR:  between 
Cuenca  and  Huigra,  Chimborazo,  2700-3000  m.,  Hitchcock  21701  (GH,  NY, 
US)  ;  Huigra,  Rose  and  Rose  22230  (GH,  NY,  US)  ;  Mt.  Azuay,  8pruce  5976 
(GH,  NY,  photo  of  Geneva  specimen  at  F,  POM).  PERU?:  Churupullana, 
Domhey  ex  Herb.  Mus.  Paris  (F).  I  can  see  no  difference  between  the  Vene- 
zuela and  Ecuador  specimens  in  spite  of  the  discontinuous  distribution. 


Vol.  XXV]  MUNZ:  TEE  GENUS  FUCHSIA  81 

(79)  Fuchsia  hirsuta  Hemsley 
(Plate  13,  fig.  70) 
Fuchsia  hirsuta  Hemsley,  Journ.  Bot.,  14 :69,  1876. 

Tuber-bearing,  climbing,  epiphytic  shrub  or  scanclent  over  faces  of  cliffs, 
few-branched,  mostly  leafless  at  anthesis,  the  younger  twigs  3-8  mm.  thick, 
knotty,  tortuose,  densely  pilose  with  white  hairs  not  over  1  mm.  long,  more  or 
less  purplish  underneath  the  hairs;  leaves  alternate,  ovate  to  oblong-ovate, 
membranous,  subentire  to  denticulate,  rounded  at  base,  acute  to  almost  acumi- 
nate at  apex,  densely  pilose  on  both  surfaces  when  young,  more  or  less  glabres- 
cent  except  for  veins  of  under  surface,  principal  lateral  veins  5-8  on  each  side 
of  midrib  ;  leaf -blades  6-12  cm.  long,  3.5-6  cm.  wide ;  petioles  pilose,  1-4  cm. 
long,  1.5-2  mm.  wide;  stipules  pilose,  lanceolate,  ca.  1  mm.  long;  flowers  few 
to  many,  crowded  near  tips  especially  of  lateral  twigs ;  pedicels  slender,  pilose, 
1-1.5  cm.  long;  ovary  cylindric,  densely  shaggy-pilose  and  pubescent,  8-12 
mm.  long,  1.5-2.5  mm.  thick ;  hypanthium  tubular-funnelform,  densely  pilose 
and  pubescent,  with  some  hairs  gland-tipped,  fiery  to  purplish  red,  3-5.5  cm. 
long,  2-3  mm.  thick  at  base,  then  slightly  narrowed  for  one-third  its  length 
and  then  rather  abruptly  widened  into  the  cylindrical  upper  part  which  is 
8-10  mm.  wide,  pilose  within  the  lower  portion ;  sepals  reddish,  ovate  to  lance- 
ovate,  obtuse  to  acute,  pilose  on  backs,  15-20  mm.  long,  7-10  mm.  wide,  con- 
nate only  a  short  distance  at  base ;  petals  0 ;  episepalous  stamens  15-18  mm. 
long,  alternate  ones  two-thirds  as  long ;  anthers  about  3  mm.  long ;  style  pilose 
above  the  base,  extending  to  as  much  as  1.5  cm.  beyond  the  sepals;  stigma 
about  2  mm.  long ;  berry  narrowly  ellipsoid,  almost  2  cm.  long. 

Type  locality,  "Peruvia" ;  I  have  not  seen  the  type,  LecJiler  1989.  The  species 
is  included  in  F.  apetala  by  Macbride  (Field  Mus.  Nat.  Hist.,  Bot.,  13,  part  4, 
no.  1 :546,  1941),  but  to  me  it  is  a  more  southern  entity  with  short  sepals  and 
more  included  stamens,  gray-pilose  and  shaggy  ovaries  and  hypanthia.  I  have 
seen  from  PERU:  Ayacucho  :  Punaccuanca  Pass,  4100  m..  West  3669  (GH). 
Cuzco :  prov.  Paucartambo,  disto.  Marcachea,  forest  of  Pucara,  3400  m., 
Vargas  11170  (F) ;  below  Tres  Cruces,  Weherlauer  6975  (F,  GH)  ;  prov. 
Urubamba,  disto.  Chincheros,  Chincheros,  3700  m.,  Vargas  9602  (F)  ;  Yucay, 
3000  m.,  Soukup  715  (F)  ;  Cuzco,  2500  m.,  Herrera2189  (GH)  ;  Saxaihuaman, 
Herrera  2189  (F)  ;  Combapata,  Canchis,  3700  m.,  Vargas  1900  (GH).  Puno, 
Soukup  525  (F),  BOLIVIA:  La  Paz:  Ingenio,  between  Sorata  and  Mapiri, 
3700  m.,  Cardenas  1142  (NY)  ;  La  Paz  :  prov.  Larecaja,  near  Sorata,  Mandon 
629  (F,  GH)  ;  Pongo,  White  248  (GH,  US). 

(80)  Fuchsia  unduavensis  Munz,  new  species 
(Plate  13,  fig.  71) 

Much  like  F.  hirsuta,  but  with  leaves  well  developed  at  time  of  flowering, 
blades  elliptic-ovate,  7-10  cm.  long,  2-5  cm.  wide.  Math  6-7  principal  lateral 


82  CALIFORNIA  ACADEMY  OF  SCIENCES  [Proc.  4th  Ser. 

veins  on  each  side  of  midrib ;  young  stems,  ovaries,  pedicels,  hypanthia  with 
spreading,  brownish  hairs  about  1.5  mm.  long;  hypanthium  4.5^5.5  cm.  long, 
7-8  mm.  wide  in  upper  part ;  sepals  2  cm.  long,  lance-ovate,  acute  to  obtusish. 

F.  hirsutae  similis,  foliis  ad  anthesin  praesentibus,  elliptico-ovatis,  laminis  7-10  cm. 
longis,  2-5  cm.  latis;  caulibus  junioribus,  ovariis,  pedicelis,  hypanthiis  capillis  fuscis  et  1.5 
mm.  longis  vestitis;  hypanthio  4.5-5.5  cm.  longo,  supra  7-8  mm.  lato;  sepalis  2  cm.  longis, 
lanceolato-ovatis,  acutis  vel  subobtusis. 

Type:  Unduavi,  North  Yungas,  Bolivia,  at  3600  m.,  Feb.  13, 1907,  Buchtien 
2925,  United  States  Nat.  Herb.  1177335.  Other  collections  from  same  place : 
at  3200  m.,  Nov.,  1910,  Buchtien  (F),  at  3600  m.,  Nov.,  1910,  Buchtien  2925 

(US). 

(81)  Fuchsia  Garleppiana  Kuntze  and  Wittmack 

(Plate  14,  fig.  72) 
Fuchsia  Garleppiana  Kuntze  and  Wittm.,  Gartenfl.,  47 : 461,  fig.  96, 1893. 

Tuberous,  the  thickened  roots  said  to  be  20-90  cm.  long ;  slender-stemmed, 
few-branched,  up  to  1  m.  high,  epiphytic  or  on  wet  slopes,  the  ultimate  branch- 
lets  reddish,  soft-pubescent,  2-4  mm.  thick,  tardily  exfoliating ;  leaves  alter- 
nate, well  developed  at  anthesis,  ovate  to  oblong-ovate,  the  blades  thin, 
rounded  to  subcordate  at  base,  subacuminate  at  apex,  subentire  to  remotely 
denticulate,  finely  pubescent  above  and  beneath,  with  8-9  principal  veins  on 
each  side  of  midrib ;  blades  6-10  cm.  long,  4-6  cm.  wide ;  petioles  pubescent, 
2.5-7  cm.  long,  about  1  mm.  wide;  stipules  lanceolate,  scarcely  1  mm.  long, 
deciduous ;  flowers  in  axils  on  main  growing  branches ;  pedicels  slender,  soft- 
pubescent,  3-5  cm.  long;  ovary  5-10  mm.  long  at  anthesis,  soft-pubescent,  2-3 
mm.  thick;  hypanthium  10-14  cm.  long,  subeylindric,  finely  pubescent  with- 
out and  within,  pink,  5-6  mm.  thick  at  base,  then  narrowed  to  2-3  mm.  and 
then  gradually  wider  until  8-10  mm.  broad  below  the  somewhat  contracted 
mouth  ;^epals  pink,  lance-ovate,  acute,  somewhat  divergent,  13-18  mm.  long, 
5-7  mm.  wide,  connate  almost  one-half  their  length,  fine-pubescent  on  both 
surfaces;  petals  0;  episepalous  stamens  about  three-fourths,  alternate  ones 
about  two-thirds  the  sepals;  anthers  2.5-3.5  mm.  long;  style  pubescent  near 
the  base,  slender,  scarcely  extending  beyond  the  sepals;  stigma  subglobose, 
2-3  mm.  thick ;  fruit  not  seen. 

Type  locality,  Cocapata,  Cochabamba,  Bolivia,  at  3000  m.  in  the  Tunari 
Mts.  The  species  is  near  to  F.  macrantha  in  its  long  hypanthium,  but  differs 
in  many  respects :  flowers  not  on  spur-like  branchlets,  plant  leafy  at  anthesis, 
petioles  longer  and  more  slender,  pubescence  finer  and  shorter,  sepals  hairy 
on  both  surfaces,  filaments  pubescent;  hypanthium  narrowed  at  end.  Material 
seen,  PERU:  Dept.  du  Cuzco,  Aout.,  1847,  Weddcll,  from  Herb.  Mus.  Paris 
(F).  BOLIVIA:  Cochabamba:  Incachaca,  2700  m.,  Cardenas  688  (US); 
Incacorral-Aduana,  Chapare,  2700  m.,  Steinbach  9538  (GH,  NY)  ;  Tunarige- 
birge,  3000  m..  Herbarium  Otto  Kuntze,  type  coll.  (F). 


Vol.  XXV]  MUNZ :  THE  GEXUS  FUCHSIA  83 

(82)  Fuchsia  macrantha  Hooker 
(Platel4,  fig.  73) 
Fuchsia  macrantha  Hook.,  Bot.  Mag.,  72  :pl.  4233, 1846. 

Low,  trailing  or  straggling,  or  epiphytic,  few-branched,  the  twigs  2.5-6 
mm.  thick,  pubescent  to  villous,  with  some  hairs  gland-tipped,  freely  ex- 
foliating; leaves  largely  lacking  at  anthesis,  alternate,  elliptic-ovate,  acute, 
subentire,  rounded  or  subcordate  at  base,  pubescent  above,  pilose  beneath 
especially  on  veins  and  when  young,  with  6-7  principal  lateral  veins  on  each 
side  of  midrib,  blades  4-9  cm.  long,  3-6  cm.  wide ;  petioles  1-2  (3.5)  cm.  long, 
more  or  less  pilose,  1-2  mm.  wide ;  stipules  lance-deltoid,  0.5  mm.  long,  decidu- 
ous ;  flowers  crowded  on  short  branchlets ;  pedicels  slender,  pubescent,  1-2  cm. 
long;  ovary  linear,  pubescent,  6-12  mm.  long,  1.5-3.5  mm.  thick;  hypanthium 
rose-red.  to  coral  or  scarlet,  subcylindric  to  cylindro-fusiform,  6-12  cm.  long, 
3-8  mm.  wide  at  base,  then  narrower  and  very  gradually  ampliate  until  5-12 
mm.  wide  below  the  mouth,  8-ribbed,  finely  pubescent,  with  some  hairs  gland- 
tipped,  villous  within  near  base ;  sepals  ovate  to  lance-ovate,  divergent,  1.5-2.5 
cm.  long,  0.7-1.2  cm.  wide,  tips  greenish  yellow,  somewhat  pubescent  on  both 
surfaces,  obtusish,  connate  about  half  their  length;  petals  0;  episepalous 
stamens  about  three-fourths  the  sepals,  alternate  ones  slightly  shorter;  fila- 
ments glabrous ;  anthers  yellow,  2.5-4.5  mm.  long ;  style  pubescent  just  above 
the  ovary,  slender,  exserted  5-20  mm.  beyond  the  calyx;  stigma  conico- 
spherical,  greenish,  2-A  mm.  long,  scarcely  lobed. 

Type  locality,  Cundimarca,  Peru.  Material  seen,  PERU:  locality  not  given, 
Pavon  1826,  as  apetala,  Geneva  specimen  (photo  F,  POM)  ;  Peru,  mixed  with 
apetala,  Buiz  and  Pavon,  Madrid  Herb.  (F).  Huanuco:  Villcabamba,  haci- 
enda on  Rio  Chinchao,  1800  m.,  Machride  4970  (F,  GH)  ;  Tambo  de  Vaca, 
4000  m.,  Machride  4901  (F,  GH,  US).  Apurimac :  prov.  de  Abancay,  bosques 
del  Ampay,  3200-3500  m.,  C.  Vargas  C.  1000  (GH).  Dept.  not  known :  Lucu- 
mayo  Valley,  1800-3600  m..  Cook  and  Gilbert  1312  (US).  Machride  4970  and 
Vargas  1000  tend  to  have  the  hypanthium  somewhat  fusiform  and  for  a  time 
seemed  to  me  worth  recognizing  as  a  separate  entity,  but  the  material  is 
insufficient. 

(83)  Fuchsia  cestroides  Schulze-Menz 

(Plate  14,  fig.  74) 

Fuchsia  cestroides  Schulzb-Menz,  Notizbl.  Bot.  Gart.,  Berlin,  15:317,  1940;  Macbride, 
Field  Mus.  Nat.  Hist.,  Bot.  13,  part  4,  no.  1 :550,  1941. 

Erect  shrub,  3  m.  tall,  the  young  twigs  densely  puberulent  to  subpilose, 
1.5-3  mm.  thick,  somewhat  reddish  in  color;  leaves  present  at  anthesis,  mostly 
ternate,  not  crowded,  firm-membranaceous,  elliptic-obovate  to  oblong-ovate, 
obtuse  to  acute  at  base,  acuminate  to  acute  at  apex,  plainly  and  rather  closely 
serrulate,  subglabrous  to  hispidulous-puberulent  above,  paler  and  short- 
pilose  beneath  especially  on  the  veins,  principal  lateral  veins  14-16  on  each 


84  CALIFORNIA  ACADEMY  OF  SCIENCES  [Proc.  4th  Ser. 

side  of  midrib,  submarginal  scarcely  developed ;  leaf -blades  4-8  (10)  em.  long, 
2-3.5  (4)  cm.  wide ;  petioles  3-6  mm.  long,  pubescent ;  stipules  subulate,  often 
reflexed,  about  1  mm.  long,  deciduous;  flowers  numerous  in  short  lateral 
racemes  or  terminal  panicles;  bracts  lanceolate  to  linear,  pilose,  3-15  mm. 
long;  pedicels  pilose,  slender,  3-5  mm.  long;  ovary  fusiform,  pilose,  2-A  mm. 
long;  hj-pantliium  dark  red,  17-18  mm.  long,  1-1.5  mm.  wide  at  base,  then 
somewhat  narrowed  until  gradually  widened  to  4  mm.  at  summit,  pilose  with- 
out, glabrous  within ;  sepals  lanceolate,  6-7  mm.  long,  2.5  mm.  wide,  pilose  on 
backs,  somewhat  divergent,  acute;  petals  0;  stamens  7-8  and  5-6  mm.  long, 
exserted;  anthers  1  mm.  long;  style  glabrous,  exceeding  sepals  by  6-7  mm. ; 
stigma  narrow,  about  1  mm.  long ;  berry  apparently  ellipsoid,  at  least  6  mm. 
long. 

Type  locality,  below  Frias,  prov.  Ayabaca,  Piura,  Peru,  at  1100-1200  m., 
the  type  collection,  Weherhauer  6423  (F,  GH,  US).  This  species  is  not  at  all 
close  to  the  other  apetalous  ones,  and  probably  should  not  be  placed  with 
them.  It  differs  in  its  many  veins,  smaller  flowers,  etc.,  and  suggests  in  general 
appearance  F.  Lehmannii. 

Section  6.  SCHUFIA  (Spach)  Munz  new  section 

Schufia,  as  genus,  Spach,  Ann.  Sci.  Nat.,  (2),  4:177, 1835. 
Schufia,  as  subsection,  Endlicher,  Gen.  PI.,  1193, 1840. 

Flowers  erect  in  a  terminal  cymose  panicle,  arranged  subtrichotomously. 
Leaves  opposite  or  verticillate.  Nectary  fused  with  base  of  hypanthium. 
One  species,  F.  arhorescens  Sims. 

(84)  Fuchsia  aborescens  Sims 
Fuchsia  arborescens  Sims,  Curtis  Bot.  Mag.,  53:2620,  1826. 

Low  shrub  to  small  tree  up  to  8  m.  tall,  sometimes  epiphytic,  glabrous  or 
nearly  so ;  branchlets,  petioles  and  veins  sometimes  tinged  with  red ;  leaves 
opposite  or  ternate,  oblong-oblanceolate  to  oblong-elliptic,  acute  or  acuminate 
at  both  ends,  paler  beneath  than  above,  entire  to  serrulate,  blades  3-20  cm. 
long,  1-7  cm.  wide,  with  9-13  principal  lateral  veins;  flowers  numerous, 
crowded  in  corymbose  panicles  5-25  cm.  long  and  equally  wide,  lower  branches 
of  inflorescence  with  few  reduced  leaves,  upper  floriferous  ones  with  much 
reduced,  deciduous,  subulate  bracts;  pedicels  reddish,  slender,  erect,  5-15 
(25)  mm.  long ;  hypanthium  tubular  to  funnelform,  rose  to  magenta,  3-5  mm. 
long,  1-3  mm.  wide  at  summit  in  pressed  specimens ;  sepals  reddish  to  wine- 
purple,  linear  to  lance-oblong,  not  apiculate  in  bud,  3-7  mm.  long,  1-2.5  mm. 
wide,  spreading-reflexed  at  anthesis;  petals  lavender  or  lilac,  lance-ovate, 
two-thirds  as  long  as  sepals,  spreading;  episepalous  stamens  exceeding  sepals, 
4-8  mm.  long,  epipetalous  two-thirds  as  long;  filaments  pinkish  lavender, 
anthers  purplish,  1-1.6  mm.  long;  style  just  longer  than  stamens,  filiform, 
glabrous ;  stigma  purplish,  conic,  about  0.5  mm.  long ;  berry  purplish,  covered 
with  blue  bloom,  subglobose,  7-10  mm.  thick. 


Vol.  XXV]  MUNZ :  THE  GENUS  FUCHSIA  85 

Key  to  Forms  of  Fuchsia  arborescens 

A.  Sepals  4.5-7  mm.  long;  liypanthium  4.5-8  mm.  long. 

B.  Sepals  1.5-2.5  mm.  wide;  liy2)antliium  2-3  mm.  wide 84a.  f.  typica. 

BB.  Sepals  1-1.5  mm.  Avide;  hypanthium  1  mm.  wide 84b.  f.  tenuis. 

A  A.  Sepals  3-4.5  mm.  long ;  hypanthium  3-4  mm.  long , 84c.  f .  parva. 

(84a)  Fuchsia  arborescens  Sims  forma  typica  new  name 
(Plate  14,  fig.  75) 

F.  arborescens  Sims,  Curtis  Bot.  Mag.,  2G20,  1826;  Lindl.,  Bot.  Eeg.,  11:943,  1826. 

Schufia  arborescens  Spach,  Ann.  Sci.  Nat.,  (2),  4:177,  1835. 

F.  hamellioides  Moc.  and  SessiS  ex.  G.  Don,  Gen.  Syst.,  2 :677,  1832,  in  synon.  Have  seen  no 

material. 
F.  syringae flora  Care.,  Eev.  Hortic,  1873 :  311,  1873  ;  judging  from  plate. 
F.  arborea  SESst  and  Moc,  PI.  N.  Hispan.,  ed.  1,  58, 1887-'90  ;  type  seen. 
F. arborescens  var.  (?)  megalantha  Donnell  Smith,  Bot.  Gaz.,  18:2,  1893.  Type  seen; 

agrees  Avell  in  broad  sepals,  etc.,  with  Sims'  plate. 
F.  Liebmanni  L6vL.,  Bull.  Georgr.  Bot.,  22  :24,  1912.  Photograph  of  type  seen. 

Hypanthium  4.5-8  mm.  long,  2-3  mm.  wide  at  summit ;  sepals  4.5-7  mm. 
long,  mostly  2-2.5  mm.  wide. 

Type  locality,  Mexico.  Ranging  at  altitudes  of  1000-2500  m.,  from  Morelos 
and  Guerrero  to  Panama ;  the  most  common  form  of  the  species.  In  this  treat- 
ment of  F.  arborescens,  forms  rather  than  varieties  are  being  recognized, 
since  there  is  no  geographical  correlation  and  the  intergradation  is  so  com- 
plete. Representative  material  of  f .  typica  is,  MEXICO,  without  definite  local- 
ity, '^F.  arliorca,"  Sesse,  Mocino,  Castillo  y  Maldonado  5216  (F).  Morelos, 
mts.  above  Cuernavaca,  Pringle  6825  (F,  GH,  MICH,  NY,  POM,  US).  Guer- 
rero, Sierra  Madre  del  Sur,  Distrito  Mina,  2d  ridge  west  of  Petlacala,  Mexia 
9052  (GH,  NY) .  Vera  Cruz,  Xalapa,  Galeotti 3035  (NY) .  Puebla,  St.  Vincent, 
Nicolas  11  (F,  NY) ;  Puebla,  Bro.  Arscne  in  1909  (US) .  Oaxaca,  Sierra  de  San 
Felipe,  Pringle  6242  (F,  NY,  GH,  CAS,  POM,  US)  ;  Barranca  de  San  Luis, 
Conzatti  1538  (US).  GUATEMALA:  Alta  Vera  Paz,  near  San  Jose,  s.e.  of 
Tactic,  Standley  69660  (F),  Tactic,  von  Tuerckheim  under  J.  D.  Smith  8395 
(F,  NY,  US).  Quezaltenango,  Volcan  de  Santa  Maria,  Steyermark  33616 
(F)  ;  Cumbre  de  Tuilacan,  Standley  67808  (F)  ;  Volcan  de  Zunil,  Skutch  877 
(F).  Chimaltenango,  Chichavac  Tecpan,  Skutch 255  (US)  ;  Chichavac,  Skutch 
59  (US)  ;  Las  Calderas,  Standley  60074  (F).  Sacatepequez,  Volcan  de  Agua, 
Maxon  and  Hay  3745  (US),  Standley  65066  (F)  ;  Volcan  de  Acatenango, 
J.  D.  Smith  2469,  type  no.  var.  megalaiitha  (GH,  US).  COSTA  RICA:  Ala- 
juela,  Volcan  de  Poas,  Cufodonti  601  (F).  San  Jose,  La  Hondura,  Standley 
36587  (US);  between  Guayabillos  and  Cabeza  de  Vaca,  Dodge  and  Thomas 
4936  (F,  MICH,  US)  ;  La  Palma,  Tonduz  under  J.  D.  Smith  7410  (F,  GH, 
US).  Cartago,  Volcan  de  Turrialba,  Pittier  under  J.  D.  Smith  7508  (F,  US)  ; 
Volcan  de  Irazu,  Oersted,  type  F.  Liebmannii  at  Copenhagen  (photographs 
F,  POM).  PANAMA :  Chiriqui,  el  Boquete,  Pittier  2974  (US)  ;  Casita  Alta, 
Volcan  de  Chiriqui,  Woodson,  Allen  and  Seihert  797  (MO,  POM). 


86  CALIFOEXIA  ACADEMY  OF  SCIEXCES  [Proc.  4th  Ser. 

(84b)  Fuchsia  arborescens  Sims  forma  tenuis  new  form 

Hypanthium  tubular,  5-6  mm.  long,  1  mm.  wide;  sepals  linear,  5-6  mm. 
long,  1-1.5  mm.  wide. 

Hypanthium  tubulosum,  5-6  mm.  longum,  1  mm.  latum ;  sepalis  linearibus,  5-6  mm.  longis, 
1-1.5  mm.  latis. 

Type :  Vara  Blanca  de  Sarapiqui,  north  slope  of  Central  Cordillera,  be- 
tween Poas  and  Barba  Volcanoes,  Costa  Rica,  alt.  1740  m.,  Oct.,  1937,  A.  F. 
Skutch  3357  (US  1643234);  isotype  (NY).  With  about  some  range  as  f. 
typica.  Representative  material,  MEXICO  :  Michoacan  and  Guerrero,  Sierra 
Madre,  Langlasse  872  (F,  GH,  US).  Orizaba,  Botteri  (F,  GH).  Chiapas,  San 
Cristobal,^.  W.  Nelson  3236a  (US);  Mt.  Pasitar,  Matuda  1695  (MICH). 
GUATEMALA:  Alta  Vera  Paz,  Coban,  von  Tuerckhcim  II  713  (F,  US). 
Quezaltenango  :  Fuentes  Georginas,  w.  slope  Volcan  de  Zunil,  Standley  67493 
(F).  Solola,  Volcan  de  Atitlan,  Hatch  and  Wilson  in  1936  (F).  Chimalte- 
nango :  Chichavac,  Skutch  750  (F).  PANAMA :  Chiriqui :  Casita  Alta,  Volcan 
de  Chiriqui,  Woodson,  Allen  and  Seihert  911  (MO,  POM);  El  Boquete, 
Maxon  5015  (NY,  GH,  US).  COLOMBIA:  Dept.  Boyaca :  Labranzgrande, 
1150  m.,  B.  Guevara  Amortegui  389,  in  1932  (F,  US).  Since  this  is  the  only 
collection  I  have  seen  from  south  of  Panama,  I  cannot  help  wonder  whether 
it  represents  an  introduction  or  not. 

Intergrades  with  f .  typica  are  numerous ;  among  such  may  be  cited,  Oaxaca  : 
Coyula  a  Cuyamecalco,  Cancim  2438  (US)  ;  Chiapas,  Mt.  Orlando,  Matuda 
0437  (F,  MICH,  US)  ;  Vera  Paz  in  Guatemala:  Chucaneb,  /.  D.  Smith  182 
(F,  GH,  NY,  US)  ;  Huehuetenango,  Yalambofroch,  Seler  2869  (GH,  US)  ; 
San  Jose  in  Costa  Rica :  Between  Aserri  and  Tarbaca,  Standley  34100  (US). 

(84c)  Fuchsia  arborescens  Sims  forma  parva  Munz,  new  form 

F.  paniculata  Lindl.,  Gard.  Chron.,  1856:301,  1856.  Type  not  seen,  but  the  "much  smaller 
flowers"  of  the  description  would  probably  mean  this. 

HjT)anthium  funnelform,  3-4  mm.  long,  1.5  mm.  wide;  sepals  3-4  mm. 
long,  1  mm.  wide. 

Hypanthium  3-4  mm.  longum,  1.5  mm.  latum ;  sepalis  3-4  mm.  longis,  1  mm.  latis. 

Type :  Finca  Pireneos,  below  Santa  Maria  de  Jesus,  Dept.  Quezaltenango, 
Guatemala,  March  11,  1939,  Standley  68287,  Pomona  College  Herb.  No. 
254648;  isotype  Field  Museum  988800.  With  much  the  same  range  as  the  other 
forms;  for  example,  MEXICO  :  Oaxaca,  Tonaguia,  Galcotti  3035  (US).  Vera 
Cruz,  Jalapa,  Walters  (NY)  ;  Barranca  de  Tenampa,  Zaeuapan,  Purpus  7647 
(NY,  US).  Chiapas,  Volcan  de  Tacana,  Matuda  2924  (MICH)  ;  Cerro  del 
Boqueron,  Purpus  6970  (NY,  US).  GAUTEMALA:  Quezaltenango,  Fuentes 
Georginas,  Volcan  de  Zunil,  Standley  67356  (F,  POM)  ;  Pirenos  above  San 
Felipe,  Standley  68287  (F,  POM).  Quiche,  San  Miguel  Uspantan,  Heyde  and 
Lux  under  J.  D.  Smith  3000  (US) .  Alta  Vera  Paz :  Coban,  von  Tuerckheim  in 
1908  (NY).  Sacatepequez,  Volcan  de  Agua,  Kellerman  in  1905  (US).  Zacapa, 


1L 


Vol.  XXV]  MUNZ :  TEE  GENUS  FUCHSIA  87 

Sierra  cle  las  Minas,  Steyermark  29996  (F).  Chiquimula,  Montana  Norte  to 
El  Jntal,  Steyermark  31042  (F).  COSTA  RICA:  Alajuela,  La  Yentolera, 
Volcan  de  Poas,  Standley  34580  (US).  PANAMA:  Chiriqui,  Rio  Chiriqui 
Viejo,  Allen  1351  (MO,  NY,  GH,  POM) ,  Vi^liUe  179  (MO,  POM) . 

Intergrades  with  f.  tenuis  are,  MEXICO,  Chiapas,  Mt.  Pasitar,  Matuda 
1695  (MICH,  NY).  COSTA  RICA  :  San  Jose,  Laguna  de  la  Chonta,  Standley 
42260  (US)  ;  Cartago,  Volcan  de  Irazu,  La  Esperanza,  Standley  35374  (US). 

Section  7.  ENCLIANDRA  (Zucc.)  Endlicher 

EncUandra  (Zucc.)  Endlicher,  Gen  PL,  1183,  1840;  Benth.  and  Hook.,  Gen.  PL,  1,  part 

3:791,1862. 
As  genus,  Zuccarini,  Abh.  Akad.  Muenchen,  11:335,  1837;  spelled  Eucliandra,  Steud., 

Nom.ed.,  2,  1:600,  1840. 
Brebissonia  Spach,  Hist.  Nat.  Veg.  Phan.,  4:401,  1835;  Ann.  Sci.  Nat.,  (2),  4:175,  1835; 

as  subsect.,  Endlicher,  Lc. 
Lyciopsis  Spach,  I.e.,  176;  as  subsect.,  Endlicher,  I.e. 
Myrinia  Lilja,  FL  sverig.  Vexter.,  Suppl.  1 :25,  1840. 

Flowers  polygamous.  Petals  flat.  Stamens  short,  equal,  the  epipetalous  re- 
flexed  into  the  hypanthium.  Berry  few-seeded.  Slender  shrubs  with  opposite 
leaves,  small  solitary  axillary  flowers. 

Type  species.  Fuchsia  EncUandra  Steud. 

Key  to  Species  of  Section  ENCLIANDEA 
A.  Hypanthium  obconic;  sepals  more  than  half  as  long  as  hypanthium,  generally  reflexed 
at  anthesis. 
B.  Leaf -blades  mostly  less  than  2  em.  long,  rounded  to  obtuse  at  base,  obtuse  to  acute 
at  apex ;  petioles  not  more  than  1  em.  long. 

C.  Hypanthium  4-6  mm.  long ;  sepals  4-5  mm.  long 85.  F.  thymifolia. 

CO.  Hypanthium  1-3  mm.  long;  sepals  1.5-3  mm.  long. 

D.  Sepal-tips  eonnivent  in  bud ;  hypanthium  2-3  mm.  long. 

E.  Petals  2-3  mm.  long;  sepals  2-3  mm.  long.  Hidalgo  to  Miehoacan. 

86.  F.  Pringlei. 
'    EE,  Petals  1  mm.  long ;  sepals  1.5  mm.  long,  Volcan  de  Colima. 

87.  F.  colimae. 
DD.  Sepal-tips  spreading  in  bud;  hypanthium  1-2  mm.  long;  petals  1-1.5  mm, 

long.  Mexico  to  Chiapas 88,  i^,  minimiflora. 

BB.  Leaf -blades  mostly  2-5  (7)  em.  long,  acute  at  base,  subaeuminate  at  apex;  petioles 
1-3  cm.  long. 
C.  Hypanthium  longer  than  wide;  stamens  included;  young  growth  subglabrous 

to  pubescent 89.  i^.  tacanensis. 

CC.  Hypanthium  not  longer  than  wide  in  pressed  flowers ;  stamens  evident ;  young 

growth  fine-strigulose 90.  F.  STcutchiana. 

AA.  Hypanthium  subcylindric ;  sepals  usually  not  more  than  half  the  hypanthium,  ascending 
to  somewhat  spreading  at  anthesis. 
B.  Leaves  subeoriaceous,  frequently  conspicuously  serrulate. 

C,  Plant  quite  glabrous ;  hypanthium  5-6  mm.  long,  1,5-2,5  mm.  wide  at  summit ; 

sepals  3-5  mm.  long.  Guerrero  to  Chiapas 91.  F.  iacillaris. 

CC.  Plant  puberulent  on  young  twigs,  hypanthia,  etc. 

D.  Leaf-blades  3-8  (10)  mm.  long;  hypanthium  3-4  mm.  long;  sepals  2-2.5 
mm.  long ;  petals  1.5-2  mm.  long.  Hidalgo,  Vera  Cruz,  Mexico, 

92.  F.  minutiflora. 


88  CALIFORNIA  ACADEMY  OF  SCIENCES  [Proc.  4th  Ser. 

DD.  Leaf -blades  7-20  or  more  mm.  long. 

E.  Hypanthium  5-9  (11)  mm.  long;  sepals  2.5-5  mm.  long. 

F.  Ilypantliium  7-9   (11)   mm.  long,  2.5-5   (6)   mm.  wide  in  the 

perfect  flowers.  Hidalgo  to  Guatemala 93.  F.  microphylla. 

FF.  Hypanthium  5-7   (9)  mm.  long,  1-2  mm.  wide.  Costa  Eica  to 

Panama 94.  F.  Hemsleyana. 

EE.  Hypanthium  3-4  mm.  long,  1  mm.  wide;  sepals  2  mm.  long.  Chiapas. 

95.  F.  striolata. 
BB.  Leaves  membranaceous,  often  subentire. 

C.  Leaf -blades  mostly  less  than  3  cm.  long,  usually  pilose  beneath ;  sepals  2-3 
mm.  long;  petals  0.6-2  mm.  long.  Michoacan  to  Costa  Eica. 

96.  F.  michoacanensis. 
CC.  Leaf-blades  not  pilose  beneath,  or  if  so,  larger. 

D.  The  leaf-blades  8-15  (20)  mm.  long;  hypanthium  of  staminate  flowers 

6-8  mm.  long,  2-3  mm.  wide.  Hidalgo  to  Oaxaca 97.  F.  Encliandra. 

DD.  Leaf -blades  20-60   (100)   mm.  long;  hypanthium  of  staminate  flowers 
mostly  8-10  mm.  long,  2.5-3.5  mm.  wide. 
E.  Hypanthium  of  pistillate  flowers  1-3  mm.  wide;  petals  red;  leaves 
pubescent  or  pilose  beneath. 
F.  Petioles  3-10  (12)  mm.  long;  branchlets  pubescent;  leaves  sub- 
glabrous  to  strigose  above,  pilose  beneath.  State  of  Mexico  to 

Oaxaca 98.  F.  cylindracea. 

FF.  Petioles  15-30   (40)   mm.  long;  branchlets  puberulent;  leaves 
pubescent  on  both  surfaces.  Oaxaca  to  Guatemala. 

99.  F.  tetradactyla. 
EE.  Hypanthium  of  pistillate  flowers  barely  1  mm.  wide ;  petals  appar- 
ently white;   leaves  subglabrous  to  minutely  puberulent  on  both 
surfaces.  Jalisco  to  Morelos 100.  F.  Mexiae. 

(85)  Fuchsia  thymifolia  H.B.K. 

(Plate  15,  fig.  76) 

Fuchsia  fhijmifolia  H.B.K.,  Nov.  Gen.  et  Sp.,  6:104,  pi.  535,  1823.  F.  ihymifolia  DC,  Prodr., 
3:37,  1828;  Lindl.,  Bot.  Eeg.,  15:1284,  1829;  Sweet,  Brit.  Fl.  Gard.,  4:25,  1829;  Hem- 
SLEY,  Bot.  Biol.  Centr.  Amer.,  1:460,  1880;  Standley,  Contr.  U.  S.  Nat.  Herb.,  23:1079, 
1924. 

Lopezia  thymifolia  "Willd.,  herb.  Link,  Jahrb.,  I.  3:25;  ex  Link  in  Schult.,  Mant.,  1:50, 
according  to  H.B.K.,  I.e., 

Lyciopsis  thymifolia  Spach,  Ann.  Sci.  Nat.,  (2)  4:176,  1835. 

Encliandra  thymfolia  Lilja,  Linnaea,  15:262, 1841. 

Fuchsia  alternans  Moc.  and  SESsfi,  Fl.  Mex.  Icon,  ined.,  ex  DC,  Prodr.,  3  :37, 1828,  in  synon. 

F.  parviflora  Lindl.,  Bot.  Eeg.,  13:1048,  1827.  Type  not  seen,  but  the  figure  given  is  good. 

F.  ovata  Sess£  and  MociNO  ex  DC,  Prodr.,  3 :37,  1828,  in  synon.  Not  seen. 

Shrub  about  1  m.  tall,  openly  branched,  with  slender  puberulent  somewhat 
reddish  twigs,  the  ultimate  ones  scarcely  1  mm.  thick;  leaves  opposite  to  alter- 
nate, not  crowded,  elliptic-ovate  to  round-ovate,  subentire,  rounded  to  obtuse 
at  base,  obtuse  to  acute  at  apex,  membranaceous,  paler  beneath  than  above, 
minutely  pubescent  to  subglabrous  above,  minutely  pubescent  beneath,  some- 
what ciliate,  the  blades  0.7-2  (3)  cm.  long,  0.5-1.6  cm.  wide,  with  3-5  pairs 
lateral  veins ;  petioles  very  slender,  puberulent,  3-10  mm.  long,  often  reddish ; 
stipules  subulate,  somewhat  glandular,  scarcely  1  mm.  long ;  flowers  solitary, 


Vol.  XXV]  MUNZ :  THE  GENUS  FUCHSIA  89 

axillary,  white  to  pink,  darker  in  age ;  pedicels  puberulent,  filiform,  5-10  (12) 
mm.  long ;  hypanthium  puberulent  without  and  within,  ob conic,  4-6  mm.  long, 
2-3  mm.  wide  at  summit  in  pressed  specimens,  whitish  to  reddish  (especially 
in  age)  ;  sepals  ovate,  puberulent,  about  4  mm.  long,  white  and  spreading- 
reflexed  at  anthesis,  later  red,  the  subulate  tips  connivent  in  bud,  1  mm.  long; 
petals  ovate-oblong,  spreading,  4  mm.  long,  white  becoming  purplish  red,  ob- 
tuse, often  undulate ;  stamens  subsessile,  the  episepalous  ones  exserted  from 
hypanthium,  1.5  mm.  long,  the  epipetalous  included,  1  mm.  long ;  style  white, 
glabrous,  exserted,  6-9  mm.  long  from  base  of  hypanthium ;  stigma  greenish, 
the  lobes  erect  or  spreading,  barely  0.5  mm.  long ;  berry  globose,  glabrate  to 
puberulent,  4-6  mm.  long. 

Type  locality,  Patzcuaro,  Michoacan,  Mexico.  Kanging  through  several 
states  of  south  central  Mexico;  representative  material,  without  locality, 
fragment  Bonpland  4106  (F)  ;  ex  horto  bot.  petropolitano,  seeds  from  Mexico, 
in  1828  (NY);  Coulter  176  (NY).  Hidalgo,  Sierra  de  Pachuca,  Rose  and 
Painter  8023  (US),  6737  (GH,  NY,  US),  PnngrZe  9472  (G,  US  in  part),  5754 
(F,  GH,  MICH,  NY,  US),  Eose,  Painter  and  Rose  8859  (GH,  NY,  US)  ;  El 
Chico,  Lyonnet  534  (US).  Michoacan,  Quinceo  near  Morelia,  Arsene  3228 
(GH,  US)  ;  Cerro  Azul,  Arsene  2453  (US) .  Mexico,  Salazar,  Rose  and  Painter 
7035  (US)  ;  Federal  District,  Cerro  Xitle,  Russell  and  Souviron  175  (US)  ; 
Eslava,  Rushij  342  (NY,  US)  ;  Pedregal,  Pringle  7206  (GH,  US)  ;  Contreras 
Canyon,  Munz  15001  (POM).  Vera  Cruz,  Pedregal  Naolingo,  Purpus  296 
(US).  Sinaloa,  El  Carrizo,  Ortega  6009  (US). 

(86)  Fuchsia  Pringlei  Robinson  and  Seaton 

(Plate  15,  fig.  77) 

Fuchsia  Pringlei  Robins  and  Seat.,  Proe.  Arner.  Acad.,  28:106,  1893;  Standley,  Contr. 
U.  S.  Nat.  Herb.,  23:1079, 1924. 

Habit  and  pubescence  of  F.  thymifolia,  but  leaves  closer  together,  blades 
mostly  less  than  1  cm.  long,  0.3-0.7  (1)  cm.  wide;  petioles  2-5  (7)  mm.  long; 
hypanthium  2-3  mm.  long ;  sepals  2-3  mm.  long ;  petals  2-3  mm.  long ;  style 
4r-o  mm.  long. 

Type  locality,  Patzcuaro,  Michoacan.  In  the  original  description  were 
cited  :  Mts.  near  Patzcuaro,  Pringle  5063 ;  barranca  near  Guadalajara,  Pringle 
5002,  and  Mts.  near  Patzcuaro,  Pringle  4140.  Since  the  first  two  cited  were  in 
fruit  and  only  the  last  had  flowers,  it  should  be  taken  as  the  type  number. 
Fruiting  specimens  alone  can  be  distinguished  from  F.  thymifolia  only  with 
difficulty.  The  species  ranges  with  F.  thymifolia.  Specimens  seen,  MEXICO  : 
"Fuchsia  microphylla  N.  ic."  Sesse,  Mocino,  Castillo,  y  Maldoyiado  5210  (F)  ; 
Salto  de  Agua,  Purpus  1780  (POM,  US) .  Hidalgo,  Pachuca,  Rose  and  Hough 
4452  (GH,  US).  Mexico,  Comunidad,  Temascaltepec,  Hinton  1814  (GH,  US)  ; 
Popocatepetl,  Purpus  3059  (NY,  US) ;  Ixtaccihuatl,  Purpus  370  (US)  ;  Cima, 
Rose  and  Painter  7189  (GH,  US)  ;  Federal  district,  Sierra  de  Ajusco,  Pringle 
6412  (CAS  in  part,  F  in  part,  MO,  NY,  US  in  part),  11988  (F,  GH,  MICH. 


90  CALIFOEXIA  ACADEMY  OF  SCIENCES  [Proc.  4th  Ser. 

US).  Vera  Cruz,  Acultzinco,  Matuda  1097  (F,  MICH,  NY);  Maltarata, 
Matilda  1218  (F,  MICH) .  Morelos,  Serrania  de  Ajusco,  Pringle  7570  (POIM)  ; 
Ti^s  Marias,  Orcutt  3754  (F,  US) .  Michoacan,  Morelia,  Cerro  Azul,  Arscne  in 
1909  (F,  GH)  ;  Patzcuaro,  Pringle  4140,  type  no.  (F,  GH,  CAS,  NY,  US)  ; 
Tarascon,  Pringle  13463  (US).  Tepie,  between* Santa  Gertrudis  and  Santa 
Teresa,  Rose  2117  (US).  Jalisco,  west  of  Bolafios,  Rose  2964  (US),  not  typi- 
cal ;  flowers  white. 

(87)  Fuchsia  minimiflora  Hemsley 

(Plate  15,  fig.  79) 

Fuchsia  minimiflora  Hemsl.,  Bot.  Biol.  Centr.  Amer.,  1:458,  1880;  5:  pi.  28,  fig.  1,  1880; 
Standley,  Contr.  U.  S.  Nat.  Herb.,  23  :  1079,  1924. 

Shrub  1-4  m,  tall,  loosely  branched  with  long  very  slender  reddish  crisp- 
puberulent  twigs;  leaves  oposite  or  alternate,  ovate  to  elliptic-ovate,  acute  at 
apex,  i^ounded  to  obtuse  at  base,  subentire  to  serrulate,  paler  beneath,  sub- 
giabrous  to  minutely  crisp-puberulent,  the  blades  mostly  1-2  cm.  long,  0.5- 
1.2  cm.  wide ;  petioles  very  slender,  puberulent,  2-5  mm.  long ;  flowers  solitary, 
axillary ;  pedicels  filiform,  puberulent,  4-12  mm.  long ;  hypanthium  obconic, 
1-2  mm.  long,  whitish  to  reddish,  puberulent  without;  sepals  lanceolate, 
reflexed-spreading,  whitish  to  reddish,  1.5-2.5  mm.  long,  the  tips  apiculate, 
spreading  in  bud ;  petals  oblong-elliptic,  white  to  red,  less  than  1.5  mm.  long; 
stmens  subsessile,  less  than  1  mm.  long ;  style  glabrous,  2-3  mm.  long ;  stigma 
4-lobed,  the  lobes  spreading,  0.5-1  mm.  long ;  berry  subglobose,  glabrate  to 
puberulent,  4-5  mm.  thick. 

Type  locality,  Chiapas,  Mexico.  A  species  of  south-central  Mexico,  known 
from  but  few  collections,  of  which  I  have  seen  :  MEXICO,  state  not  known  to 
me,  Salto  de  Agua,  Purpus  1780  in  part  (F) .  Mexico,  Cima,  Rose  and  Painter 
7189  (US)  ;  Federal  District,  Sierra  de  Ajusco,  Rose  and  Hay  5521  (US, 
Pringle  6412  in  part(  F,  GH,  MO,  US)  ;  District  of  Temascaltepec,  Hint  on 
1351  (F,  NY) .  Puebla,  Boca  del  Monte,  Piirpus  5828  (F,  GH,  NY,  US) .  Chia- 
pas, without  definite  locality.  Linden  680,  type  coll.,  photo  of  Geneva  specimen 
(F). 

(88)  Fuchsia  colimae  Munz,  new  species 
(Plate  15,  fig.  78) 

Habit  and  puberulence  of  F.  minimiflora;  leaves  orbicular-ovate,  the  blades 
5-12  mm.  long,  4-11  mm.  wide,  obscurely  serrulate,  obtuse  to  rounded  at  both 
ends;  petioles  3-5  mm.  long;  hypanthium  whitish-green,  2-2.5  mm.  long; 
sepals  white,  reflexed-spreading,  1.5  mm.  long,  the  tips  not  free  in  the  bud ; 
petals  white,  elliptic-oblong,  1  mm.  long ;  stigma-lobes  spreading. 

Habitus  et  puberulencia  Fuchsiae  minimiflorae,  foliis  orbiculare-ovatis,  laminis  5-12 
mm.  longis,  4-11  mm.  latis,  obscure  serrulatis,  obtusis  vel  rotundatis  in  extremis  utrisque; 
petiolis  3-5  mm.  longis;  hypanthio  albido,  2-2.5  mm.  longo  ;  sepalis  albidis,  reflexo-expansis, 
1.5  mm.  longis;  petalis  albis,  elliptico-oblongis,  1  mm.  longis;  lobis  stigmatis  expansis. 

Type:  from  Volcano  of  Colima,  Jalisco,  Mexico,  July  13, 1892,  M.  E.  Jones 


Vol.  XXV]  MUNZ :  TEE  GENUS  FUCHSIA  91 

224,  Pomona  College  Herbarium  83711 ;  isotypes  United  States  National 
Herbarium  237441  and  at  New  York  Botanical  Garden.  This  species  is  near 
F.  minimiflora,  but  the  sepals  are  shorter  than  the  hypanthium,  the  tips  are 
not  free  in  the  bud,  and  the  leaves  are  more  rounded. 

(89)  Fuchsia  tacanensis  Lundell 

(Plate  15,  fig.  80) 
Fuchsia  tacanensis  Lundei^l,  Contr.  Univ.  Mich.  Herb.,  4:20, 1940, 

Shrub  up  to  4  m.  high,  open-branched,  the  ultimate  branchlets  slender, 
greenish,  subglabrous  to  pubescent;  leaves  opposite,  the  blades  membranace- 
ous, rhombic-ovate,  abruptly  cuneate  at  base,  acuminate  at  apex,  rather  evenly 
and  coarsely  sinuate-serrulate,  paler  beneath  than  above,  somewhat  strigose 
above,  subglabrous  to  pilose  beneath,  2.5-5  (7)  cm.  long,  1.2-3.5  (4.2)  cm. 
wide,  with  7-9  pairs  of  lateral  veins ;  petioles  slender,  subglabrous  to  pubes- 
cent, 1-2  cm.  long;  pedicels  filiform,  glabrous  to  pubescent,  mostly  1-2  cm. 
long;  flowers  perfect  or  imperfect;  hypanthium  obconic,  3-5.5  mm.  long, 
2-3.5  mm.  wide  at  summit,  glabrous  to  puberulent,  greenish  to  reddish ;  sepals 
reflexed-spreading  at  anthesis,  lanceolate,  acuminate,  usually  somewhat  pu- 
bescent, whitish  or  greenish,  usually  shorter  than  hypanthium,  2.5-6  mm. 
long,  apiculate,  the  tips  1-2  mm.  long,  connivent  in  bud ;  petals  oblong-ovate, 
white  to  pink,  red  in  age,  2-3.5  mm.  long,  more  or  less  emarginate;  stamens 
included,  anthers  about  1  mm.  long ;  style  in  perfect  flowers  4-7  mm.  long ; 
stigma  4-lobed,  scarcely  1  mm.  long ;  fruit  subglobose,  black,  4-5  mm.  thick. 

Type  locality,  Volcan  de  Tacana,  Chiapas,  Mexico ;  the  species  is  known 
from  the  following  collections,  all  from  MEXICO  :  without  locality,  Schiede 
(US)  ;  Morelos,  barranca  above  Cuernavaca,  Pringle  6380  (F,  GH,  CAS, 
MICH,  NY,  US)  ;  Chiapas,  Volcan  de  Tacana,  Matuda  2399  (MICH,  type) ; 
Mt.  Pasitar,  Matuda  1686  (MO,  NY,  MICH,  US).  F.  tacanensis  is  variable  in 
flower-size,  pubescence,  length  of  hypanthium,  etc.,  but  is  distinguished  by  the 
large  thin  toothed  leaves,  wide  hypanthium,  long  sepal-tips,  light-colored 
flowers,  sepals  shorter  than  hypanthium.  Pringle  6380  has  staminate  more 
crowded  flowers,  pedicels  less  than  1  cm.  long. 

(90)  Fuchsia  Skutchiana  Munz,  new  species 

(Platel5,  fig.  81) 

Much  like  F.  tacanensis  in  habit,  foliage,  etc.,  but  finely  strigulose  on  young 
branchlets,  hypanthia,  sepals;  leaves  more  finely  strigose;  petioles  2-3  cm. 
long;  pedicels  1.5-3  cm.  long,  strigulose;  hypanthium  not  longer  than  wide 
in  pressed  specimens,  2-3  mm.  long,  slightly  wider ;  sepals  at  least  as  long  as 
hypanthium,  reflexed,  2.5-4  mm.  long,  white  to  pink,  aging  red,  the  tips  1-1.5 
mm.  long;  petals  about  as  long  as  sepals,  white  to  pink,  aging  red,  emarginate ; 
stamens  not  included,  about  1  mm.  long,  anthers  0.5  mm.  long;  style  4—5  mm. 
long ;  fruit  black,  4-5  mm.  thick. 


92  CALIFOENIA  ACADEMY  OF  SCIENCES  [Proc.  4th  Ser. 

Habitus  et  folia  Fuchsiae  tacanensis;  ramulis,  hypanthiis  sepalisque  strigulosis;  foliis 
strigulosis;  petiolis  2-3  cm.  longis;  pedicellis  1.5-3  cm.  longis,  strigulosis;  hypanthio  2-3 
mm.  longo,  latiore;  sepalis  hypanthio  aequalibus,  reflexis,  2.5-4  mm.  longis,  albis  vel  carneis, 
in  aetate  roseis,  cum  apicibus  1-1.5  mm.  longis;  petalis  sepalis  aequalibus,  emarginatis, 
albis  vel  carneis,  in  aetate  roseis;  staminibus  non  inclusis,  1  mm.  longis;  antheris  0.5  mm. 
longis;  stylo  4-5  mm.  longo;  bacca  nigra,  4^5  mm,  diametro. 

Type:  from  near  Quezaltenang-o,  Quezaltenango,  Guatemala,  at  2500  m., 
July  24, 1934,  A.  F.  Skutch  812  (NY)  ;  isotype  (F) .  Other  material  from  same 
place  and  on  same  date,  Skutch  813  (F)  ;  Chiapas,  Mexico,  Ghieshreght  730, 
in  1864-70  (F,  Gil,  MO). 

(91)  Fuchsia  bacillaris  Lindley 

(PlatelS,  fig.  82) 

Fuchsia  bacillaris  Lindl.,  Bot.  Eeg.,  18:1480,  1832;  F.  bacillaris  in  Curtis,  Bot.  Mag.,  76: 
4506,  1850;  Hemsley,  Biol.  Centr.  Amer.,  Bot.,  1:456,  1880;  Standley,  Contr.  U.  S.  Nat. 
Herb.,  23:1079, 1924. 

Shrub  1-2  m.  tall,  practically  glabrous  throughout,  openly  and  finely 
branched ;  leaves  opposite,  subcoriaceous,  sinuate-denticulate,  paler  beneath 
than  above,  lance-ovate  to  elliptic-ovate,  more  or  less  cuneate  at  base,  acute  to 
obtuse  at  apex,  sometimes  with  line  of  minute  hairs  on  upper  surface  of  mid- 
rib ;  blades  1-3  (4.5)  cm.  long,  0.5-1.2  (1.5)  cm.  wide;  petioles  3-6  mm.  long, 
sometimes  with  lines  of  puberulence ;  flowers  solitary,  axillary,  apparently 
perfect  or  pistillate ;  pedicels  glabrous,  filiform,  5-15  mm.  long ;  hypanthium 
subcylindric,  glabrous,  red,  5-6  mm.  long,  1.5-2.5  mm.  wide  at  summit ;  sepals 
spreading-reflexed  at  anthesis,  rose,  glabrous,  lanceolate,  3-5  mm.  long,  apicu- 
late,  the  tips  connivent  in  the  bud ;  petals  rose,  round-obovate,  refuse,  2.5-5 
mm.  long;  stamens  included,  the  anthers  1-1.5  mm.  long ;  style  8-13  mm.  long ; 
stigma  reddish,  the  4  lobes  tending  to  spread ;  fruit  spherical,  4-5  mm.  thick. 

Type  locality,  Mexico,  seeds  sent  by  J.  S.  Mill,  Esq.  This  species  is  an  un- 
certain entity,  the  exact  location  of  its  origin  being  unknown.  It  has  always 
been  keyed  out  as  being  glabrous.  The  plate  by  Lindley  shows  it  as  having 
relatively  large  flowers  and  is  pretty  well  matched  by  Purpus  6708.  Material 
seen,  MEXICO  :  "seed  from  Mexico,  1829,"  Ex  horto  bot.  Petropolitano  (NY) , 
matching  the  Lindley  plate  and  description  and  is  likely  authentic  material. 
Guerrero:  Mts.  near  Chilpancingo,  E.  W.  Nelson  2258  (US),  with  leaves 
rather  small;  Sierra  Madre,  near  Chilpancingo,  Nelson  2232  (US)  ;  crest  of 
Sierra  Madre,  Langlasse  890  (F,  GH,  US).  Chiapas:  Cerro  del  Boqueron, 
Purinis  6708  (F,  GH,  NY,  US).  GUATEMALA:  Iluehuetenango,  Sierra 
Cuchumatanes,  Skutch  1247  (F),  twigs  slightly  pubescent. 

(92)  Fuchsia  minutiflora  Hemsley 

Fuchsia  minutiflora  Hemsl.,  Diag.  PI.  Nov.,  1:15,  1878;  Bot.  Biol.  Centr.  Amer.,  1:458, 
1880;  5: pi.  28,  1880;  Standley,  Contr.  U.  S.  Nat.  Herb.,  23:1079,  1924. 

Shrub  1-2  m.  tall,  with  very  slender  rather  numerous  branches,  the  ultimate 
ones  scarcely  1  mm.  thick  and  covered  with  a  loose  brownish  puberulence,  the 


Vol.  XXV]  MUNZ:  TEE  GENUS  FUCHSIA  93 

hairs  more  or  less  spreading,  often  with  incurved  tips ;  leaves  mostly  opposite, 
numerous,  mostly  longer  than  the  internodes,  the  blades  thickish,  lance-ovate 
to  -elliptic,  conspicuously  serrulate  with  4-5  teeth  on  each  edge,  glabrous, 
somewhat  paler  beneath,  3-8  (10)  mm.  long,  2-5  (7)  mm.  wide,  obtuse  to  acute 
at  both  ends;  petioles  1-2  (4)  mm.  long;  flowers  solitary,  axillary;  pedicels 
filiform,  sparsely  puberulent,  2-8  mm.  long;  hypanthium  tubular-funnel- 
form,  3-4  mm.  long,  1.5  mm.  thick  at  base,  gradually  ampliate,  subglabrous 
to  puberulent,  apparently  reddish ;  sepals  lanceolate,  acuminate,  apiculate, 
somewhat  spreading  in  anthesis,  2-2.5  mm.  long,  the  tips  connivent  in  the  bud ; 
petals  oblong,  obtuse,  entire,  1.5-2  mm.  long,  apparently  white ;  stamens  very 
small,  almost  included ;  style  glabrous,  scarcely  as  long  as  petals,  stigma  deeply 
4-lobed ;  berry  sub-globose,  subglabrous,  3-4.5  mm.  thick. 

Ket  to  Varieties  of  Fuchsia  minutiflora 

A.  Hypanthium  and  sepals  white;  hypanthium  pubescent 92a.  var.  hidalgensis. 

AA.  Hypanthium  red,  subglabrous 92b.  var.  typica. 

(92a)  Fuchsia  minutiflora  Hemsley,  var.  hidalgensis  Munz,  new  variety 

(Plate  15,  fig.  83) 

Hypanthium  pubescente,  album ;  sepalis  albis. 

Type:  from  Trinidad,  Hidalgo,  Mexico,  at  1850  m.,  April  30, 1904  Pringle 
8808,  Pomona  College  Herbarium  59867  ;  isotypes  (F,  GH,  CAS,  MICH,  NY, 
US),  the  label  on  the  Michigan  specimen  reads  "Honey  Station,  Puebla." 
Material  seen,  Trinidad,  Pringle  8807  (F,  GH,  CAS,  NY,  US)  ;  Zacualtipan, 
Selerl50  (GH,  US). 

(92b)  Fuchsia  minutiflora  Hemsley,  var.  typica  Munz,  new  name 
Fuchsia  minutiflora  Hemsl.,  Diag.  PI.  Nov.,  1 :15,  1878. 

Hypanthium  red,  subglabrous. 

Type  locality,  "in  monte  Orizaba,  alt.  5000  ped.,"  based  on  Linden  631. 
Material  seen,  MEXICO :  without  locality,  Mueller  1339  (NY)  ;  garden  of 
Mrs.  Nuttall,  Mexico  City,  Safford  1511  (US)  ;  Vera  Cruz,  Chinantla,  Lieh- 
mann  3231  (US) ;  Prov.  Jalapa,  com.  Schlechtendal  in  1840,  coll.  Schiede  525 
(GH,  NY)  ;  Aseradero  de  Santa  Cruz,  Mueller  1550  (NY)  ;  Orizaba,  Mueller 
1702  (GH,  NY),  640  (NY)  ;  Maltrata,  Matuda  1358  (MICH,  NY).  Mexico, 
Ixtaccihuatl,  Purpus  160  (POM). 

(93)  Fuchsia  microphylla  H.B.K. 

Fuchsia  microphylla  H.B.K.,  Nov.  Gen.  et  Sp.,  6 :103,  pi.  534,  1823. 

Subshrub  or  shrub,  3-20  dm.  tall,  rather  densely  branched,  the  ultimate 
branchlets  slender,  barely  1  mm.  thick,  with  a  dense  subappressed  puberu- 
lence;  leaves  crowded,  opposite,  subcoriaceous,  lance-oblong,  to  elliptic-obo- 
vate,  usually  sharply  serrulate,  sometimes  subentire,  deep  green  above,  paler 
beneath,  acute  to  obtuse  at  both  ends,  subglabrous  except  sometimes  for  line 
of  puberulence  along  upper  surface  of  midrib,  the  blades  5-20  (25)  mm.  long. 


94  CALIFORNIA  ACADEMY  OF  SCIENCES  [Proc.  4th  Ser. 

3-8  (10)  mm.  wide;  petioles  glabrous  to  puberulent,  2-8  (15)  mm.  long; 
flowers  solitary,  axillary,  perfect  or  pistillate ;  pedicels  appressed-puberu- 
leut,  3-6  (10)  mm.  long;  hypanthiiim  deep  red,  subcylindric,  constricted 
above  the  ovary,  then  usually  7-9  (11)  mm.  long,  2.5-5  (6)  mm.  wide  in 
pressed  specimens,  sometimes  shorter  and  narrower  in  pistillate  flowers,  gla- 
brous to  minutely  strigose  ;  sepals  deep  red,  mostly  4,  sometimes  3,  lanceolate 
to  ovate,  2.5-5  (7)  mm.  long,  acuminate,  usually  apiculate,  the  tips  up  to  1 
mm.  long  and  connivent  in  the  bud,  glabrous  to  strigulose ;  petals  rose,  broadly 
elliptic  to  almost  round,  usually  toothed,  3-5  (7)  mm.  long;  stamens  nearly 
or  quite  included ;  anthers  1.5-2  mm.  long,  sometimes  lacking ;  style  8-15  (20) 
mm.  long;  stigma  reddish,  deeply  4-lobed;  fruit  subglobose,  glabrous  to 
strigulose,  purple-black  when  ripe,  up  to  8  mm.  thick. 

Characterized  by  its  coriaceous  glabrous  usually  serrulate  and  smallish 
crowded  leaves  and  rather  large  flowers,  but  variable;  flowers  small  if  pistil- 
late only. 

Key  to  VAiiiETiEs  of  Fuchsia  microphylla 

A.  Ovary  and  hypantliium  glabrous 93a.  var.  typica. 

A  A.  Ovary  and  lower  part  of  hypantliium  minutely  strigulose 93b.  var.  aprica. 

(93a)  Fuchsia  microphylla  H.B.K.  var.  typica  Munz,  new  name 

(Plate  15,  fig.  84) 

F.  microphylla  H.B.K.,  Nov.  Gen.  et  Sp.,  6:103,  pi.  534,  1823;  Lindl.,  Bot.  Eeg.,  15:1269, 
1829;  Sweet,  Brit.  PI.  Gard.,  ser.  2,  pi.  16,  1829?;  Hemsley,  Bot.  Biol.  Centr,  Amer., 
1 :  458,  1880  in  part ;  Standley,  Contr.  U.  S.  Nat.  Herb.,  23  :  1079,  1924  in  part. 

Brebissonia  microphylla  Spach,  Ann.  Sci.  Nat.,  ser.  II,  4:175,  1835. 

Myrinia  microphylla  Lilja,  F1.  Sverig.  Suppl.,  1:25,  1840. 

F.  gracilis  Sess£  and  Mocino  ex  DC,  Prodr.,  3:37,  1828  in  sjTion. 

F.  mixta  Hemsley,  Diag.  PI.  Nov.,  1:14,  1878;  Biol.  Centr.  Amer.  Bot.,  1:459,  1880;  5:pl. 
28,  fig.  2,  1880.  Based  on  Galeotti  3025  from  Orizaba,  which  I  have  not  seen,  but  which 
is  in  the  range  of  the  typical  form  if  referred  to  F.  microphylla.  I  agree  with  Standi.,  I.e., 
that  such  must  be  the  case. 

Ovary  and  hypantliium  glabrous. 

Type  locality,  Volcan  de  Jorullo,  which  I  believe  is  in  Michoacan,  Mexico. 
Ranging  from  Hidalgo  to  states  of  Mexico,  Michoacan  and  Jalisco.  Material 
seen,  MEXICO  :  without  definite  locality,  Sesse,  Mocino,  Castillo  y  Maldonado 
5197  (F)  ;  Lithrum  vulnerarium,  N.,  Sesse  et  at  1842  (F)  ;  ex  horto  bot.  Petro- 
politano,  introd.  1828  from  Mex.  (NY).  Hidalgo,  El  Chico,  Lyonnet  2204 
(US) ;  between  Real  del  Monte  and  Chico,  Hartweg  285  (NY)  ;  Lagunas 
Zempoala,  Lyonnet  2510  (US).  Vera  Cruz,  Pico  de  Orizaba,  Vacqueria  de 
Jacal,  Lielmann  3230  (US,  GH).  Federal  District,  El  Desierto,  Conviento, 
Russell  and  Souviron  133  (CAS,  US)  ;  La  Cima,  Barnes  and  Land  395  (F) ; 
San  Nicolas,  Valle  de  Mexico,  Bourgeau  974  (GH,  US)  ;  Pass  near  Desierto 
de  los  Leones,  Munz  15062  (POM)  intergrading  toward  aprica;  Contreras, 
Lyonnet  1637  (US).  Mexico,  Salazar,  Pringle  13462  (US),  Rose  and  Painter 
7036  (US)  ;  Montes  de  Gavia,  Asphmd  356  (NY)  ;  Nevada  de  Toluca,  Rose 


Vol.  XXV]  MUNZ :  THE  GENUS  FUCHSIA  95 

and  Painter  7935  (NY,  US)  ;  Sierra  de  las  Cruces,  Pringle  4314  (F,  GH, 
MICH,  NY,  US) ;  Ixtaccihuatl,  Punms  1779  (F,  GH,  NY,  POM,  US)  ;  Meson 
Viejo,  Temascaltepec,  Hinton  1902  (US)  ;  San  Rafael,  Lyonnet  and  Elcoro 
1923  (US).  Morelos,  Popocatepetl,  Purpus  3060  (F,  NY,  GH,  US),  Rose  and 
Hay  6256  (US).  Michoacan,  Mt.  Patamban,  Nelson  6566  (NY,  US);  "in 
monte  ignivomo  JoruUo,  alt.  550  hex.,"  Humboldt  et  al,  type  coll.  (scrap  at 
F).  Jalisco,  volcano  of  Colima,  Jones  225  (POM,  US). 

(93b)  Fuchsia  micropliylla  H.B.K.  var.  aprica  (Lundell),  new  comb. 
F.  aprica  Lundell,  Contr.  Univ.  Mich.  Herb.,  4:18-19,  1940. 

Ovary  and  lower  part  of  hypanthium  minutely  appressed-puberulent. 

Type  locality,  Volcan  de  Tacana,  Chiapas,  Mexico.  Ranging  from  Oaxaca 
and  Chiapas  to  Guatemala.  Representative  material  seen,  MEXICO,  Puebla, 
Tezuitlan,  Orcutt  4001  (F,  MO,  US).  Oaxaca,  Yavezca,  Galeotti  3039  (US)  ; 
Cerro  de  los  Frailes,  between  Teotitlan  del  Camino  and  Huantla  de  Jimenez, 
Schultes  and  Reko  370  (F,  GH) .  Chiapas,  San  Cristobal,  Goldman  985  (US)  ; 
Mt.  Tacana,  Matuda  2378,  type  coll.  aprica  (MICH,  NY).  GUATEMALA: 
Alta  Vera  Paz,  Mts.  east  of  Tactic,  Standley  71428  (F).  Huehuetenango,  San 
Juan  Ixcoy,  Skutch  1069  (F).  Quiche,  San  Miguel  Uspantan,  Heyde  and  Lux 
under  J.  D.  Smith  2999B  (NY,  US)  ;  Nebaj,  Skutch  1711  (F)  pistillate.  San 
Marcos,  Volcan  de  Tajumulco,  Schmidt  1934  (F)  ;  Volcan  de  Tacana,  Steyer- 
mark  36159  (F).  Quezaltenango,  Volcan  de  Santa  Maria,  Steyermark  34187 
(F).  Chimaltenango,  Volcan  de  Acatenango,  Standley  61954  (F) ;  Volcan  de 
Agua,  Johnston  896  (F).  Guatemala,  between  San  Martin  and  Todos  Santos, 
Nelson  3627,  pistillate  (F,  US).  Sacatepequez,  Antigua,  Kellerman  6535  (F)  ; 
Volcan  de  Agua,  Kellerman  7577  (NY). 

This  pubescent  plant  intergr^des  freely  with  F.  michoacanensis  which  has 
thinner  more  membranaceous  leaves  with  coarser  longer  hairs  on  their  lower 
surface,  for  example:  GUATEMALA:  Quezaltenango,  Hartweg  527  (NY)  ; 
Sacatepequez,  Volcan  de  Agua,  Shanyion  3655  (US)  ;  Alta  Vera  Paz,  Coban, 
von  Tuerckheim  II 1295  (F,  MICH,  NY)  ;  Mts.  east  of  Tactic,  Standley  71357 
(F).  Oaxaca,  Cerro  de  la  Virgen,  Conzatti  4352  (US).  In  these  and  many 
others,  one  gets  varying  combinations  of  thick  leaves  and  pilosity. 

(94)  Fuchsia  Hemsleyana  Woodson  and  Seibert 
(Plate  15,  fig.  85) 

Fuchsia  Hemsleyana  Woodson  and  Seibert,  Ann.  Mo.  Bot.  Gard.,  24:194,  1937. 

F.  pulcliella  "Woodson  and  Seibert,  I.e.,  195-196.  Based  on  Seibert  183,  which  I  have  seen. 

Shrub  1-3  m.  tall,  rather  openly  branched,  the  ultimate  branchlets  slender, 
ca.  1  mm.  thick,  minutely  pubescent,  sometimes  strigulose;  leaves  mostly 
opposite,  rather  crowded,  subcoriaceous,  rhomboid-elliptic  to  -obovate,  cuneate 
at  base,  acute  to  obtusish  at  apex,  serrulate  to  sinuate-dentate,  paler  beneath 
than  above,  subgiabrous  except  on  upper  surface  of  midrib  and  along  margin, 
the  blades  7-15  (25)  mm.  long,  4-8  (12)  mm.  wide ;  petioles  sparsely  puberu- 


96  CALIFORNIA  ACADEMY  OF  SCIENCES  [Proc.  iTH  Ser. 

lent,  3-8  (12)  mm.  long;  flowers  solitary,  axillaiy,  perfect  or  pistillate;  pedi- 
cels filiform,  more  or  less  puberulent,  4-6  (8)  mm.  long;  hypanthium  rose, 
tubular,  constricted  above  ovary,  then  5-7  (9)  mm.  long,  1-2  mm.  wide  at 
summit,  glabrous  to  puberulent;  sepals  deltoid-lanceolate,  rose,  spreading- 
reflexed  at  anthesis,  3.5-4  mm.  long,  apiculate,  the  tips  connivent  in  the  bud ; 
petals  rose  or  with  purplish  tinge,  oblong-obovate,  2.5-4  mm.  long,  undulate 
and  rounded  at  apex;  stamens  usually  included;  anthers  1-1.2  mm.  long; 
style  8-10  mm.  long;  stigma  deeply  4-lobed,  exserted,  1  mm.  long;  ovary  sub- 
glabrous  to  strigulose ;  fruit  subglobose,  black-purple,  4-5  mm.  thick. 

Type  locality,  Rio  Chiriqui  Viejo,  near  Monte  Lirio,  Chiriqui,  Panama- 
Found  in  the  mountain  of  Costa  Rica  and  Panama.  Representative  material 
may  be  cited,  COSTA  RICA :  Vara  Blanca  de  Sarapiqui,  Skiitch  3225  (NY). 
Alajuela,  Volcan  de  Poas,  Standley  34629  (VQ),  34620  (US),  Allen  600  (F). 
Cartago,  Cerros  de  Irazu,  Pittier  13048  (US)  ;  Volcan  de  Turrialba,  Standley 
34949  (US).  San  Jose,  La  Guaba,  Valerio  1526  (F)  ;  Laguna  de  la  Clionta, 
Standley  42275,  strigulose  (F,  US)  ;  Cerro  de  Piedra  Blanca,  above  Escasu, 
Standley  32598  (US),  Solis  154  (F)  strigulose;  Finca  La  Cima  above  Los 
Lotes,  Standley  42758  (US);  Cerro  de  Las  Vueltas,  Standley  and  Valerio 
43523  (F,  \^^),  43499  (US)  ;  Laguna  de  la  Escuadra,  n.e.  of  El  Copey,  Stand- 
ley  42078  (F,  US),  42057  (US)  ;  between  Guayabillos  and  Cabeza  de  Vaca, 
Dodge  and  Thomas  4940  (F).  PANAMA:  Chiriqui,  upper  Rio  Chiriqui 
Viejo,  Seibert  162,  type  no.  Hemsleyana  (MO,  NY)  ;  near  Monte  Lirio,  Scibert 
182,  type  pulchella  (MO)  ;  Rio  Chiriqui  Viejo  valley.  White  2  (GH,  POM)  ; 
trail  Paso  Ancho  to  Monte  Lirio,  Allen  1497  (MO,  NY,  POM)  ;  Volcan  de 
Chiriqui,  Terry  1331  (F,  MO),  Davidson  876  (F,  MO) ;  Casita  Alta,  ^yoodson, 
Allen  and  Seibert  873  (MO,  POM)  ;  Bajo  Chorro,  Boquete  District,  Davidson 
319  (F,  MO.)  ;  Cuesta  de  Cerro  Quemado,  Pittier  3114  (US,  GH). 

This  species  is  near  F.  microphylla  in  its  coriaceous,  smallish,  usually 
toothed  leaves,  but  has  smaller  flowers  and  a  more  southern  range.  It  often  is 
much  like  F.  michoacanensis,  but  the  leaves  are  thicker,  smaller,  not  pilose ; 
however,  it  intergrades  with  it  (see  under  michoacanensis) .  Plants  that  I 
refer  to  Hemsleyana  have  largely  been  determined  as  F.  parviflora  (F. 
Encliandra  of  this  paper),  but  that  species  belongs  in  the  thinner  or  mem- 
branaceous-leaved  group. 

(95)  Fuchsia  striolata  Lundell 
(Plate  15,  fig.  86) 
Fuchsia  striolata  Lundell,  Contr.  Univ.  Mich.  Herb.,  4:20, 1940. 

Shrub,  openly  branched,  more  or  less  scandent,  the  ultimate  branchlets 
somewliat  strigulose ;  leaves  opposite,  subcoriaceous,  ovate  to  lance-ovate,  the 
blades  1.5-9.0  cm.  long,  1-5  cm.  wide,  cuneate  at  base,  acuminate  to  acutish  at 
apex,  minutely  strigulose  above,  subglabrous  and  paler  beneath,  subentire 
to  remotely  denticulate  in  upper  half,  ciliolate,  with  6-7  pairs  of  lateral  veins ; 
petioles  strigulose,  1-2.5  cm.  long;  flowers  solitary,  red;  pedicels  strigulose. 


Vol.  XXV]  MUNZ :  THE  GENUS  FUCHSIA  97 

6-12  mm.  long;  hypanthium  tubular,  strigulose,  3-4  mm.  long,  barely  1  mm. 
wide ;  sepals  red,  deltoid,  ea.  2  mm.  long,  apieulate ;  petals  red,  1  mm.  long, 
rounded ;  anthers  included,  0.6  mm.  long ;  style  4-5  mm.  long;  stigma  exserted, 
4-lobed,  less  than  1  mm.  long;  fruit  subglobose,  ca.  7  mm.  thick,  strigulose. 
T.^'pe  locality,  Volcan  de  Tacana,  Chiapas,  Mexico,  the  type  number  being 
Matilda  2765  (F,  ]\IICH,  NY)  entirely  without  flowers,  although  the  sheet 
at  New  York  has  one  young  bud.  On  the  whole,  though,  Matuda  2765  seems  to 
match  Matuda  4004,  from  Mt.  Ovando,  Chiapas  (MICH,  NY)  which  does 
have  flowers.  From  GUATEMALA :  San  Marcos,  Volcan  Tacana,  Quebrada 
Canjula,  between  Sibinal  and  Canjula,  SteyermarJc  3608  (F)  also  without 
flowers,  seems  to  be  this  species.  It  is  characterized  by  the  large  thickish  leaves 
which  are  quite  glabrous  beneath  and  with  short  pedicels  and  small  flowers. 
A  collection  from  Jalisco,  San  Sebastian,  Canon  El  Ranchito,  Mexia  1465  (F, 
MICH,  NY,  US)  has  certain  characters  of  F.  striolata  (leaf -shape,  size,  pubes- 
cence; pedicel -length ;  flower-size,  shape,  color;  fruit-size),  but  the  leaves  are 
very  much  thinner  than  in  the  Chiapas  specimens. 

(96)  Fuchsia  michoacanensis  Sesse  and  Moeino 

(Plate  15,  fig.  87) 

Fuchsia  michoacanensis  Sess6  and  MociNO,  PI.  N.  Hispan.,  ed.  1,  57, 1887-90. 

Fuchsia  iiflora  Sess6  and  MociNO,  PI.  Mex.,  ed.  2,  93,  1894  given  as  same  as  F.  michoacan- 
ensis. Based  on  specimen  no.  5209,  which  I  have  seen. 

F.  chiapensis  T.  S.  Brandg.,  Univ.  Calif.  Pub.  Bot.,  6:59, 1914;  Standley,  Contr.  U.  S.  Nat. 
Herb.,  23 :  1079,  1924.  Based  on  Purpus  6707,  which  I  have  seen. 

F.  heterotricha  Lundell,  Contr.  Mich.  Herb.,  4:19,  1940.  Based  on  Matuda  S457,  which  I 
have  seen.  It  may  have  somewhat  finer  pubescence  than  usual. 

Shrub  1-3  m.  tall,  with  slender  branches,  the  ultimate  ones  about  1  mm. 
thick,  adpressed-puberulent  to  densely  spreading-pubescent  or  pilose,  usually 
with  both  kinds  of  hairs  at  least  on  the  petioles  and  veins  of  lower  leaf- 
surfaces;  leaves  opposite,  the  blades  orbicular-ovate  or  elliptic-ovate,  mem- 
branaceous, subentire,  to  denticulate,  bieolored,  rounded  to  cuneate  at  base, 
rounded  to  acute  at  apex,  subglabrous  to  puberulent  or  sparingly  strigose 
above,  sparsely  to  densely  pilose  beneath,  7-25  (40)  mm.  long,  5-20  (25)  mm. 
wide ;  petioles  3-25  mm.  long,  usually  pilose  and  strigulose ;  flowers  axillary, 
perfect  or  pistillate;  pedicels  pilose  to  puberulent,  filiform,  5-15  (25)  mm. 
long ;  hypanthium  red,  cylindrical  in  pistillate  flowers  to  tubular-f unnelf orm 
in  perfect  ones,  often  more  or  less  pilose,  pubescent,  4-6  (10)  mm.  long,  1-3 
mm.  wide  at  apex  in  pressed  specimens,  constricted  below  the  rounded  base ; 
sepals  red,  usually  pilose,  sometimes  only  puberulent,  more  or  less  spreading, 
deltoid,  acuminate,  2-3  mm.  long,  the  apieulate  tips  connivent  in  bud ;  petals 
coral  red,  round-ovate  to  -elliptic,  erect,  0.6-2  mm.  long;  stamens  subsessile, 
almost  or  quite  included;  style  6-8  (12)  mm.  long,  the  stigma  somewhat  ex- 
serted, deeply  4-lobed,  more  or  less  red ;  berry  subglobose,  black-purple,  4-7 
mm.  thick. 


98  CALIFORNIA  ACADEMY  OF  SCIENCES  [Paoc.  4th  Ser. 

Type  locality,  not  given,  probably  southern  Mexico.  Ranging  from  southern 
Mexico  to  Costa  Rica.  Representative  material,  MEXICO :  without  definite 
locality,  ''Ftichsia  hiflora  N.  t.  2.  f.  159,"  Sesse,  MociTio,  Castillo  and  Maldon- 
ado  5209  (fragment  at  F).  Mexico,  district  of  Temascaltepec,  Pantoja,  Hinton 
2846  (NY) .  Michoacan,  Tarascon,  Pringle  11989  (F,  GH,  US) .  Chiapas,  Cerro 
del  Boqueron,  Piirpus  6707,  type  no.  chiapensis  (F,  GH,  NY,  US),  7168  (F, 
NY,  US)  ;  Volean  de  Tacana,  Matuda  2457,  type  heterotricha  (MICH).  Jal- 
isco, Real  Alto,  trail  to  El  Tajo  de  Santiago,  Mexia  1744  (F,  CAS,  MICH,  NY, 
US).  Morelos,  El  Parque,  Rose  and  Rose  11118  (NY,  US)  very  smooth. 
GUATEMALA :  San  Marcos,  Quebrada  Conjula,  Volean  Tacana,  Steyer- 
mark  36047  (F)  ;  Rio  Vega,  Volean  Tacana,  Steyermark  36233  (F)  ;  El 
Boqueron,  Standley  66305  (F,  POM) .  Quiche,  so.  of  Chiehicastenango,  Stand- 
ley  62408  (F).  Chimaltenango,  Chichivac,  Skutch  43  (US),  710  {¥)';  Cerro 
de  Tecpam,  Standley  58716  (F).  Guatemala,  Finca  La  Aurora,  Aguilar  80 
(F);  Volean  de  Pacaya,  above  Las  Calderas,  Standley  58400  (F,  POM). 
Quezaltenango,  Volean  Santa  Maria,  Steyermark  33915  (F)  ;  above  Santa 
Maria  de  Jesus,  Standley  67264  (F),  67188  (F)  ;  Volean  Zunil,  Steyermark 
34929  (F).  Jalapa,  Volean  Jumay,  Steyermark  32353  (F),  Amatitlan,  be- 
tween San  Vicente  Pacaya  and  El  Cedro,  Tonduz  428  (US).  Sacatepequez, 
Volean  de  Fuego,  J.  D.  Smith  1460  (GH,  US)  ;  Volean  de  Agua,  so.  of  Santa 
Maria  de  Jesus,  Standley  59494  (F,  POM)  ;  Finca  Carmona,  s.e.  of  Antigua, 
Standley  63805  (F,  POM).  Sacapa,  Sierra  de  las  Minas,  Steyermark  29741 
(F).  EL  SALVADOR  :  San  Vicente,  Volean  de  San  Vicente,  Standley  21549 
(NY,  GH,  US).  COSTA  RICA:  San  Jose,  Las  Nubes,  Standley  38705  (US). 

Material  which  I  refer  to  this  species  has  largely  passed  as  F.  minutiflora 
which  is  a  smaller-leaved  and  smaller-flowered  plant  of  more  northern  range. 
F.  michoacanensis  intergrades  freely  with  F.  microphylla  var.  aprica  espe- 
cially in  Guatemala,  where  much  of  the  material  is  intermediate.  In  Costa 
Rica,  it  intergrades  likewise  with  F.  Hemsleyanu,  but  in  general,  F.  michoa- 
canensis has  thin  leaves  while  the  other  two  have  more  coriaceous  ones.  Ex- 
amples of  intergradation  with  microphylla  are,  Chiapas,  Ohiesbraght  697 
(F)  ;  Barranca  Eminencia,  San  Marcos,  Guatemala,  Standley  68586  (F)  ; 
Cerro  Quemado,  Quezaltenango,  Kellerman  5939  (US);  San  Juan  de  Dios, 
Sonsonate,  El  Salvador,  Pittier  1995  (US)  ;  Volean  de  San  Salvador,  Stand- 
ley  22810  (NY,  GH,  US).  In  these  there  is  a  tendency  toward  the  pubescence 
of  michoacanensis  and  toward  the  small  thickish  leaves  of  microphylla.  With 
the  pubescence  of  F.  michoacanensis,  but  the  leaf -thickness  of  F.  Hemsleyana 
may  be  cited,  COSTA  RICA:  Cartago,  Torres  Rojas  48  (US)  ;  Volean  de 
Turrialba,  Pittier  13191  (GH,  NY,  US)  ;  Irazu,  Knntze  2343  (NY). 

(97)  Fuchsia  Encliandra  Steudel 

(Plate  15,  fig.  88) 

Fuchsia  Encliandra  Steud.,  Nomen.  Bot.,  1:649,  1840  (pul)lished  as  Enclinndra) . 
Encliandra  parvi flora  Zucc,  Abli.  Akad.  Wiss.  Muench.,  2:335,  1831-1836. 


Vol.  XXV]  MVNZ:  THE  GENUS  FUCHSIA  99 

Fuchsia  parvi flora  Zucc,  I.e.,  337;  Hemsley,  Biol.  Centr.  Amer.,  Bot.,  1:460,  1880;  Stand- 
ley,  Contr.,  U.  S.  Nat.  Herb.,  23:1080,  1924.  Not  F.  parviflora  Lindl.,  Bot.  Eeg.,  13: 
1048,  1827  (see  F.  thymi folia). 

Eierschlegeria  Lindleyi  Spach,  Ann.  Sci.  Nat.,  (2),  4:176, 1835. 

Shrub  1-4  ni.  tall,  densely  branched,  with  ultimate  branchlets  very  slender, 
puberuleut,  somewhat  strigulose  and  dark  colored  under  the  whitish  puberu- 
lence ;  leaves  mostly  opposite,  membranaceous,  lance-ovate  to  round-elliptic, 
entire  to  inconspicuously  serrulate,  minutely  stri^lose  to  almost  glabrous, 
bicoloi-ed,  obtusish  to  acute  at  both  ends,  blades  8-15  (20)  mm.  long,  4-10 
(15)  mm.  wide;  petioles  puberuleut,  4-8  (12)  mm.  long;  flowers  solitary, 
axillary,  dioecious  or  polygamous;  pedicels  filiform,  puberuleut,  5-15  mm. 
long ;  hypanthium  cylindrical,  strigulose,  red,  not  constricted  at  base  in  stam- 
inate  flowers,  6-8  (10)  mm.  long,  2-3  mm.  wide,  in  pistillate  flowers  con- 
stricted above  ovary,  then  5-7  mm.  long,  1-2  mm.  wide ;  sepals  deltoid-ovate, 
ascending  in  anthesis,  puberuleut,  reddish,  2.5-3.5  mm.  long,  apiculate,  the 
short  tips  connivent  in  the  bud ;  petals  sub  orbicular,  2-3  mm.  long,  reddish, 
more  or  less  undulate ;  stamens  included,  anthers  ca.  1.5  mm.  long ;  style  7-9 
mm.  long;  stigma  exserted,  deeply  4-lobed;  berry  dark,  7-8  mm.  thick. 

Type  locality,  Mexico.  Ranging  from  Hidalgo  to  Oaxaca,  and  character- 
ized by  its  small  entire,  thin  leaves  and  cylindrical  flowers.  As  here  under- 
stood this  species  is  not  nearly  so  inclusive  as  determinations  on  herbarium 
material  would  indicate  as  to  previous  treatment.  Material  seen,  MEXICO, 
Hidalgo,  Zimapan,  Coulter  175  (NY,  GH),  possible  mistake  on  label.  Mexico, 
base  of  Ixtaccihuatl,  Arsene  5539  (MO,  US).  Morelos,  Valle  del  Tepeite, 
Lyonnet  1358h  (US)  ;  Huitzilac,  Lyonnet  1994  (US)  ;  Valle  de  Cuernavaca, 
Lyonnei  735  (NY,  US)  ;  kilometer  61,  road,  Mexico  City  to  Cuernavaca, 
Mexia  2718  (F,  MICH,  GH,  CAS,  NY,  POM,  US)  ;  El  Parque,  Orcutt  4406 
(F,  GH).  Oaxaca,  Sierra  de  San  Felipe,  Pringle  7485  (US),  Nelson  1179 
(US)  ;  San  Felipe,  Rose  and  Hough  4605  (US) ;  Oaxaca,  Conzatti  and  Gon- 
zales 20  (MO,  US)  ;  Reyes,  Nelson  1803  (GH,  US)  ;  Yotado,  Galeotti  3068  H 
(fragment  at  F). 

(98)  Fuchsia  cylindracea  Lindley 
(Plate  15,  fig.  89) 
Fuchsia  cylindracea  Lindl.,  Bot.  Keg.,  24:66,  1838. 

Shrub,  rather  openly  branched,  up  to  5  m.  high,  the  ultimate  branchlets 
purplish,  subglabrous  to  sparingly  pubescent,  the  youngest  tips  more  densely 
but  rather  coarsely  pubescent,  rather  slender ;  leaves  membranaceous,  oppo- 
site, obovate  to  elliptic,  subentire,  cuneate  at  base,  obtuse  to  acutish  at  apex, 
strongly  bicolored,  subglabrous  to  somewhat  strigose  above,  more  or  less  pilose 
beneath,  the  blades  2-5  (6.5)  cm.  long,  1-2  (2.5)  cm.  wide ;  petioles  somewhat 
pilose  or  pubescent,  3-10  (12)  mm.  long;  plants  apparently  dioecious;  flowers 
solitary  axillary,  dioecious ;  pedicels  filiform,  pubescent,  10-25  mm.  long ; 
staminate  flowers  with  narrow  basal  portion,  then  widened  into  hypanthium 
8-10  mm.  wide,  cylindrical,  deep  red,  somewhat  pubescent ;  sepals  ovate- 
deltoid,  3-3.5  mm.  long,  red,  somewhat  ascending  in  anthesis,  short-apiculate, 


100  CALIFORNIA  ACADEMY  OF  SCIENCES  [Proc.  4th  Ser. 

the  tips  connivent  in  the  bud ;  petals  red,  suborbieular,  more  or  less  apiculate 
and  notched,  3-4  mm.  long ;  stamens  included,  anthers  1.2-1.4  mm.  long;  style 
about  1  mm.  long ;  stigma  4-lobed,  about  0.5  mm.  long ;  pistillate  flowers  con- 
stricted above  the  ovary,  then  expanded  into  a  cylindric  hj^anthium  4-5  mm. 
long  and  2-3  mm.  wide ;  sepals  and  petals  as  above,  but  somewhat  smaller ; 
stamens  present,  the  anthers  about  0.5  mm.  long,  not  functional  ? ;  stigma  ex- 
serted,  4-lobed,  ca.  1  mm.  long;  fruit  subglobose,  dark,  6-8  mm.  thick. 

Type  locality,  not  known;  seed  of  original  plants  from  Mexico.  I  have  seen 
none  of  Lindley's  herbarium  material,  but  his  plate  fits  well  the  specimens  I 
refer  to  this  species,  with  large  leaves,  long  flowers  (cylindrical,  %  inch  long 
ace.  to  Lindley),  pubescence,  etc.  The  species  is  near  F.  Enclianclra  (parvi- 
flora)  but  has  larger  leaves  and  coarser  hair ;  there  is  some  intergradation,  as 
Ghieshreght  696  (MO)  from  Chiapas  with  leaves  of  cylindracea  and  flowers  of 
Encliaoidra;  "F.  puhescens,"  SessS  and  Mocino  5215  (F).  Kepresentative  of 
cylindracea  may  be  cited,  MEXICO,  without  locality,  Coulter  174  (GH,  NY) . 
Mexico,  District  of  Temascaltepec,  Timbres,  Hint  on  3055  (US)  ;  Cajones, 
Hinton  4044  (US).  Michoacan,  Campanario  near  Morelia,  Arsene  5665  (NY, 
US),  6523  (US),  Arsene  in  1910  (F)  ;  Patzcuaro,  Pringle  4266  (F,  GH, 
]\riCH,  NY,  US) ,  3583  (F) .  Oaxaca,  18  miles  s.w.  of  Oaxaca  City,  Nelson  1391 
(US). 

(99)  Fuchsia  tetradactyla  Lindley 
(Platel5,  fig.  90) 

Fuchsia  tetradactyla  Lindl.,  Joum.  Hort.  Soc,  1:304,  1846. 

F.  Seleriana  Loesener,  Verb.  Bot.  Ver.  Brandenb.,  55:179,  1913;  Standley  and  Steyeb.- 
MARK,  Field  Mus.  Nat.  Hist.,  Bot.,  22 :  363, 1940. 

Shrub  6-15  (24)  dm.  tall,  openly  branched,  the  ultimate  branchlets  red- 
dish, 1-2  mm.  thick,  puberulent,  more  or  less  strigulose ;  leaves  opposite  or 
nearly  so,  broadly  ovate,  somewhat  rhomboid,  membranaceous,  not  crowded, 
subentire  or  remotely  denticulate,  soft  pubescent  on  both  surfaces,  bicolored, 
rounded  to  broad-cuneate  at  base,  abruptly  acute  to  obtuse  at  apex,  the  blades 
2-6  (10)  cm.  long,  1.5-5  cm.  wide,  with  about  6  pairs  of  lateral  veins;  petioles 
slender,  1.5-3  (4)  cm.  long;  flowers  solitary,  axillary,  the  plants  dioecious; 
pedicels  filiform,  strigulose  to  pubescent,  1-1.8  cm.  long;  staminate  flowers 
with  hypanthium  subcylindric,  7-10  mm.  long,  2.5-3.5  mm.  wide,  puberulent, 
rose-orchid  to  deep  red ;  sepals  red  to  orchid,  puberulent,  2.5-3  (4)  mm.  long, 
ascending,  deltoid,  apiculate,  the  tips  about  0.5  mm.  long,  somewhat  separate 
in  bud;  petals  rose-scarlet  or  paler,  suborbieular,  ca.  3  mm.  long;  stamens 
included,  anthers  2  mm.  long ;  stigma  included ;  pistillate  flowers  smaller, 
constricted  above  ovary;  hypanthium  5-6  mm.  long,  1-2  mm.  wide;  anthers 
less  than  1  mm.  long ;  style  8-9  mm.  long,  stigma  exserted,  4-lobed,  1  mm.  long ; 
fruit  purplish  or  black,  up  to  1  cm.  thick. 

Type  locality  "Guatemala,"  type  collected  by  G.  W.  Skinner,  April  4,  1846. 
I  have  not  seen  the  type,  but  am  taking  the  name  tetradactyla  for  the  plant 
recently  called  Seleriana.  The  large  leaves  (2.7  inches  long,  including  peti- 


Vol.  XXV]  MUNZ :  THE  GENUS  FUCHSIA  101 

oles),  the  calyx  (^  inch  long,  that  is  hypanthium  plus  sepals) ,  pedicels  equal- 
ing petioles,  and  origin  Guatemala,  all  these  points  mentioned  in  the  original 
description  can  indicate  only  this  species.  Material  seen,  MEXICO,  without 
definite  locality,  '^Fuchsia  tomentosa  N.,"  Sesse,  Mociiio,  et  al.  5207  (F). 
Oaxaca,  between  Suchistepec  and  Miahuatlan,  Nelson  2515  (US)  ;  between 
Plunia  and  San  Miguel  Suchistepec,  Nelson  2505  (US).  Chiapas,  Teopisca, 
Goldman  970  (US) .  GUATEMALA :  Baja  Vera  Paz,  Santa  Rosa,  von  Tuerck- 
heim  under  J.  D.  Smith  1315  (US).  Quezaltenango,  Volcan  Zunil,  Steyermark 
34741  (F)  ;  near  Quezaltenango,  Seler  2894,  type  of  Seleriana  (photos  F, 
POM,  specimen  CAS)  ;  Quezaltenango,  Skiitch  810  (F,  NY)  ;  Volcan  de 
Santa  'NLaria,8teijermark  34031  (F,  POM).  Chimaltenango,  Tecpam,  John- 
ston 761a  (F),  Skutch  489  (F,  GH,  MICH,  US).  Sacatepequez,  Santa  Lucia, 
Popenoe  689  (US) ;  Volcan  de  Acatenango,  Shannon  under  J.  D.  Smith  3656 
(US)  ;  Finca  El  Hato,  ne.  of  Antigua,  Standley  61202  (F)  ;  Santiago,  Gomez 
under  J.  D.  Smith  872  (US).  Guatemala,  Carmela,  Morales  1100  (F).  Jalapa, 
Cerro  Alcoba,  Steyermark  32512  (F,  POM).  Zacapa,  Sierra  de  las  Minas, 
Steyermark  29704  (F). 

(100)  Fuchsia  Mexiae  Munz,,  new  species 
(Plate  15,  fig.  91) 

Leafy  shrub  apparently  rather  large,  ultimate  twigs  very  slender,  some- 
what reddish,  somewhat  fine-pubescent;  leaves  opposite,  petioles  slender, 
pubescent,  7-20  (25)  mm.  long;  leaf -blades  membranaceous,  rhomboid-ovate 
to  -obovate,  bicolored,  acute  to  acuminate  at  both  ends,  remotely  sinuate- 
serrulate,  subglabrous  to  minutely  puberulent  on  both  surfaces,  especially 
on  the  veins  and  margins,  1.5^  (4.5)  cm.  long,  1-2  cm.  wide  ;  flowers  reddish, 
axillary,  dioecious ;  pedicels  filiform,  strigulose,  about  1  cm.  long;  hypanthium 
of  pistillate  flowers  tubular,  strigulose,  3-4.5  mm.  long,  barely  1  mm.  wide ; 
sepals  ovate,  1.5-2  mm.  long ;  petals  white,  rounded,  ca.  1  mm.  long ;  style  5-6 
mm.  long;  stigma  deeply  4-lobed,  well  exserted,  ca.  1  mm.  long;  staminate 
flowers  not  seen,  but  undoubtedly  larger;  fruit  dark,  5-7  mm.  thick. 

Frutex  f  oliosus ;  ramulis  tenuibus,  rubescentibus,  puberulentis ;  f  oliis  oppositis ;  petiolis 
tenuibus,  pubescentibus,  7-20  (25)  mm.  longis;  laminis  membranaceis,  rhomboideo-ovatis 
vel  -obovatis,  acutis  vel  acuminatis  in  extremis  utrisque,  remote  sinuato-serrulatis,  sub- 
glabris  vel  puberulentis  in  superficiebus  utrisque,  1.5-4  cm.  longis,  1-2  cm.  latis;  floribus 
rubescentibus,  in  axillis  solitariis;  pedicelis  filiformibus,  strigulosis,  1  cm.  longis;  hypan- 
thio  florium  pistillatorum  tubuloso,  striguloso,  3-4.5  mm.  longo,  1  mm.  lato ;  sepalis  ovatis, 
1.5-2  mm.  longis;  petalis  albis,  orbiculatis,  1  mm.  longis;  stylo  5-6  mm.  longo;  stigmate 
4-lobato,  exserto,  1  mm.  longo ;  floribus  staminatis  inconnatis ;  f  ructu  nigro,  5-7  mm.  crassi. 

Type:  from  open  dry  pine  forest,  at  1500  m.,  Canon  El  Ranchito,  San 
Sebastian,  Jalisco,  Mexico,  Jan.  12, 1927,  Ynez  Mexia  1465,  U.  S.  Herbarium 
no.  1319484 ;  isotypes  F,  GH,  CAS,  MICH,  NY.  Another  collection  is  from 
Morelos,  El  Parque,  Rose  and  Rose  11118  (NY,  US).  The  proposed  species  is 
much  like  F.  striolata,  but  has  membranaceous  leaves,  more  rhomboid  leaf- 
blades,  white  petals,  and  more  western  range. 


INDEX  TO  NAMES 
The  number  given  after  each  name  refers  to  the  species  number  used  in  this  paper. 


Brebissonia  Spach,  see  Sect.  7,  Encliandra. 

micropliylla  Spach,  see  93a. 

thymifolia  Steud.,  see  85. 
Dorvalla  eucharis  Comm.,  see  3a. 
EUobium  Lilja,  see  Sect.  2,  Eufuchsia. 

fulgens  Lilja,  see  47. 
Encliandra  Zucc,  sect.  7. 

parvi flora  Zucc,  see  97. 

thymifolia  Lilja,  see  85. 
Fuchsia  abrupta  Johnston,  no.  43. 

acynifoUa  Scheidw.  in  Otto  and  Dietr., 
Allg.  Gartenz.,  15:226,  1847.  I  have 
seen  no  material  so  labeled;  no  type 
number  was  designated.  It  was  said  to 
have  come  from  Mexico  and  the  char- 
acters mentioned  (small  ovate  leaves 
dentate  toward  the  tips;  calyx  cylin- 
drical, 6  lines  long;  leaves  with  some 
puberulence  above,  glabrous  beneath ; 
pedicels  short)  suggest  F.  microphylla 
H.B.K.,  no.  93. 

affinis  Cambess.,  see  6c. 

alpestris  Gardn.,  see  6d. 

alternans  Moc.  and  Sesse,  see  85. 

amoena  Hort.,  mentioned  in  DC.  Prodr., 
3:37,  1828  a  ssynonym  of  F.  arbores- 
cens. 

ampliata  Bouth.,  see  21. 

Andrei  Johnston,  no.  54. 

apetala  Kuiz  and  Pavon,  no.  78. 

apiculata  Johnston,  see  16. 

aprica  Lundell,  see  93b. 

araucana  F.  Pliil.,  see  3a. 

arborea  Sess6  and  Moc,  see  84a. 

arborescens  Sims,  no.  84. 
Fuchsia  arborescens  var.  megalantha  Donn. 
Smith,  see  84a 
forma  parva  Munz,  no.  84c. 
forma  tenuis  Munz,  no.  84b. 
forma  typica  Munz,  no.  84a. 

asperifnlia  Krause,  no.  59. 

Aspiazui  Macbr.,  no.  33. 

Asplundii  Macbr.,  no.  26. 

atrorubra  Johnston,  see  64. 

austromontana  Johnston,  no.  9. 

ayavacensis  H.B.K.,  no.  21. 

bacillaris  Lindl.,  no.  91. 

biflora  Sesse  and  Moc,  see  96. 

boliviano  Britton,  see  52. 


boliviana  Carr.,  no.  46. 

f .  puberulenta  Munz,  no.  46b. 

var.  luzurians  Johnston,  no.  46c. 

var.  typica  Munz,  no.  46a. 
Bracelinae  Munz,  no.  2. 
Brittoni  Johnston,  see  52. 
Campos-Portoi  Pilger  and  Schulze,  no.  1. 
canesccns  Benth.,  no.  14. 
caracasana  Field,  and  Gardn.,  see  40. 
cestroides  Schulze-Menz,  no.  83. 
chiapensis  Brandg.,  see  96. 
chloroloba  Johnston,  see  73a. 
chonotica  Phil.,  see  3a. 
coccinea  Curtis,  see  3b. 
coccinea  Soland.  in  Ait.,  no.  4. 

var.  chronotica  Eeiche,  see  3a. 
Colensoi  Hook,  f .,  no.  68. 
colimae  Munz,  no.  88. 
confertifolia  Field,  and  Gardn.,  no.  32. 
conica  Lindl.,  see  3a. 
corallina  Hort.  ex  Lynch,  Gard.  Chron., 

2:604,  1883;  said  by  Lynch  to  be  syn-j 

onym  of  F.  exoniensis. 
Fuclisia  cordi folia  Benth.,  no.  8. 
cordifolia  /3  Hook.,  see  7. 
corollata  Benth.,  see  14. 
corymbi flora  R.  and  P.,  no.  42. 
corymbosa  Pritz.,  see  42. 
Cuatrecasasii  Munz,  no.  44. 
curviflora  Benth.,  see  24a. 
cuspidata  Fawcett  and  Eendle,  see  46c. 
cylindracea  Lindl.,  no.  98. 
cyrtandroides  Moore,  no.  67. 
decidua  Standley,  no.  72. 
decussata  R.  Grah.,  see  3b. 
decnssata  R.  and  P.,  no.  48. 
denticulata  R.  and  P.,  no.  10. 
dependens  Hook.,  see  42. 
discolor  Lindl.,  see  3a. 
dolichantha  Krause,  see  32. 
Bominiana  Hort.  ex.  Fl.  des  Serres,  ser.  1, 

95,  pi.  1004,  1854-5 ;  said  to  be  a  horti- 
cultural hybrid. 
elegans  Paxt.  ex.  Steud.;  see  globosa  var. 

elegans. 
elegans  Salisb.,  see  4. 
Encliandra  (spelled  Encliandra)  Steud., 

no.  97. 
excorticata  L.  f.,  no.  66. 


[102] 


Vol.  XXV] 


MUNZ:  THE  GENUS  FUCHSIA 


103 


exoniensis  Paxt.,  Mag.  Bot.,  10 :  153, 1843 ; 
hybrid  between  F.  cordifolia  and  F. 
globosa. 

filipes  Kusby,  see  52. 

Fischeri  Macbr.,  no.  36. 

fontinalis  Macbr.,  see  48. 

fulgens  Moc.  and  Sesse,  no.  47. 

furfuracea  Johnston,  no.  38. 

fusca  Krause,  see  48. 

Garleppiana  Kuntze  and  Wittm.,  no.  81. 
Fuchsia  Gelirigeri  Munz,  no.  30. 

glaierrima  Johnston,  no.  61. 

Glazioviana  Taub.,  see  4. 

gloiosa  Lindl.,  Bot.  Beg.,  18: pi.  1556, 
1833 ;  apparently  a  plant  of  horticul- 
tural origin.  I  have  seen  no  wild  speci- 
mens like  it;  it  apparently  has  arisen 
from  F.  magellanica. 
var.  elegans  Paxton,  Mag.  Bot.,  4:75, 
1838 ;  apparently  horticultural. 

gracilis  Lindl.,  see  3b. 
var.  multiflora  Lindl.,  see  3a.  var.  ten- 
ella  Lindl.,  see  3a. 

gracilis  Moc  and  Sesse,  see  93a. 

grandiflora  Euiz  ex  Dahlgren,  see  10. 

hamellioides  Moc.  and  Sesse,  see  84a. 

Hartwegii  Benth.,  no.  57. 

Hemsleyana  Woodson  and  Seibert,  no,  94. 

heterotricha  Lundell,  see  96. 

hirsuta  Hemsl.,  no.  79. 

hirtella  H.B.K.,  no.,  40. 

Hitchcoclcii  Johnston,  see  21. 

liortensis  Bergmans,  Vaste  PI.  Eotshee- 
sters,  227,  1924 ;  horticultural. 

hyhrida  Hort.,  no.  5. 

hypoleuca  Johnston,  no.  49. 

inflata  Schulze-Menz,  see  7.3b. 

insignis  Hemsl,,  see  78. 

integrifolia  Cambess.,  see  6a. 

intermedia  Hemsl.,  see  7. 

involucrata  Sw.  Prodr.  Veg.  Ind.  Occ,  62, 
1788,  is  Schradera  Cephalotes  of  Eubi- 
aceae. 

involucrata  K.  and  P.,  Field  Mus.  Nat. 
Hist.,  Bot.  21 :  212, 1940 ;  nomen.  nudum 

Jahnii  Munz,  no.  29, 

juntasensis  Kuntze,  no.  74. 

Killipii  Johnston,  no.  45. 

Kirhii  Hook,  f.,  no.  71. 

Lehmannii  Munz,  no.  55. 

Lenneana  Warcz.  in  Otto  and  Dietr.,  Allg. 
Gartenz.,  20:293,  1852;  novien  sub- 
nudum. 


Fuchsia  leptopoda  Krause,  no.  11. 
Liebmanni  Levi.,  see  84a. 
Llewelynii  Macbr.,  no.  27. 
longiflora  Benth.,  see  20a. 
Lowei  Hort.  ex  L.  H.  Bailey,  Standard 

Cycl.  Hort.,  1301,  1915,  where  given  in 

synonymy   under   F.   magellanica  var. 

discolor, 
loxensis  H.B.K.,  no.  16. 
lycioides  Andr.,  no.  65. 
macrantha  Hook.,  no.  82. 
macropetala  Presl.,  see  42. 
macrophylla  Johnston,  no.  62. 
macrostema  E.  and  P.,  see  3b. 

var.  conica  Sweet,  see  3a. 

var.  gracilis  Sweet,  see  3b. 

var.  recurvata  Hook.,  see  3b. 

var.  tenella  DC,  see  3a. 
macrostigma  Benth.,  no.  20. 

var.  longiflora  Munz,  no.  20a. 

var.  typica  Munz.,  no.  20b. 
magdalenae  Munz,  no.  13. 
magellanica  Lam.,  no.  3. 

var.  conica  Bailey,  see  3a. 

var.  discolor  Bailey,  see  3a. 

var.  globosa  Bailey,  see  F.  globosa. 

var.  gracilis  Bailey,  see  3b. 

var.  inacrostema  Munz,  no.  3b. 

var.  Molinae  Espinosa,  no.  3c. 

var.  Biccartoni  Bailey,  see  F.  Riccar- 
toni. 

var.  typica  Munz,  no.  3a. 
Mathewsii  Macbr.,  no.  35. 
Mattoana  Krause,  see  74. 
membranacea  Hemsl.,  no.  75. 
Mexiae  Munz,  no.  100. 
michoacanensis  Sesse  and  Moc,  no.  96. 
microphylla  H.B.K.,  no.  93. 

var.  aprica  Munz,  no.  93b. 

var.  typica  Munz,  no.  93a. 
Fuchsia  miniata  Planch  and  Linden,  see  40. 
minimiflora  Hemsl.,  no.  87. 
minutiflora  Hemsl.,  no.  92. 

var.  hidalgensis  Munz,  no.  92a. 

var.  typica  Munz,  no.  92b. 
mitifolia  (mutifolia  in  index)  E.  and  P., 

in  Field  Mus.  Pub,,  Bot.,  21:80,  1940; 

nomen  nudum, 
mixta  Hemsl.,  see  93a. 
mollis  Krause,  see  6d. 
montana  Cambess.  ,see  4. 
multiflora  Lodd.,  see  3a. 


104 


CALIFORNIA  ACADEMY  OF  SCIENCES 


[Proc.  4th  Ser. 


multiflora  Murr.,  Syst.  ed.  13,  299,  1774; 
nomen  nudum. 

Munzii,  Macbr.,  no.  39. 

myrtifolia  Hort.  ex  Koehne,  Deutsche 
Dendrol.,  429,  1893 ;  said  to  be  near  F. 
discolor. 

nigricans  Linden,  see  64. 

Notarisii  Lehm.,  Hamb.  Gartenz,,  8:459, 
1852;  I  have  not  seen  authentic  mate- 
rial; description  suggests  a  member  of 
section  Encliandra. 

Osgoodii  Macbr.,  no.  53. 

ovalis  E.  and  P.,  no,  58. 

var.  dberrans  Macbr.,  see  53. 

ovata  Moc.  and  Sesse,  see  85. 

pallescens  Diels,  no.  17. 

paniculata  Lindl.,  see  84c. 

parviflora  Lindl.,  see  85. 

parviflora  Zucc,  see  97. 

pendula  Salisb.,  see  4. 

pendulae flora  Hort.,  Gard.  Chron.,  1:521, 
1877;  hybrid. 

perirevis  Johnston,  see  51. 

persicandens  Cockayne  and  Allan,  no.  69. 

peruviana  Carr.  in  Johnston,  Contr.  Gray 
Herb.,  75:39,   1925,  apparently  a  slip 
for  F.  boliviana. 
Fuchsia  petiolaris  H.B.K.,  no.  24. 
var.  bolivarensis  Munz,  no.  24b. 
var.  typica  Munz,  no.  24a. 

pilosa  Field,  and  Gardn.,  no.  60. 

platypetala  Johnston,  no.  19. 

polyantJia  Killip,  no.  41. 

polyanthella  Johnston,  see  58. 

Pringlei  Eobinson  and  Seaton,  no.  86. 

Pringsheimii  Urb.,  no.  22. 

procumbens  Cunn.,  no.  70. 

prostrata  Baill.,  see  70. 

pubescens  Cambess.,  see  4. 

pubescens  R.  and  P.,  in  Field  Mus.  Nat. 
Hist.,  Bot.,  21:212, 1940,  nomen  nudum. 

pulchella  Woodson  and  Seibert,  see  94. 

pumila  Hort.  ex.  Muenissier,  Rev.  Hort., 
n.s.  20:173, 1926;  horticultural. 

pumila  Hort.  ex  Vilmorin,  Blumeng.,  ed. 
3,  Sieb.  and  Voss,  1 :  332, 1894,  in  synon- 
ymy under  F.  gracilis  f.  pumila  Voss. 

punicea  R.  and  P.,  Field  Mus.  Nat.  Hist., 
Bot.,  21:212,  1940,  nomen  nudum. 

putumayensis  Munz,  no.  56. 

pyrifolia  Presl,  see  6b. 

quinduensis  H.B.K.,  see  24a. 


racemosa  Lam.,  Encye.  2:565,  1788;  pi. 
282,  fig.  1,  1793.  Given  with  F.  triphylla 
as  synonym,  but  as  coming  from  Carta- 
gena, Colombia.  It  may  be  an  earlier 
name  for  F.  petiolaris,  but  I  have  not 
had  access  to  a  specimen  since  I  dis- 
covered that  fact. 

racemosa  Sesse  and  Moc,  PI.  N.  Hispan., 
ed.  1,  58,  1887-'90.  Description  given 
suggests  F.  fulgens,  although  it  was 
referred  to  F.  arborescens  by  G.  Don, 
Gen.  Syst.,  2:677,  1832.  Leaf  at  Field 
(ex  herbario  Musei  Parisiensis)  labeled 
"F.  racemosa  Lam."  can  well  be  F.  ful- 
gens in  size,  veins,  pubescence. 

radicans  Miers,  see  6b. 
Fuchsia  recurvata  Niven,  see  3b. 

regia  (Veil.)  Munz,  no.  6. 
var.  affinis  Munz,  no.  6c. 
var.  alpestris  Munz,  no.  6d. 
var.  radicans  Munz,  no.  6b. 
var.  typica  Munz,  no.  6a. 

Biccartonensis  Bergmans,  Vaste  PI.  Rot- 
sheesters,  227,  1924;  see  F.  Riccartonii. 

Biccartoni  Hort.  ex  Tillery,  in  Flor.  and 
Pom.,  217,  1871;  horticultural  name 
with  inadequate  description. 

Biccartoniana  Hort.  ex  Koehne,  Deutsche 
Dendrol.,  429, 1893 ;  horticultural  name. 

rivularis  Macbr.,  no.  15. 

robusta  Tengbergen,  Tuinb.  FL,  2:345, 
1855;  hybrid. 

rosea  R.  and  P.,  see  65. 
var.  spinosa  Eeiche,  see  65. 

salicifolia  Hemsl.,  no.  76. 

sanctae-rosae  Kuntze,  no.  52. 

scabriuscula  Benth.,  no.  50. 

scandens  Krause,  see  48. 

Scherffiana  Andre,  Rev.  Hort.,  1888:233 
and  268,  from  Ecuador ;  would  seem 
nearest  to  F.  Townsendii  Johnston,  over 
which  it  has  priority,  but  the  description 
sounds  as  if  it  had  narrower  petals.  I 
have  not  seen  type  material. 

Seleriana  Loes.,  see  99. 

semperfiorens  Rozain,  Wien.  Illustr.  Gar- 
tenz., 1888:244;  hybrid. 

serratifolia  R.  and  P.  ,see  10. 

serratifolia  Hook.,  see  9. 

sessilifolia  Benth.,  no.  63. 

simplicicaulis  R.  and  P.,  no.  31.  '^ 

siphonantha  Krause,  see  11. 


Vol.  XXV] 


MUNZ:  TEE  GENUS  FUCHSIA 


105 


STcutchiana  Munz,  no.  90. 
Smithii  Munz,  no.  25. 
Fuchsia  speciosa  Hort.,  see  5. 
spectaiilis  Hook.,  see  20a. 

var.  pubens  Johnston,  see  20b. 
spinosa  Presl,  see  65. 
splendens  Zucc,  no.  7. 
Standishii  Paxton,  Bot.  Mag.,  11 :  33, 1844 ; 

garden  hybrid. 
Steinbachii  Johnston,  see  74. 
Storlcii  Munz,  no.  37. 
striolata  Lundell,  no.  95. 
sylvatica  Benth.,  no.  64. 
syringae flora  Carr.,  see  84a. 
tacanensis  Lundell,  no.  89, 
tacsoniiflora  Krause,  see  10. 
tenella  Hort.,  see  3a. 
tetradactyla  Lindl.,  no.  99. 
Thompsoni   Hort.    ex   Koehne,   Deutsche 

Dendrol.,  430,  1893 ;  nomen  nudum.  No 

description,  3  varieties  figured. 
thymifoUa  H.B.K.,  no.  85. 
tincta  Johnston,  no.  34. 
Townsendii  Johnston,  no.  18. 
tripliylla  L.,  no.  23. 
tuierosa  Krause,  no.  73. 

var.  inflata  Munz,  no.  73b. 

var.  typica  Munz,  no.  73a. 
tunariensis  Kuntze,  no.  77. 
umhrosa  Benth.,  see  16. 
unduavensis  Munz,  no.  80. 
uniflora  Sesse  and  Moc,  PI.  N.  Hispan., 

ed.  1,  58,  1887-'90.  Unknown  to  me. 
velutina  Johnston,  see  42. 
venusta  H.B.K.,  no.  28. 

var.  huilensis  Munz,  no.  28b. 

var.  typica  Munz,  no.  28a. 


Fuchsia  verrucosa  Hartw.,  no.  51. 

verticillata  E.  and  P.,  in  Field  Mus.  Nat. 

Hist.,  Bot.,  21:46, 1940;  nomen  nudum. 

violacea  E.  and  P.,  in  Field  Mus.  Nat. 

Hist.,  Bot.,  21 :  128, 1940 ;  nomen  nudum. 

virgata  Sweet,  Hort.  Brit.,  ed.  1,  485, 1827 ; 

nomen  nudum, 
vulcanica  Andre,  see  14. 
Weberbaueri  Krause,  see  52. 
WoytTcowsMi  Maebr.,  no.  12. 
Kierschlegeria  Spach,  section  no.  3. 
Lindleyi  Spach,  see  97. 
lycioides  Spach,  see  ^^5. 
Lopesia  thymifoUa  Willd.,  see  85. 
Lyciopsis  Spach,  see  Encliandra,  sect.  7. 

thymifoUa  Spach,  see  85. 
Myrinia  Lilja,  see  EncUandra,  sect.  7. 

microphylla  Lilja,  see  93a. 
Nahusia  Schneev.,  see  Quelusia,  sect.  1. 

coccinea  Schneev.,  see  4. 
Quelusia  Vandelli,  section  no.  1. 

regia  Vand.,  see  6. 
Schufla  Spach,  section  no.  6. 

arborescens  Spach,  see  84a. 
SJcinnera  Forst.,  section  no.  4. 

antartica  Forst.   said  to  be  BlacTcweUia 

antartica  Gaertn. 
coccinea  Moench,  see  4. 
exeorticata  Forst,  see  66. 
Spachia    Lilja,    in    Traedgardtidning,    62, 
1840;    ex   Linnaea,   15:262,   1841.   No 
species  named;  synonym  of  Fuchsia. 
Thilcum  (or  Thileum'?)  Molina,  see  Nahusia, 
sect.  1. 
tinctorium  Molina,  see  3b. 
Tilco  Adans.,  Fam.  2:498,  1763;  no  species 
named;    synonym   of  Fuchsia,  ace.  to 
Kew  Index. 


EXPLANATION  OF  PLATES 

Measurements  given  for  flower  are  from  base  of  ovary 
to  tip  of  stigma,  for  leaf  include  petiole  and  blade. 

PLATE   1 
Fig.  1.  F.  Campos-Fortoi,  drawn  from  Brade  15676 ;  flower  32  mm. ;  leaf  27  mm. 
Fig.  2.  7*'.  Bracelinae,  drawn  from  Mexia  4013  ;  flower  42  mm. ;  leaf  45  mm. 
Fig.  3.  F.  magellanica,  from  "Werdermann  61 ;  flower  56  mm. ;  leaf  53  mm. 
Fig.  4.  F.  coccinea,  from  Hoehne  6278;  flower  41  mm.;  leaf  41  mm. 
Fig.  5.  F.  regia,  from  Eegnell  794 ;  flower  62  mm. ;  leaf  66  mm. 
Fig.  6.  F.  splendens,  from  Standley  61808  ;  flower  60  mm. ;  leaf  80  mm. 
Fig.  7.  F.  cordi folia,  from  Skutcli  867 ;  flower  83  mm. ;  leaf  120  mm. 
Fig.  8.  F.  auiitromontana,  from  Pennell  14110 ;  flower  56  mm. ;  leaf  104  mm. 


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Cmunz]  plate  I 


[107] 


PLATE  2 

Fig.    9.  F.  denticulata,  from  Weberbauer  7587 ;  flower  72  mm. ;  leaf  103  mm. 
Fig.  10.  F.  leptopoda,  from  Weberbauer  6934;  flower  80  mm.;  leaf  107  mm. 
Fig.  11.  F.  W oyiTcoivskii,  from  Woytkowski  coll. ;  flower  58  mm. ;  leaf  73  mm. 
Fig.  12.  F.  magdalenae,  from  Seifriz  392;  flower  95  mm.;  leaf  65  mm. 
Fig.  13.  F.  canescens,  from  Penland  623 ;  flower  80  mm. ;  leaf  55  mm. 
Fig.  14.  F.  loxensis,  from  Penland  959 ;  flower  30  mm. ;  leaf  68  mm. 


[108] 


PROC.  CALIF.  ACAD.  SCI..  4TH  SERIES.  VOL.  XXV.  NO.    1 


[MUNZ]  PLATE  2 


[109] 


PLATE  3 

Fig.  15.  F.  Townsendii,  from  Townsend  A93  ;  flower  75  mm.;  leaf  128  mm. 

Fig.  16.  F.  platypetala,  from  West  3705 ;  flower  78  mm. ;  leaf  91  mm. 

Fig.  17.  F.  macrostigma,  from  Eose  22479 ;  flower  83  mm. ;  leaf  145  mm. 

Fig.  18.  F.  ayavacensis,  do  not  have  record  of  specimen ;  flower  56  mm. ;  leaf 
95  mm. 


[110] 


PROC.  CALIF.  ACAD.  SCI..  4TH  SERIES.  VOL.  XXV.  NO.   1 


[MUNZ]  PLATE  3 


[111] 


PLATE  4 

Fig.  19.  F.  Prm^rs/ietrnw,  from  von  Tuerckheim  3151 ;  flower  65  mm. ;  leaf  25  mm. 

Fig.  20.  F.  triphylla,  from  Leonard  3681;  flower  50  mm.;  leaf  70  mm. 

Fig.  21.  F.  petiolaris,  from  Pennell  2379;  flower  69  mm.;  leaf  57  mm. 

Fig.  22.  F.  petiolaris  var.  iolivarensis,  from  Pennell  4324;  flower  80  mm.;  leaf 
76  mm. 

Fig.  23.  F.  Smithii,  from  Killip  and  Smith  17300 ;  flower  75  mm. ;  leaf  57  mm. 

Fig.  24.  F.  Asplundii,  from  Weberbauer  6054 ;  flower  63  mm. ;  leaf  75  mm. 


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PROC.  CALIF.  ACAD.  SCI..  4TH  SERIES.  VOL.  XXV.  NO.    1 


[MUNZ]  PLATE  4 


[  113  ] 


PLATE  5 

Fig.  25.  F.  LZcwdi/nM,  from  Llewelyn  Williams  7594;  flower  68mm.;  leaf  87mm. 

Fig.  26.  F.  venusta,  no  record  of  specimen  from  which  drawn;  flower  62  mm.; 
leaf  125  mm. 

Fig.  27.  F.  Jahnii,  from  Jahn  506 ;  flower  60  mm. ;  leaf  71  mm. 

Fig.  28.  F.  Gehrigeri,  from  Gehriger  322;  flower  62  mm.;  leaf  80  mm. 

Fig.  29.  F.  simplicicaulis,  from  Macbride  4014;  flower  68  mm.;  leaf  141  mm. 


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[MUNZ]  PLATE  5 


[1]5] 


I 


PLATE  6 

Fig.  30.  F.  con ferti folia,  from  Weberbauer  7153  ;  flower  74  mm. ;  leaf  17  mm. 
Fig.  31.  F.  Aspiazui,  from  Weberbauer  7042;  flower  78  mm.;  leaf  133  mm. 
Fig.  32.  F.  tincta,  from  West  7092  ;  flower  44  mm. ;  leaf  134  mm. 
Fig.  33.  F.  StorTcii,  from  Stork  10073 ;  flower  41  mm. ;  leaf  75  mm. 
Fig.  34.  F.  furfuracea,  from  Bang  674 ;  flower  68  mm. ;  leaf  100  mm. 


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PROC.  CALIF.  ACAD.  SCI..  4TH  SERIES.  VOL.  XXV.  NO.   1 


CMUNZ]«PLATE  6 


[117] 


PLATE  7 

Fig.  35.  F.  Munzii,  from  Weberbauer  6648;  flower  52  mm.;  leaf  115  mm. 
Fig.  36.  F.  hirtella,  from  Cuatrecasas  6657  ;  flower  64  mm.;  leaf  125  mm. 
Fig.  37.  F.  polyantha,  from  Mexia  7571 ;  flower  55  mm. ;  leaf  69  mm. 
Fig.  38.  F.  corymbiflora,  from  Mexia  7765  ;  flower  68  mm. ;  leaf  148  mm. 
Fig.  39.  F.  dbrupta,  from  Macbride  4541 ;  flower  58  mm. ;  leaf  116  mm. 
Fig.  40.  F.  Cuatrecasasii,  from  Cuatrecasas  8738;  flower  47  mm.;  leaf  84  mm. 


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PROC.  CALIF.  ACAD.  SCI..  4TH  SERIES.  VOL.  XXV.  NO.    1 


[MUNZ]  PLATE  7 


[119] 


PLATE  8 

Fig.  41.  F.  Killipii,  from  Killip  16695;  flower  62  mm.;  leaf  102  mm. 

Fig.  42.  F.  boUviana,  from  Balls  6715 ;  flower  84  mm. ;  leaf  140  mm. 

Fig.  43.  F.  fulgens,  from  Pringle  4123 ;  flower  150  mm. ;  leaf  180  mm. 

Fig.  44.  F.  decussata,  from  Macbride  and  Featherstone  2124;  flower  28  mm.; 
leaf  65  mm. 

Fig.  45.  F.  hypoleuca,  from  Hitchcock  21440 ;  flower  42  mm. ;  leaf  70  mm. 

Fig.  46.  F.  scabriuscula,  from  unknown  source;  flower  45  mm.;  leaf  106  mm. 

Fig.  47.  F.  verrucosa,  from  Cuatrecasas  8465 ;  flower  24  mm. ;  leaf  65  mm. 


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PROC.  CALIF.  ACAD.  SCI..  4TH  SERIES.  VOL.  XXV,  NO.    1 


[MUNZ]  PLATE  8 


[121] 


i 


PLATE  9 
rig.  48.  F.  sanctae-rosae,  from  West  6414 ;  flower  40  mm. ;  leaf  96  mm. 
Fig.  49.  F.  Osgoodii,  from  Osgood  and  Anderson  47 ;  flower  38  mm. ;  leaf  99  mm. 
Fig.  50.  F.  Andrei,  from  Andre  K  820 ;  flower  45  mm. ;  leaf  122  mm. 
Fig.  51.  F.  Lehmannii,  from  Lehmann  5498 ;  flower  35  mm. ;  leaf  140  mm. 
Fig.  52.  F.  putumayensis,  from  Garcia  4639;  flower  28  mm. ;  leaf  91  mm. 
Y\g.5i.F.Bartwegu,  from  Killip  6829;   flower  37  mm.;  leaf  130  mm. 


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PROC.  CALIF.  ACAD.  SCI..  4TH  SERIES,  VOL.  XXV.  NO.   1 


[MUNZ]  PLATE  9 


[123] 


PLATE  10 

Fig.  54.  F.  ovalis,  from  Webcrbauer  6721 ;  flower  24  mm. ;  leaf  189  mm. 
Fig.  55.  F.  glaberrima,  from  Hitchcock  21750;  flower  42  mm.;  leaf  161  mm. 
Fig.  56.  F.  viacrophylla,  from  Maebride  5616;  flower  31  mm.;  leaf  225  mm. 


[124] 


PROC.  CALIF.  ACAD.  SCI..  4TH  SERIES.  VOL,  XXV.  NO.    I 


[MUNZ]  PLATE  ID 


[125] 


PLATE  11 

Fig.  57.  F.  sessilifolia,  from  Mexia  7597;  flower  30  mm.;  leaf  138  mm. 

Fig.  58.  F.  sylvatica,  from  Dawe  769;  flower  42  mm. ;  leaf  131  mm. 

Fig.  59.  F.  lycioides,  from  Garaventa  1641 ;  flower  14  mm. ;  leaf  26  mm. 

Fig.  60.  F.  excortieata,  from  Petrie  4110  ;  flower  40  mm. ;  leaf  95  mm. 

Fig.  61.  F.  ciirtandroidea, from  MacDaniels  in  1927 ;  flower  34 mm. ;  leaf  130 mm. 


[126] 


PROC.  CALIF.  ACAD.  SCI.,  4TH  SERrES.  VOL.  XXV.  NO.   1 


[MUNZ]  PLATE  1  1 


[127] 


PLATE  12 

Fig,  62.  F.  Colensoi,  from  Petrie  4109 ;  flower  28  mm. ;  leaf  48  mm. 
Fig.  63.  F.  perscandens,  from  Allan  in  1924;  flower  32  mm.;  leaf  38  mm. 
Fig.  64.  F.  procumiens,  from  Meebold  25160 ;  flower  17  mm.;  leaf  40  mm. 
Fig.  65.  F.  decidua,  from  Mexia  1601 ;  flower  45  mm. 
Fig.  66.  F.  tuberosa,  from  Pennell  13973 ;  flower  82  mm. ;  leaf  135  mm. 
Fig.  67.  F.  juntasensis,  from  Steinbach  5038 ;  flower  82  mm. ;  leaf  91  mm. 


[128] 


PROC.  CALIF.  ACAD.  SCI..  4TH  SERIES.  VOL.  XXV.  NO.    1 


[MUNZ]  PLATE  12 


[129] 


PLATE  13 

Fig.  68.  F.  tiinariensis,  from  Cardenas  3369;  flower  91  mm.;  leaf  76  mm. 

Fig.  69.  F.  apetala,  from  Hitchcock  21701;  flower  72  mm.;  leaf  108  mm.  and  ■ 
from  Eose  22230. 

Fig.  70.  F.  hirsuta,  from  Weberbauer  6975;  flower  67  mm.;  leaf  131  mm. 

Fig.  71.  F.  unduavensis,  from  Buehtien  2925;  flower  80  mm.;  leaf  88  mm. 


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PROC.  CALIF.  ACAD.  SCI..  4TH  SERIES.  VOL.  XXV.  NO.   1 


[MUNZ]  PLATE  13 


[131] 


PLATE  14 

Fig.  72.  F.  Garleppiana,  from  Cardenas  688;  flower  152  mm,;  leaf  153  mm. 
Fig.  73.  F.  macrantha,  from  Cook  and  Gilbert  1312;  flower  136  mm.;  leaf  87  mm. 
Fig.  74.  F.  cestroides,  from  Weberbauer  6423 ;  flower  35  mm. ;  leaf  111  mm. 
Fig.  75.  F.  arhorescensva.T.typica,  from  Skutch59;  flower  15mm.;  leaf  125mm. 


[132] 


=ROC.  CALIF.  ACAD.  SCI..  4TH  SERIES.  VOL.  XXV.  NO.   I 


[MUNZ]  PLATE  14 


[133] 


PLATE  15 

Fig.  76.  F.  thymifoUa,  from  Pringle  9794;  flower  11  mm.;  leaf  21  mm. 

Fig.  77.  F.  Pringlei,  from  Pringle  4140 ;  flower  7  mm. ;  leaf  20  mm. 

Fig.  78.  F.  colimae,  from  Jones  224;  flower  6  mm.;  leaf  20  mm. 

Fig.  79.  F.  minimiflora,  from  Rose  and  Hay  5521 ;  flower  5  mm. ;  leaf  14  mm. 

Fig.  80.  F.  tacanensis,  from  Matuda  1686;  flower  13  mm. ;  leaf  80  mm. 

Fig.  81.  F.  STcutchiana,  from  Skutch  812;  flower  7  mm.;  leaf  75  mm. 

Fig.  82.  F.  bacillaris,  from  Hort.  Bot.  Petrop.,  1829;  flower  16  mm. ;  leaf  30  mm. 

Fig.  83.  F.  minutiflora  var.  hidalgensis,  from  Pringle  8807 ;  flower  11  mm. ;  leaf 
14  mm. 

Fig.  84.  F.  microphylla,  from  Pringle  4314;  perfect  flower  18  mm.,  leaf  17  mm. ; 
pistillate  flower  10  mm.,  from  Hartweg  285. 

Fig.  85.  F.  Hemsleyana,  from  Davidson  876 ;  flower  11  mm. ;  leaf  15  mm. 

Fig.  86.  F.  striolata,  from  Matuda  4004 ;  flower  7  mm. ;  leaf  95  mm. 

Fig.  87.  F.  michoacanensis,  from  Standley  67188;  perfect  flower  11  mm.,  leaf 
40  mm. ;  pistillate  flower  from  Standley  58934,  8  mm. 

Fig.  88.  F.  Encliandra,  from  Mexia  2718 ;  staminate  flower  13  mm.,  leaf  25mm. ; 
pistillate  flower  from  Nelson  1803,  12  mm.  long. 

Fig.  89.  F.  cylindracea,  from  Arsene  6523  ;  flower  15  mm. ;  leaf  65  mm. 

Fig.  90.  F.  tetradactyla,  from  Popenoe  689 ;  staminate  flower  13  mm.,  leaf  40 
mm.;  pistillate  flower  from  Skutch  810,  15  mm. 

Fig.  91.  F.  Mexiae,  from  Mexia  1465 ;  flower  7  mm. ;  leaf  75  mm. 


[  134  1 


PROC.  CALIF.  ACAD.  SCI..  4TH  SERIES,  VOL.  XXV.  NO.   1 


[MUNZ]  PLATE  15 


[  135  ] 


PLATE  16 
(Photographs  by  Dr.  G.  Dallas  Hanua) 

Fig.  92.  Photograph  of  title-page  of  Plumier's  "Nova  Plantarum  americanarum 
Genera." 

Fig.  93.  Photograph  of  Plate  14  of  same  with  first  illustrations  of  the  Genus 
Fuchsia. 

Figs.  94  and  95.  Photographs  of  pages  14  and  15  of  same  containing  the  descrip- 
tion of  Fuchsia  by  Plumier. 


[136] 


PROC.  CALIF.  ACAD.  SCI..  4TH  SERIES.  VOL.  XXV.  NO.    1 

92 
NOVA 

PLANTARUM 

A  M  E  R  I  C  A  N  A  R  U  M 

GENERA. 

Authore  P.  Carolo  PluMIER  Oidinis 
Minimorum  in  Fiovincia  Francix,  &C  apud  Infulas 
AiTiericanasBotanico  Regio. 


[MUNZ]  PLATE  16 

93 


PARISIIS, 

Apud  JOANNEM    BOUDOT,  Regis  Si  Rcgi*  Sclcnnirunv 
Aadema-  Typographum >  via  Jacobxa.ad  SoIldi  A.iicjm. 

M.    D  C  C  1 1 1."""" 

CVM    r  RIV  1  LE  G  I  O    REGIS. 


la*.  11. 


ViUi'ir 
Mm. 


Vt.  M. 


94 

14  NovxPlant. 

mus  Viofioridis  ^  fatlidjfmtts  inda^aior  fUntdritfH  nort  dnees 
co^nttarvm  fuit.  yid  qHoa  pater  exemplo  Inflammavu  ,  tjm  eum 
ab  inndcun^thulis  inter  iffu  hcrbas  0"  fores  zoiutt  educart.  Rome 
moritur  I544.  £tat.  19.  Edidit  HiJ^oridftirpiuni Lib.  ^.  Extant 
Argentine  ijtfj.  lal. 

T  R  A  G    I   A. 

TRagia  eft  plantar  genus  flore  A  monopecilo  ,  infundibu- 
li  toimi ,  pIcru!Tiqiietnfa,riamdiviro,  fed  fterili.  Embryoncs 

cnim  C  a  floribus  fcjunguntur  in  cadem  plsnta  &:abcunt  dcin- 

de  in  frutftum  D  tricoccum  ,  ex  tribus  fcilicet  capfulis  E  compo- 

Ctum  ,  femine  foccis  fphartico  F. 
Tragii  fpecles  func. 

Tragia  fcandens  longo  Becontcx  folio. 

Tragia  alia  fcandens  uriicx  folio. 

Hitronimus  Tragus  alias  Hifronimus  Bo^ius  vemdcuJe  Sock,  dc 
Meidejpach ,  na:us  cnim  eji  HeideJpachUfub  an.  1 498.  B:ponti  w- 
xic  per  alitjuot  artnos  uhl  hcrrum  Ducis  Ludovid  Palatini  Jtheni 
varji generis  plantis  injlruxit.  Inde  Harnhatum^  poj^ea  Saraponturrt 
fc  confer  em  Htjloriam  hcrbarum  condidit,  in  e^ua  in  tres  Ltbros  divi- 
fa  de  Jiirpium  dijferenriis  &  facutiatibus  fufe  dijferir.  Mortuits^ 
eft  autem  an.  1554.  21  Febr.  £tat.  ^6.  txtar  o^us  jirgentorad 
IJJI.  in  ^u^rr. 

Fuchsia. 

FUchfiaeft  plmtx  genus  flore  A  monoperalo  ,  infundibuli- 
forrci  &  mulu-fido  ;  cujus  calyx  B  abic  delnde  in  fruftuiu 

C  fubrorundum  ,  mollem  ,  carnofum  ,  in  quacuoi  loculamenta 

divifum  D,  fcmimbjlque  factum  fubrocundis  E. 
Fuchfix  unlearn  fpccietn  agnovi. 

Fuchfia  iriphylla  ,  flore  coccineo. 

leonartuj  Furhfus  Vembding£  ,  Rhxda  Oppido  in  dicione  Vucnrri 
Baviridt  natui  1501.  Mcdlcin£  Do6lor  renuncia/us  Monachum 
adiit  t  deinde  Tubir.^m  1  ubi  35.  annis  prxcUre  docuit.  Vir 
fwt  d^idul  (abo/is  ,  piantarum  Cermanit  dili^ens  explordtor. 
Magna  litrbdriorHm  comrnodo  hdrum  iconcs  jio.  ampUorit  for- 


A  M  E  R  r  c.    G  E  N  E  a.  A.  ty 

mt  exhibttlt.   Tiibing£  mortuus  eft  anno  ij65.  lo.  Maji  ,  dta-  M*Uhm 
tis  6j.  Scripfit  de  Hiftoria  ftirpium  Commtntari^sinfignss  \  Bd"'^^^ 
files  i|4i<  in  folio* 

RONDELETIA. 

ROndeletia  eft  p!antar  genus  fiore  A  raonopetalo  ,  hypocra-r**.  m 
tcri-formi ,  cubulaco  8c  calyci  C  infidcnce.    Is  autem  ca- 
lyx abit  deinde  in  fruiium  D  fubrotundum  >  coronatum ,  bicap* 
fularem  E  >  &  feminibus  factum  exiguis  F. 

Rondeletis  unicam  fpeciem  novi. 
Rondcletia  arborefcens  ,  tini  facie. 

Cuillflmus  Rondelctius  MonfpeJJiiU  natus  >  dnno  1^07.  die  17. 
Sept.  Medicind  arte  prtceilem ,  de  pijcibus  &•  de  pifcium  natu- 
Yd  pr£cUrum  Opus  condidit  1  fed  etiam  fmplicibus  medicament 
tis  dignofcendis  ^  inveniendis  fumtnam  operam  coUocavit  ,  at-  TamffAn^, 
que  in  eo  multum  excelluit ,  Diofcoridrm  primus  MonfpeUt  enar-*  '  '*'" 
rans.  Plura  in  Diofcoridem  Jcripffte  apparet  ,  ex  Epiftch's  GeJ^ 
neri  ad  I.  Bauhinum  ,  a  C.  Bauhino  edltts.  Cancellarius  ScnO' 
Id  Monfpelienfts  mortuus  eft  anno  1^66. 

T  U   R   N    E   R   A. 

TUrnera  eft  plants  genus  flore  A  monopctalo  ,  inftindibu-  Tab.  n: 
H-formi  fie  multi-fido  ;  ex  cujus  calycc  C  bicoini  furgjc 
piftillum  infima:  flciis  parti  B ,  ad  inftar  clavl  intixum  .  quod 
deinde  abic  in  frudum  D  fere  globofum  aut  turbinatum  ,  trila- 
riam  dehifcentem  E  ,  feminibufquc  fcctum  F  fubrotundis  ,  te* 
Duibus  capillamemis ,  ceu  placenta:  affixis. 

Turners  unicam  vidi  fpecism. 
Turaera  fi-utefccns  ,-  ulmi-folia. 

Cuiltelmus  TumerHS  yinglus  y  Medicine  VoSlor  ,  vir  JoUdt  eru- jtalut  fr-fl 
ditionis  (St-  judtcli  ,.  emijn  plantarum  Hlftoriam  jfngitt ,  anno  f^'**- 
1^51.  in  qua  Flgurds  Fuchfii  plerumque  adhtbitit  t  nomlna  expref 
fit  latlne ,  grdce,  dngUce »  germankc  >  gdllicc  ,  ordinem  atplutm 
heticum  JecHtus. 


95 


[137] 


PROCEEDINGS 

OF  THE 

CALIFORNIA  ACADEMY  OF  SCIENCES 

Fourth  Series 
Vol.  XXV,  No.  2,  pp.  139-146  Jiiiie  1,  1944 


ALICE  EASTWOOD 
SEMI-CENTENNIAL  PUBLICATIONS 

No.  2 

RELATIONS  OF  THE  TEMPERATE  FLORAS  OF 
NORTH  AND  SOUTH  AMERICA 

BY 

DOUGLAS  HOUGHTON  CAMPBELL 

Professor  of  Botany,  Emeritus 
Stanford  University 

THE  DiSTRiBUTiox  of  the  existing  land  floras  of  the  world,  as  well  as  their 
fossil  record,  have  been  studied  by  many  botanists  for  a  long  time.  These 
botanists  have  been,  for  the  most  part,  Europeans  or  Americans,  and  have 
especially  studied  the  floras  of  the  Northern  Hemisphere ;  and  the  history  of 
the  Holarctic  vegetation  is  fairly  well  known.  There  is  much  less  information 
concerning  the  history  of  the  floras  of  the  Southern  Hemisphere,  especially 
the  fossils.  Since  we  now  have  as  allies  most  of  the  South  American  countries, 
it  might  be  well  for  us  to  become  better  acquainted  with  the  botanists  of  South 
America  and  their  studies  of  the  floras  of  those  countries. 

Great  changes  have  occurred  during  past  ages  in  the  distribution  of  land 
and  water  of  the  globe;  and  great  mountain  ranges  now  stand  where  once  the 
sea  covered  the  land.  These  changes  in  the  distribution  of  the  main  bodies  of 
land  and  water  have  undoubtedly  had  much  to  do  with  fluctuations  of  climate 
and  the  distribution  of  many  plants. 

Thus  the  invasion  of  the  sea  in  the  Cretaceous  Age  over  much  of  the  west- 
ern United  States,  and  later,  the  elevation  of  the  Rocky  Mountains  and  the 
Sierra  Nevada  and  Cascades,  had  a  very  marked  effect  on  the  climate  of  those 
regions  and  the  distribution  of  the  vegetation.  During  the  Mid-Tertiary,  for 
example,  the  redwood  (Sequoia)  now  practically  restricted  to  California, 
occurred  over  much  of  North  iVmerica  and  Eurasia,  and  the  southern  cypress 
(Taxodium)  now  confined  to  the  southeastern  States  was  equally  widespread. 

The  great  Pleistocene  glaciation  caused  a  great  redistribution  of  the  floras 

[  139  ] 


140  CALIFOEXIA   ACAD  f:  MY  OF  SCIENCES        [Proc.  4th  ser. 

of  North  America  and  Eurasia,  and  many  American  trees,  like  the  tulip  tree 
(Liriodendron),  magnolia  and  walnut,  before  the  Ice  Age  lived  also  in  Eu- 
rope but  were  completely  exterminated  there  at  that  time. 

Existing  Factors  ix  Distribution 

Among  the  factors  of  prime  importance  in  the  distribution  of  the  existing 
floras,  the  first  is,  perhaps,  the  relations  of  the  great  continental  masses  to 
each  other,  and  to  the  surrounding  oceans.  The  latter  are  effective  barriers  to 
the  migration  of  all  but  a  very  few  plants,  and  high  mountains  and  extensive 
areas  of  desert  and  arid  regions  such  as  occur  in  the  center  of  the  continents, 
also  are  barriers  to  migration  ;  but  mountains  may  also  serve  as  liigliAvays — as 
may  be  seen  in  the  occurrence  of  arctic  and  sub-arctic  species  in  the  Sierra 
Nevada  and  Rocky  Mountains,  far  south  of  their  normal  habitat,  and  many 
plants  are  common  to  Chile  and  California. 

The  major  part  of  the  land  surface  of  the  globe  is  in  the  Northern  Hemi- 
sphere and  comprises  North  America  and  Eurasia.  It  is  generally  recognized 
that  these  two  continents  have  for  a  very  long  time  been  more  or  less  inti- 
mately connected.  This  is  indicated  by  the  evident  close  relationship  of  their 
floras,  especially  in  the  higher  latitudes.  Throughout  the  Arctic  and  Sub- 
arctic Zones  most  of  the  trees,  shrubs  and  herbaceous  species  are  closely  re- 
lated, or  even  identical.  It  has  even  been  proposed  to  consider  North  America 
and  Eurasia  as  parts  of  an  ancient  primary  continent,  "Laurasia." 

The  southern  continents.  South  America,  Africa,  Australia  and  Antarctica, 
are  separated  from  each  other  by  the  width  of  the  great  oceans.  Except  for 
Antarctica  which  is  destitute  of  terrestrial  vegetation,  the  southern  continents 
have  varied  and  peculiar  floras,  indicating  a  long  period  of  isolation.  Never- 
theless they  all  have  many  obviously  related  genei-a,  and  even  species,  point- 
ing to  some  ancient  land  connections. 

It  has  been  held  by  some  geologists  that  in  the  late  Palaeozoic  there  existed 
a  great  southern  continent,  "Gondwana,"  which  included  all  the  existing 
southern  continents,  and  also  India.  Gondwana  was  separated  from  the  north- 
ern land-masses  by  a  broad  oceanic  belt,  the  "Tethys  Sea"  which  persisted  to 
near  the  end  of  the  Mesozoic.  This  perhaps  explains  the  radical  differences 
between  the  present  temperate  floras  of  North  and  South  America. 

How  the  existing  southern  continents  became  so  completely  isolated  has 
caused  much  speculation.  In  a  recent  study  of  the  subject  by  a  South  African 
geologist,'  the  author  concludes  that  Gondwana  split  into  segments  which 
drifted  apart  and  developed  into  the  present  continents.  This  theory  of  Con- 
tinental Drift,  put  forward  by  Wegener  in  1912,  is  a  plausible  explanation  of 
many  puzzling  facts  in  geographical  distribution,  both  of  animals  and  plants. 

The  greater  part  of  North  America  and  Eurasia  lies  north  of  the  Tropic  of 
Cancer,  and  extends  beyond  the  Arctic  Circle.  Almost  the  whole  of  the  United 
States  lies  in  the  "Temperate"  Zone,  between  30°  and  50°  N.  For  the  most 

^  Our  Wandering  Continents.  .\.  T..  r>n  Toit :  London,  1937. 


Vol.  XXV]   CAMPBELL:  FLORAS  OF  NORTH  AND  SOUTE  AMERICA    141 

part  the  climate  is  distinctly  "continental,"  with  great  annual  range  of  tem- 
perature. In  general,  winter  is  a  season  of  complete  cessation  of  plant  activity. 
In  the  United  States  milder  climates  prevail  in  the  Southern  States  and  espe- 
cially on  the  Pacific  Coast,  where  weather  conditions  are  very  different  from 
those  over  most  of  the  country.  From  Sitka  to  San  Diego,  the  climate  is 
remarkably  equable.  The  January  isotherm  of  0°C,  which  on  the  Atlantic 
Coast  is  in  the  region  of  New  York  and  Philadelphia,  on  the  Pacific  Coast  is 
pushed  north  to  Sitka  (lat.  57°  N.).  In  San  Francisco,  there  is  only  a  differ- 
ence of  ten  degrees  Fahrenheit  between  the  coldest  and  warmest  months  (50° 
to  60°)  ;  in  Washington,  D.C.,  about  the  same  latitude  (32°  to  78°). 

The  lofty  barrier  of  the  great  Cordillera  protects  the  coastal  areas  from  the 
great  temperature  fluctuations  of  the  interior  regions,  and  the  prevailing 
westerly  winds  from  the  Pacific  have  a  great  influence  on  both  temperature 
and  precipitation. 

In  the  strictly  arctic  regions,  e.g.,  Greenland,  northern  Canada  and  Alaska, 
trees  are  absent  and  the  vegetation  is  made  up  for  the  most  part  of  perennials, 
either  herbaceous  species  with  subterranean  roots  or  root-stocks  which  during 
the  brief  growing  season  of  perhaps  two  months,  complete  their  season's 
growth;  or  a  few  woody  species,  e.g.,  dwarf  willows  and  birches,  dwarf  rhodo- 
dendrons, cranberries  and  a  few  other  bog-plants  may  be  found.  In  specially 
favorable  places  Iceland  poppies,  buttercups,  saxifrage,  with  showy  flowers,- 
as  well  as  some  grasses,  quickly  develop  from  their  perennial  roots  during  the 
very  brief  summer  months. 

Further  south  in  the  Sub-arctic  Zone,  trees  begin  to  play  an  important  role. 
Thus  in  Canada,  Scandinavia,  Russia,  there  are  extensive  forests  of  spruce 
and  pine,  birches,  willows  and  poplars  and  various  deciduous  shrubs,  much 
the  same  throughout,  from  Alaska  to  Russia. 

The  Sub-arctic  merges  gradually  into  the  North  Temperate  which  occupies 
much  of  North  America  and  Eurasia.  In  the  United  States,  especiallj^  in  the 
eastern  portion,  this  zone  is  characterized  by  an  extraordinary  variety  of 
deciduous  trees — oaks,  beech,  ash,  elm,  walnut,  hickory,  chestnut,  and  many 
others,  which  occur  over  most  of  the  region  east  of  the  Mississippi. 

About  two-thirds  of  the  United  States  lie  between  the  Rocky  Mountains  and 
the  Atlantic  Ocean.  This  includes  the  great  Mississippi  Valley  and  the  Great 
Lakes.  The  vast  area  between  the  Rocky  Mountains  and  the  Appalachians  has 
no  barriers  to  plant  migration  except  climatic  ones,  e.g.,  temperature  and 
rain-fall,  and  many  species  are  distributed  over  most  of  the  area.  Thus  several 
species  of  oak,  the  black  walnut  and  American  elm  occur  over  most  of  the  area 
eastward  of  the  Rocky  Mountains,  except  where  the  moisture  is  insuffieient. 

The  rainfall  decreases  westward  and  the  forest  gives  way  to  the  prairies. 
West  of  the  Mississippi  trees  usually  occur  only  along  streams.  The  prairies 
finally  pass  into  the  semi-arid  steppes  which  occur  at  the  base  of  the  Rocky 
Mountains.  These  plains  are  characterized  by  scattered  bunch  grasses  and  low 
shrubs,  together  with  some  herbaceous  species. 


LIBRARY    a 


142  CALIFOBXIA  ACADEMY  OF  SCIEXCES        [Proc.  4th  ser. 

The  "western  tliird  of  North  America  is  very  different  from  the  East.  It  is 
a  region  of  mountains  and  elevated  plateaus  between  the  Rocky  JMountains 
and  the  Pacific  Coast.  This  region  includes  extensive  plateaus  and  secondary 
mountain  ranges.  Much  of  the  plateau  region  is  arid  and  in  some  places,  as 
in  the  Salt  Lake  region,  and  parts  of  Nevada,  true  deserts. 

The  great  Pacific  Cordillera  which,  with  few  breaks,  extends  from  Alaska 
to  Patagonia,  sharply  sets  off  the  Coastal  Region  from  the  rest  of  the  Conti- 
nent. The  whole  of  the  western  Mountain  area  is  very  different  climatically 
and  floristically  from  Atlantic  North  America,  and  is  the  only  portion  of  the 
United  States  in  which  the  highest  mountains  have  perpetual  snow  and  a  true 
alpine  flora. 

Southern  California  and  western  Mexico  have  very  light  rainfall  and  much 
of  the  country  is  desert,  but  the  precipitation  increases  rapidly  northward, 
and  in  northwestern  California  the  rainfall  is  very  heavy;  and  from  central 
California  northward  through  Oregon,  and  Washington  to  Alaska,  there  is 
heavy  rainfall,  resulting  in  the  development  of  the  greatest  coniferous  forest 
in  the  world. 

In  the  Gulf  region,  especially  Florida,  there  are  many  tj'pes,  like  the  palms, 
Ficus  and  Bromeliads,  evidently  derived  from  the  tropical  regions  to  the 
south. 

Central  California  is  a  meeting  ground  for  the  boreal  floras  of  Noi'tli 
America  and  the  subtropical  floras  of  Mexico.  There  is  also  a  considerable 
number  of  genera  and  even  species  common  to  California  and  central  Chile. 

The  coastal  ranges  of  central  California,  with  their  cool,  moist  forests,  con- 
tain many  distinctly  boreal  types,  like  most  of  the  conifers,  as  well  as  such 
deciduous  trees  as  maples,  oaks  and  alders,  and  deciduous  shrubs,  honey- 
suckle, roses,  elder,  spiraea,  azalea,  rhododendron,  dogwood,  and  others,  and 
mau}^  characteristic  herbaceous  flowers,  e.g.,  Trillivm,  Erythronium,  Ranun- 
cnlns,  Oxalis,  saxifrage,  AquiJegia,  violets,  and  many  others. 

With  decreasing  precipitation  southward,  these  northern  types  are  gradu- 
ally replaced  by  genera  and  species  absent  from  the  eastern  States,  or  only 
occasionally  found,  e.g.,  Acacia,  cacti,  Yucca,  Prosopis,  Mimtihts,  Penstemon, 
Castilleja,  and  others. 

The  occurrence  in  California  of  a  considerable  number  of  species  found  also 
in  Chile  is  remarkable.  Among  these  may  be  noted  Fragaria  chilensis,  Pro- 
sopis juliftora,  Mesemhryanthemum  aequilaterale,  Calandrinia  Menziesii. 

The  temperate  regions  of  South  America  include  a  small  part  of  southern 
Brazil,  Paraguay,  Uruguay,  Argentina,  and  Chile ;  the  two  latter  comprise 
the  greater  part  of  temperate  South  America.  Between  Chile  and  Argentina 
is  the  great  barrier  of  the  Andes,  a  condition  similar  to  that  separating  the 
coastal  region  of  California  from  Nevada  and  Arizona,  and  there  is  much  the 
same  difference  between  the  coastal  and  inland  climates  in  both  cases,  as  well 
as  in  the  vegetation. 

The  central  and  southern  parts  of  Chile  have  abundant  rainfall,  while  in 


Vol.  XXV]   CAMPBELL:  FLOBAS  OF  NOBTH  AXB  SOUTH  AMERICA   143 

most  of  Argentina  the  precipitation  is  scanty,  and  much  of  the  country  is  a 
desert.  The  continent  diminishes  in  breadth  southward,  its  southernmost 
point,  Cape  Horn,  being  less  than  60°S. 

Owing  to  the  proximity  of  the  oceans,  the  climate  is  insular  rather  than 
continental,  and  there  is  nothing  comparable  to  the  Arctic  Zone  of  the  north- 
ern continents.  Similar  conditions  prevail  in  temperate  South  Africa  and 
Australia,  where  the  southernmost  regions  are  under  50°S.  Excepting  in  the 
high  mountains  there  is  no  such  difference  between  winter  and  summer  as  ex- 
ists in  corresponding  latitudes  in  the  Northern  Hemisphere.  Thus  in  Ushuaia, 
in  Tierra  del  Fuego  (lat.  55.5°S)  the  coldest  monthly  average  is  28.9°F,  the 
warmest  50°  ,•  at  Nain  (Labrador)  in  about  the  same  latitude  North,  the  range 
is  -7°F  to  46°;  in  Buenos  Aires  (lat.  35.5°S)  the  coldest  month  is  50°F,  the 
hottest,  73.6°.  St.  Louis,  about  the  same  latitude  North,  has  the  range  of  31° 
to  79.1°. 

As  a  whole  the  temperate  floras  east  of  the  Andes  are  very  poor  compared 
with  corresponding  areas  in  North  America.  While  the  flora  of  Argentina  is 
mainly  related  to  the  Brazilian  regions  to  the  north,  there  are  also  elements 
related  to  those  of  Chile,  especially  in  the  extreme  south,  where  there  is  abun- 
dant rainfall,  and  where  the  so-called  "Subantarctic"  flora  becomes  evident. 
This  flora  is  also  related  to  that  of  New  Zealand  and  South  Australia. 

The  transition  from  southern  Brazil  to  Argentina  is  a  gradual  one  with 
increasing  aridity  southward.  The  strip  at  the  base  of  the  Andes  is  very  dry, 
with  scanty  vegetation,  and  might  be  compared  to  the  most  extreme  deserts 
of  Arizona  and  Nevada.  It  has  been  described  as  one  of  the  most  dismal  and 
sterile  regions  that  one  could  imagine.  The  precipitation  increases  somewhat 
toward  the  east,  and  much  of  central  Argentina  is  occupied  by  the  great 
pampas,  recalling  the  great  plains  of  the  central  United  States,  and  like  the 
prairies  there  is  a  great  development  of  grasses  and  other  herbaceous  plants. 
The  constituents  of  the  pampas  floras  are  for  the  most  part  quite  different 
species  from  those  in  North  America.  In  general  the  flora  of  northern  Argen- 
tina is  evidently  related  to  that  of  southern  Brazil. 

Southward  the  pampas  merge  into  the  arid,  sterile  plains  of  Patagonia, 
whete  only  such  scanty  vegetation  is  found  as  can  survive  the  stony,  exposed, 
and  excessively  dry  conditions  which  prevail. 

The  conditions  in  Chile,  west  of  the  Andes,  are  very  different  from  those 
in  Argentina.  The  coast  of  Chile  extends  from  16°  57'  S  to  55  S  (the  southern- 
most point  in  South  America — Cape  Horn).  This  corresponds  to  the  north 
Pacific  Coast  from  Lower  California  to  Alaska. 

The  northern  coast  of  Chile  is  practically  rainless,  and  much  of  the  region 
is  quite  destitute  of  vegetation.  Passing  southward,  the  precipitation  gradu- 
ally increases,  and  in  the  central  region,  e.g.,  Valparaiso,  Santiago,  conditions 
are  very  similar  to  those  in  Central  California,  and  the  general  character  of 
the  vegetation,  both  native  and  cultivated,  is  very  much  the  same. 

Trees  and  shrubs  from  California,  e.g.,  Sequoia,  Monterey  Cypress,  Ceano- 


144  CALTFOFXIA  ACADEMY  OF  SCIEXCES        [Proc.  4th  ser. 

thus,  are  planted  in  the  parks  and  gardens,  and  the  California  poppy  {Esch- 
scholtzia)  is  quite  naturalized.  In  California  the  Chilean  pepper  tree  {Schinus 
molle),  and  Araucaria  imhricata,  are  familiar  trees,  and  Escalloriia  spp.  and 
Buddleia  glohosa  are  common  ornamental  shrubs. 

Central  Chile,  both  in  climate  and  topography,  much  resembles  the  coi-re- 
sponding  regions  in  California,  and  the  great  central  valley  between  the 
Andes  and  the  Coast  ranges  may  be  compared  with  the  great  central  valley 
of  California.  Santiago  (lat.  35°S),  the  capital,  resembles  almost  exactly  in 
its  rainfall  and  mean  temperature,  San  Jose  (lat.  37°N)  in  the  Santa  Clara 
Valley  of  California. 

The  rainfall  in  southern  Chile  is  very  heavy  and  there  is  present  a  luxuri- 
ant rain  forest  very  much  as  the  heavy  coniferous  forest  is  developed  on  the 
northern  Pacific  Coast.  The  constituents  of  these  forests  are  very  different ; 
while  the  northern  Pacific  forests  are  composed  almost  exclusively  of  con- 
ifers— redwood,  spruce,  fir,  hemlock,  etc. ;  conifers  are  relatively  rare  in  the 
Chilean  forests  and  are  all  different  from  the  northern  genera,  and  are  most 
nearly  related  to  those  of  New  Zealand  and  Australia.  The  rain  forest  of  Chile 
is  mostly  composed  of  broad-leaved  evergreens,  mostly  lacking  in  the  northern 
forests,  and  includes  many  genera  common  to  New  Zealand,  e.g.,  evergreen 
beech  (Xothofagus),  Drimys  (Magnoliaceae) ,  Laurelia  and  Weinmannia,  both 
important  genera  in  New  Zealand,  and  also  Fuchsia  and  Griselinea.  Charac- 
teristic of  the  forests  of  the  South  Temperate  Zone  are  many  species  of  the 
Myrtle  family,  almost  completely  absent  from  the  North  Temperate  Zone, 
Eucalyptus  from  Australia  being  perhaps  the  best  known  example.  Another 
exclusively  southern  family,  Proteaceae,  especially  developed  in  South  Africa 
and  Australia,  is  well  represented  in  South  America.  Emhothrium  and  Lo- 
matia  occur  in  Chile. 

While  there  is  nothing  corresponding  to  the  Arctic  Zone  of  the  Northern 
Hemisphere,  southern  Chile  has  a  flora  which  is  unmistakably  related  to  that 
of  New  Zealand  and  southern  Australia.  This  flora  has  been  called  "Subant- 
arctic"  and  indicates  some  former  land  connections  with  New  Zealand  and 
southern  Australia. 

While,  in  general,  there  is  nearly  a  complete  absence  in  South  America  of 
characteristic  North  American  genera  and  species,  in  the  Pacific  Coastal  Re- 
gion there  is  a  remarkable  number  of  genera  and  even  species  common  to 
Chile  and  central  and  southern  California,  and  the  adjacent  southwestern 
areas.  These  may  be  explained  as  due  to  migration  along  the  great  mountain 
system  of  the  Pacific  Coast,  subsequent  to  the  union  of  South  America  and 
Central  America. 

Some  of  these  are  cosmopolitan  tyi)es — like  Caltha,  Linnm,  Geranium, 
Oxalis,  but  a  considerable  number  are  species  common  to  California  and  Chile 
but  unknown  elsewhere,  e.g.,  Fragaria  Chilensis,  Calandrinia  Menziesii,  Lupi- 
nus  microcarpa,  Madia  sativa,  Plantago  magellanica,  Gilia  pusilla,  Prosopis 
juH  flora. 


Vol.  XXV]   CAMPBELL:  F LOEAS  OF  NOETH  AND  SOLTR  AMEEICA   145 

Fossil  Evidence 

While  the  fossil  record  is  necessarily  incomplete  it  has  added  much  to  our 
knowledge  of  the  history  of  the  higher  plants,  especially  in  North  America 
and  Eurasia. 

The  fossil  remains  of  plants  are  restricted  largely  to  such  parts,  e.g.,  stems, 
leaves,  roots,  as  have  firm  tissues,  resistant  to  decay.  Occasionally  the  tissues, 
even  the  more  delicate  ones,  are  replaced  by  infiltration  of  mineral  substances, 
lime  or  silica,  which  replace  the  organic  matter  of  the  cell-walls,  and  result 
in  petrifactions ;  but  as  a  rule  the  more  delicate  organs,  such  as  the  flowers, 
are  very  seldom  found  as  fossils. 

Much  has  been  learned  of  the  fossil  floras  of  the  geological  eras,  from  the 
Devonian  where  the  earliest  certain  remains  of  land  plants  have  been  found, 
to  the  Mesozoic  and  Tertiary  formations,  which  contain  many  remains  of 
plants,  evidently  closely  related  to  living  species. 

From  a  study  of  these  ancient  fossils  it  is  evident  that  as  a  whole  plants  are 
very  conservative,  and  a  remarkably  large  number  of  forms  have  come  down 
from  a  very  remote  past  to  the  present  with  little  change.  The  common  "horse- 
tail" {Equisetum)  and  the  "club-mosses"  (Lycopodium)  are  very  similar  to 
some  of  their  relations  from  the  Carboniferous,  and  many  fossils  of  the  mod- 
ern flowering  plants  (Angiosperms)  as  far  back  as  the  Cretaceous,  when  they 
are  first  certainly  recognizable,  can  be  referred  to  such  living  genera  as  syca- 
more (Plat an  us) ,  sassafras,  oak,  poplar,  and  many  others.  It  is  to  be  expected 
that  many  common  herbaceous  plants  which  now  are  associated  with  these 
woody  species  also  existed,  but  owing  to  their  delicate  perishable  tissues  have 
not  survived  as  fossils. 

Many  of  these  fossils  occur  in  regions  now  unsuited  to  their  growth,  showing 
that  great  climatic  changes  have  taken  place.  Thus  among  the  earliest  Angio- 
sperms that  have  been  found  were  some  described  from  Greenland,  from  rocks 
supposed  to  be  of  Cretaceous  age.  These  fossils  represent  genera  now  found 
in  regions  many  degrees  southward.  Among  these  fossils  were  sycamores, 
magnolias,  cinnamon,  and  others.  Sycamore  (Plafanus)  is  now  represented 
in  the  United  States  by  two  species,  one  occurring  from  New  England  to  the 
Mississippi,  the  other  in  central  and  southern  California.  It  is  hard  to  im- 
agine any  possible  conditions  of  climate  in  which  these  trees  could  have  grown, 
within  a  few  hundred  miles  of  the  North  Pole,  deprived  of  sunlight  for  many 
months.  A  similar  problem  is  offered  by  fossils  found  near  the  Antarctic. 

While  our  knowledge  of  the  fossil  floras  of  South  America  is  much  less  com- 
plete than  that  of  North  America,  there  have  been  some  important  contri- 
butions, especially  Professor  E.  W.  Berry's  studies  in  the  Tertiary  floras  of 
Argentina  (Tertiary  Flora  from  the  Rio  Pichileufu,  Argentina,  Proc.  TJ.  8. 
Nat.  Mu.se7(m,  Vol.  72,  1928). 

This  flora  includes  elements  now  belonging  to  the  present  subtropical  floras 
of  Brazil,  Paraguay,  and  also  genera  now  confined  to  the  extreme  south  of 


140  CALIFOBXIA  ACADEMY  OF  SCIENCES        [Proc.  4th  ser. 

Patagonia,  Tierra  del  Fuego,  and  Chile,  representing  the  "Snbantarctic" 
regions.  This  indicates  that  the  Tertiary  elimate  in  Patagonia  was  both 
warmer  and  more  humid  than  at  present.  There  were  no  North  American 
genera  present,  but  the  most  remarkable  case  was  Ginkgo,  unknown  else- 
where in  South  America. 

A  recent  study  of  fossil  conifers  from  Chile  (The  Tertiary  Fossil  Conifers 
of  South  Chile  and  their  Phytogeographical  Significance)  by  Rudolph  Florin, 
Stockholm,  1940,  shows  that  the  genera  were  the  same  as  now  exist,  and  that 
none  of  the  characteristic  North  American  genera,  e.g.,  Pinus,  Abies,  Tsuya, 
Picea,  were  present.  Florin  states  that  the  same  complete  separation  of  the 
northern  and  southern  types  has  existed  since  the  late  Palaeozoic. 

From  a  study  of  the  existing  floras  as  well  as  the  fossil  forms,  one  may 
conclude  that  the  floras  of  South  America  have  developed  independently,, 
except  for  the  so-called  "Subantarctic"  floras,  which  probably  originated  in 
some  common  land-mass,  perhaps  the  hj'pothetical  Gondwana.  This  seems 
much  more  likely  than  that  they  originated  in  some  northern  extension  of 
the  present  Antarctica. 

Acceptance  of  the  recent  hypothesis  of  Du  Toit,  that  there  were  two  pri- 
mordial continents,  Laurasia  in  the  Northern  Hemisphere  and  Gondwana  in 
tlie  South,  and  from  these  primary  continents,  the  existing  continents  were 
separated  and  shifted  to  their  present  positions,  would,  if  true,  remove  most 
of  the  difficulties  in  explaining  the  present  distribution  of  many  existing 
l^lant  families. 


I 


PROCEEDINGS 
OF  THE 

CALIFORNIA  ACADEMY  OF  SCIENCES 

Fourth  Series 
Vol.  XXV,  No.  3,  pp.  147-170,  pi.  17  Ji^ii^e  1,  1944 


ALICE  EASTWOOD 
SEMI-CENTENNIAL  PUBLICATIONS 

No.  3 
PUGILLUS  ASTRAGALORUM  ALTER 

BY 

R.  C.  BARNEBY 


THE  OBSERVATIONS  on  Astragalus  contained  in  the  present  paperf  are  based 
primarily  on  material  gathered  by  Mr.  H.  D.  Ripley  and  the  writer  in 
California,  Nevada,  Utah,  and  Arizona  during  the  past  two  or  three  years. 
Through  the  kindness  of  Dr.  P.  A.  Munz  it  has  been  possible  to  make  critical 
comparisons  with  a  number  of  types  in  the  herbarium  of  Marcus  E.  Jones 
at  Pomona  College  (PO),  without  which  these  notes  could  not  have  been  pre- 
pared. In  addition,  the  writer  has  had  an  opportunity  to  visit  several  herbaria 
in  the  East,  and  there  to  study  type  and  other  specimens  not  represented  in 
California  collections.  To  the  gentlemen  in  charge  of  these  herbaria,  in  par- 
ticular to  Prof.  M.  L.  Fernald  of  the  Gray  Herbarium  at  Harvard  Universitj^ 
(G),  Dr.  H.  A.  Gleason  of  the  New  York  Botanical  Garden  (NY)  and  Dr. 
P.  C.  Standley  at  the  Field  Museum  (F),  who  generously  placed  at  my  dis- 
posal the  collections  in  their  care,  I  wish  to  express  my  lively  appreciation. 
I  am  also  especially  grateful  to  Miss  Alice  Eastwood  for  the  communication 
of  the  type  specimen  of  Astragalus  Bryantii,  described  below.  The  capital 
letters  in  parentheses  have  been  used  throughout  the  paper  to  denote  the 
herbarium  in  which  cited  material  has  been  seen. 

Types  of  all  species  and  varieties  described  as  new  are  deposited  in  the 
herbarium  of  the  California  Academy  of  Sciences  together  with  duplicates 
of  the  collections  discussed. 

t  Mr.  Barneby's  earlier  paper  on  Astragalus.  "Pugillus  Astragalorum  Nevadensium," 
was  published  in"  Leaflets  of  Western  Botany  3  :  97-114,  figs.  A-F.  1942. 


[147  1 


148  CALIFORNIA   ACADEMY  OF  SCIENCES        [Proc.  4th  ser. 


Astragalus  diversif olius  A.  Gray 

Astragalus  diversif  olius  A.  Gray,  Proc.  Amer.  Acad.  6:230.  1864.  HomnJohus  orthocarpns 
NuTT.  ex  T.  and  G.,  Fl.  N.  Amer.  1:351.  1838.  A.  convallarius  Greene  var.  diversifolius 
(A.  Gray)  Tidestr.,  Proc.  Biol.  Soc.  Wash.  50:20.  1937. 

Utah:  clay  slopes  among  junipers,  3  miles  south  of  Enterprise,  Washington  Co.,  alt. 
6300  ft.  Ripley  and  Barnehy  4967. 

This  station  extends  the  range  of  tlie  species  as  known  to  the  writer  almost 
to  the  borders  of  Arizona,  to  which  state  it  is  accredited  by  Tidestrom  (Fl. 
Ariz,  and  N.  Mex.  197.  1941),  thouyh  not  admitted  by  Kearney  and  Peebles, 
who  record  only  A.  jnnciformis  A.  Nels.  of  this  immediate  group  as  native 
to  the  state.  Prof.  Tidestrom's  plant  is  unknown  to  me,  but  his  combination 
is  in  any  case  untenable.  If  the  plants  originally  named  by  Nuttall  as  Homa- 
lobns  orthocarpus  and  H.  campestris  cannot  be  separated  specifically,  the 
oldest  valid  binomial  for  either  element  of  the  complex  must  be  given  the 
rank  of  species,  not  reduced  to  varietal  status  under  a  subsequently  published 
name.  A.  diversifolius  A.  Gray,  founded  on  H.  ortJiocarpus,  appeared  29  years 
earlier  than  A.  convallarius  Greene  (1893),  which  was  proposed  to  replace 
A.  campestris  (Nutt.)  A.  Gray  (1864)  non  Linne  (1753).  Even  if  the  com- 
bination A.  diversifolius,  often  wrongly- applied,  were  discarded  as  a  nomen 
confusum,  A.  ihapensis  M.  E.  Jones,  in  all  probability  an  exact  synonym  of 
H.  orthocarpus  and  published  in  January  of  1893,  Avill  take  precedence  over 
A.  convallarius  published  in  October  of  the  same  year.  But  the  taxonomy  and 
involved  nomenclature  of  this  group  are  still  in  need  of  critical  revision. 

Astragalus  Whitedii  Piper  fma.  speirocarpoides  Barneb}',  forma  nova 

(Platel7,  fig.  31) 

Astragalus  Whitedii  Piper  fnia.  speirocarpoides  Barneby  fina.  nov.  a  prole  tvpica  (seu 
A.  speirocarpo  var.  falciformi  A.  Gray)  legumine  in  annulum  saltem  conipletuni  vel 
saepius  sesquialterum  contorto,  ncc  falcate  tantum,  diversa. 

Nevada:  in  deep  sandy  soil  among  sagebrush,  10  miles  north  of  Winnemucca,  Hum- 
boldt Co.,  alt.  4500  ft.,  23  May,  fl.  and  fr.  Riphi/  ai\d  Barnehy  45.',2.  Type  in  Herb. 
Calif.  Acad.  Sci.  No.  300414.  Also  at  Austin,  Lander  Co.,  Jones,  16  June  1882  (PO, 
annotated  as  A.  Gibbsii  var.). 

With  its  fruit  twisted  into  one  or  one  and  a  half  spiral  coils,  the  proposed 
forma  is  very  similar  in  aspect  to  the  true  A.  speirocarjms  A.  Gray  of  Wash- 
ington and  northern  Oregon,  but  the  long  calyx  (tube  8  mm.),  rather  coarse, 
procumbent  stems  and  broad  leaflets  are  all  characteristic  of  A.  Whitedii, 
in  the  typical  form  of  which  the  pods  are  not  more  than  strongly  falcate.  It 
should  be  noticed  that  the  fma.  speirocarpoides  invalidates  Rydberg's  ke.y  to 
Homalohus  Ser.  Collini  (N.  Amer.  Fl.  24:  258.  1929),  from  which  this  plant 
would  appear  to  belong  to  A.  speirocarpus. 


Vol.  XXV]     BABNEBY:  PUGILLUS  ASTRAGALOEVM  ALTEE  149 

Astragalus  zionis  M.  E.  Jones 

Astragalus  zionis  M.  E.  Jones.  Xylophacos  zionis  (Jones)  Eydb. 

Arizona:  sandy  declivities  under  the  Echo  Cliffs  near  Limestone  Tanks,  about  13 
miles  south  of  Navajo  Bridge,  alt.  5200  ft.,  7  June,  fr.  Eipley  and  Barneby  4S68. 
Also  observed,  but  not  collected,  under  the  same  cliffs  11  miles  north  of  Cedar  Kidge. 

An  addition  to  the  flora  of  Arizona.  The  specimens  are  slightly  abnormal 
in  having  the  peduncles  much  shorter  than  is  usual  in  material  from  Zion 
and  the  Virgin  Valley,  while  the  whole  plant  is  more  copiously  pubescent. 

Astragalus  panguicensis  M.  E.  Jones 

Astragalus  panguicensis  (M.  E.  Jones),  M.  E.  Jones,  Contrib.  W.  Bot.  10:62,  tab.  4.  1902. 
A  Chamaeleuce  var.  panguicensis  M.  E.  Jones,  Proc.  Calif.  Acad.  Sci.  (II),  5:671.  1895. 
A.  argophyllus  var.  panguicensis  M.  E.  Jones,  Contrib.  W.  Bot.  8:5.  1898;  Eev.  Astrag. 
207,  tab.  47.  1923.  Xylophacos  vespertinus  Eydb.,  N.  Amer.  Fl.  24:299.  1929,  pro  parte, 
quoad  syn.  cit.,  non  A.  vespertinus  Sheld. 

Utah:  gravelly  benches  in  yellow  pine  forest  west  of  Alton,  Kane  Co.,  alt.  6800  ft., 
5  June,  fl.  and  fr.  Ripley  and  Barnehy  4822. 

It  is  not  without  sqme  misgiving  that  the  collection  is  referred  to  this 
obscure  species,  for  the  type,  Jones  6023  f,  and  only  specimen  of  A.  Chamae- 
leuce var.  panguicensis  at  Pomona  is  a  very  mature  and  imperfect  plant  with- 
out either  flower  or  fruit.  In  an  envelope  attached  to  the  sheet  there  is  a  single 
pod  marked  in  Jones'  hand  "probably  the  type  of  A.  'panguicensis,"  but  this 
creates  a  further  difficulty,  for  the  pod  does  not  agree  as  closely  with  the 
figures  or  description  published  by  Jones  as  do  the  fruits  of  our  specimens, 
and  it  may  never  have  belonged  there.  The  plant  itself  differs  from  ours  in 
being  evidently  caulescent  and  in  having,  for  the  most  part,  broader  and  more 
obtuse  leaflets.  Manj^  species  of  this  group,  however,  even  if  quite  stemless  at 
first,  tend  to  become  caulescent  late  in  the  season,  and  we  have  one  specimen 
which  in  shape  of  leaflets  closely  matches  the  type.  In  both  collections  the 
stipules  and  the  characteristically  dense,  silvery,  subappressed  pubescence 
are  identical,  and  it  should  be  remembered  that  the  type  locality,  Panguitch 
Lake,  is  in  the  same  massif  as  our  station  and  not  more  than  twenty  miles 
distant.  Certainly  our  plant  can  be  referred  to  no  other  described  species. 

If  our  determination  is  accepted  as  correct  it  becomes  evident  that  A.  pan- 
guicensis has  been  universally  misunderstood.  Jones  first  described  it  as  a 
variety  of  A.  Chamaeleuce  A.  Gray,  three  years  later  transferred  his  variety 
to  A.  argophyllus  Nutt.,  then  raised  it  to  specific  rank,  and  finally,  in  the  Re- 
vision of  Astragalus,  returned  it  as  a  variety  to  A.  argophyllus.  Rydberg, 
evidently  without  first-hand  knowledge  of  the  plant,  reduced  it  to  the  very 
different  A.  vespertinus  Sheld.  Actually  the  small,  pale  purple  or  ochro- 
leucous  flowers  and  the  quality  of  the  pubescence  suggest  an  affinity  with 
A.  castaneiformis  S.  Wats.,  from  which  it  is  immediately  distinguished  by 
the  curved,  lanceolate  and  much  longer  pod. 


150 


CALIFOBKIA  ACADEMY  OF  SCIEXCES        [Proc.  4th  ser. 


Astragalus  uncialis  Barneby 

Nevada:  Currant  (Callaway),  Nye  Co.,  12  May,  flor.  Bipley  and  Barneby  4421,  topo- 
type.  Also  20  miles  to  the  west,  in  the  foothills  of  the  Pancake  Eange  near  Lockes. 
No.  4423. 

A.  uncialh  was  described  from  fruiting  material  and  the  fresh  flower  was 
unknown.  As  surmised  by  the  writer,  the  corolla  is  of  a  brilliant  purple,  simi- 
lar to  that  found  in  A.  utahensis  (Torr.)  T.  &  G.  though  perhaps  of  a  some- 
what more  violet  tinge. 

Astragalus  marianus  (Rydb.)  Barneby  comb.  nov. 

Xylophacos  marianus  Eydb.,  Bull.  Torr.  CI.  52:233.  1925;  N.  Amer.  Fl.  24:301.  1929.  A. 
Purshii  X  Newberryi  M.  E.  Jones,  Eev.  Astrag.  216.  1923. 

Nevada:  frequent  on  dry  benches  under  pines  and  aspens  along  Snake  Creek,  near 
Treasure  Lake,  Snake  Range,  White  Pine  Co.,  alt.  ca.  8200  ft.,  July,  fr.  Ripley  and\ 
Barneby  4036. 

This  rather  well-marked  species  has  been  reported  only  from  the  mountains 

of  southern  and  central  Utah,  where  it  is  of  not  uncommon  occurrence  inj 

sagebrush  valleys  between  six  and  eight  thousand  feet.  The  plants  from! 

Nevada  are  not  identical  with  the  type,  but  have  the  broader  leaflets  of  I 

Fremont  406  (NY)  from  Utah  Lake  referred  to  X.  marianus  by  Rydbergl 

{I.  c,  1925),  and  without  doubt  represent  a  shade  form  of  the  species.  Another] 

gathering  from  the  state,  our  no.  3515  from  talus  slopes  beneath  limestone! 

cliffs  in  the  canyon  of  the  Muddy  River  near  Caliente,  Lincoln  County,  atj 

4500  feet  may  also  belong  here,  but  the  determination  is  not  certain.  The] 

specimens  resemble  A.  marianus  closely  in  habit,  in  the  slender,  often  well-j 

developed  caudices  and  numerous,  small,  appressed-sericeous  leaflets,  but  inj 

the  dense  floccose  pubescence  of  the  pods  they  recall  A.  Newherryi  A.  GrayJ 

or  even  A.  Blyae  (Rose)  Tidestr.  The  locality  is  well  below  the  normal  alti- 

tudinal  limit  of  A.  marianus  and  it  may  be  that  the  plants  had  seeded  down! 

from  a  higher  level  of  the  Highland  Range  into  a  soil  where  lack  of  humidityj 

and  greater  temperature  might  account  for  the  increased  density  of  vesture.! 

Further  investigation,  particularly  of  the  flowers,  may  reveal  an  undescribedj 

species. 

Astragalus  sabinarum  (Rydb.)  Barneby  comb.  nov. 

Batidophaca  sabinarum  Eydb.,  N.  Amer.  Fl.  24:320. 1929. 

Only  the  type  specimen  from  Iron  County,  Utah  {Garrett  B2660,  NY),  hasi 
been  seen  by  the  writer,  but  this  clearlj^  represents  a  good  species.  The  com-j 
bination  in  Astragahts  is  proposed  in  order  to  facilitate  comparison  with  its] 
nearest  ally,  A.  niusimnnitm  nob.,  described  below. 

Astragalus  musimonum  Barneby  sp.  nov. 
(Plate,  17,  figs.  1-9) 

Astrar/ahis  mv.'timninim  Barnobv  sp.  nov.  ex  affinitate  A.  sabinarnm  (Eydb.)  nob.  a  quo! 
pendunculis  validis  elongatis  folia  multo  supcrantibus,  racemis  pluritloris,  dentibusj 
calycinis  abbreviatis  hand  nigro-eiliatis,  et  praesertim  legumine  parcius  strigoso  ad 


VOL.XXV]     BABNEBY:  PUGILLUS  ASIEAGALOEVM  ALTER  15] 

apicem  abrupte  hamato-recurvo  facillime  diagnoscenda.  Habitu  A.  amphioxyn  A.  Gray 
affinesque  nonniillas  species  aliquantulum  refert,  sed  calyce  campanulato  cariiiaque  lata 
ab  Argophyllis  (sive  Xylophacis  Eydb.)  omnibus  graviter  diserepat. 

Herba  perennis  depressa  caespitosa  breviter  caulescens,  pilis  argenteis  adscendentibus 
basifixis  imprimis  arete  appressis  paucisque  patulis  commixtis  undique  strigoso-cana: 
caulibus  compluribus  prostratis  saepe  flexuosis  incanis  e  coUo  radieis  lignosae  verticalis 
emissis,  2-7  cm.  longis,  fere  ex  ipsa  basi  racemigeris :  stipulis  priminn  iml)ricatis  demum  in- 
ternodio  3-9  mm.  longo  separatis,  laneeolatis  vel  ovato-lanceolatis  acutis,  3-5  mm.  longis, 
praeter  nervum  medianum  viridem  omnino  hyalinis  vel  superne  herbaceis,  mox  scariosis, 
inferne  glabratis  intus  glabris  longe  ciliatis  ad  basin  petiolo  breviter  adnatis :  f oliis  humi 
appressis  4-10  cm.  longis,  petiolo  gracili  elongato  leviter  canaliculato  rachin  saepissime 
superanti:  foliolis  5-9-jugis  parvis  remotiusculis  inferioribus  plerumque  alternis  articulatis 
obovatis  ellipticis  oblanceolatisve,  apice  rotundatis  acutis  vel  apiculatis  3-7  mm.  longis, 
2-3  mm.  latis,  secus  racliin  sursum  decrescentibus :  pedunculis  numerosis,  praecocioribus 
subradicalibus,  prostratis  validis  striatis  parcius  strigosis  viridulis,  iam  ad  anthesin  folia 
superantibus  fructiferis  elongatis:  racemis  brevibus  7-12-floris  primum  subcapitatis  ca. 
1.5  cm.  longis,  mox  laxioribus,  demum  4-6  cm.  longis :  bracteis  ovato-acuminatis  ea.  3  mm. 
longis  hyalinis  ciliatis:  pedicellis  firmulis,  vix  1  mm.  longis,  adscendentibus:  calycis  pur- 
pureo-tincti  extus  strigoso-cani  tubo  campanulato  postice  valde  gibbo  3.5  mm.  longo,  2  mm. 
lato,  legumine  accreseenti  nunquam  rupto,  dentibus  subulatis  acutis  1.5  mm.  longis:  petalis 
laete  violaeeo-purpureis,  vexillo  macula  magna  elliptica  alba  superne  pulchre  pluristriata 
instructo:  vexillo  12-14  mm.  longo,  supra  unguiculum  brevem  et  angustum  in  limbum 
plicatum  late  oblongum  4  mm.  latum  obtusum  apice  emarginatum  abrupte  expanso,  infra 
medium  plus  minusve  retro-arcuato,  marginibus  reflexis:  alls  ca.  10  mm.  longis,  carinam 
pauUulum  superantibus,  lamina  anguste  lanceolato-kmata  ca.  1.5  mm.  lata,  auriculo  parvo 
reflexo  ineluso  7  mm.  longa  :  petalorum  carinalium  lamina  oblique  ovato-lunata  6  mm.  longa, 
saltern  2.5  mm.  lata,  marginibus  superioribus  leviter  concavis,  inferioribus  connatis  supra 
medium  in  apicem  porrectum  plerumque  obtusissimum  per  angulum  fere  rectum  sursum 
arcuatis:  legumine  patulo  sessili  uniloculari,  ambitu  anguste  elliptico,  falcato,  1.5-2.0  cm. 
longo,  utrinque  angustato  atque  compresso,  medium  versus  admodum  obcomresso  vel  saltern 
dorsaliter  sulcato,  in  rostrum  abrupte  hamato-reeurvum  stylo  persistent!  cuspidatum  sur- 
sum attenuato,  sutura  ventrali  per  longitudinem  totam  acuta  sensim  concava,  dorsali  inferne 
subrecta,  ad  medium  prominula  sed  ob  leguminis  obcompressionem  ibi  depressa,  ad  rostrum 
acutissima  abrupte  convexa,  sectione  transversali  anguste  obcordata  5-6  mm.  lata,  ca.  2 
mm.  alta,  valvulis  coriaceis  sparsiuscule  albostrigosis  transverse  striatis  immaculatis: 
seminibus  maturis  ignotis. 

Nevada:  calcareous  gravel  slopes  of  the  Sheep  Mts.,  near  the  entrance  to  Deadman 
Canyon,  Clark  Co.,  alt  5700  ft.,  5  May,  tl.  &  fr.  Ripley  and  Barnehy  3332.  Type  in_ 
Herb.  Calif.  Acad.  Sci.  No.  300411. 

A.  musimonum,  with  its  cespitose,  very  shortly  caulescent,  prostrate  stems, 
silvery-strigose  herbage  and  racemes  of  purple  flowers,  has  at  first  sight  the 
aspect  of  a  slender  form  of  A.  amphioxys  A.  Gray,  but  the  companulate  calyx- 
tube,  broad  keel  and  thin-walled  pod  indicate  a  relationship  with  that  group 
of  Astragali  segregated  by  Kydberg  as  the  genus  Batidophaca,  and  among 
those  species  it  finds  a  natural  place  beside  A.  sahinariini  in  the  series  Villosae 
(cf.  Rj'db.,  I.  c).  From  the  latter  it  differs  in  the  short  calyx-lobes,  more 
numerous  flowers  and  more  copious  pubescence  of  straight,  appressed  hairs, 
while  the  mature  fruits,  laxly  racemose  on  the  stout,  elongate  and  at  length 
reclining   peduncles,   are   comparatively   glabrate,   attenuate   at  base   and 


152  CALIFORXIA   ylCADEMY   OF  SCIENCES        [Proc.  4th  ser. 

abruptly  hooked  at  apex,  in  contrast  with  the  canescent,  regularly  lunate  pod 
of  A.  sahinarum. 

A.  musimomim  has  been  known  in  the  type  locality  on  the  western  slope 
of  the  Sheep  ]\Iountains  for  several  years,  but  adequate  fruiting  material  has 
only  recently  been  obtained.  In  this  station  it  is  local  but  abundant  on  cal- 
careous benches  in  the  Upper  Sonoran  Life-zone  and  seems  to  belong  to  the 
calciphile  foothill  flora  of  which,  in  southern  Nevada,  Actinea  Cooperi  (A. 
Gray),  0.  Kze.,  Senecio  stygius  Greene  and  Astragalus  Tidest^^omii  (Rydb.) 
Clokey  are  characteristic  elements.  Immature  specimens  from  northern  Ari- 
zona, found  growing  under  similar  conditions  at  5300  feet  in  Mokiak  Pass, 
Mohave  County  (No.  4321),  are  almost  certainly  to  be  referred  to  A.  musi- 
momim, but  in  absence  of  the  legume  the  determination  remains  in  doubt. 

The  proposed  trivial,  from  the  Latin  'musimon,'  alludes  to  that  magnificent 
race  of  the  Mountain  Sheep,  now  happily  preserved  from  extinction,  which 
is  native  to  the  Sheep  ]\Iountains,  and  from  which  the  name  of  the  range  itself 
is  certainly  derived. 

Astragfalus  pinonis  M.  E.  Jones 

Pisophaca  pinonis  (M.  E.  Jones)  Eydb. 

Nevada:  Highland  Eange  west  of  Caliente,  Lincoln  Co.,  alt.  5200  ft.,  20  May,  fl.  &  fr. 
Ripley  and  Barnrhy  34S7a.  Currant  Creek,  N.  E.  Nye  Co.,  alt.  6200  ft.  No.  3640. 

This  rare  species  has  been  known  hitherto  only  from  Beaver  (Frisco,  Jones, 
the  type  collection)  and  eastern  Juab  (Tintic,  Jones)  Counties,  Utah.  Our 
specimens  are  taller  and  better  developed  than  Jones'  evidently  depauperate 
type,  with  pods  up  to  2.5  cm.  in  length,  but  they  are  essentially  alike  in  struc- 
ture and  pubescence.  The  plants  were  found  only  as  isolated  individuals  and 
appeared  very  scarce. 

Astragalus  oophorus  S.  "Wats. 

Nevada:  Head  of  Lee  Canyon,  Charleston  Mts.,  Clark  Co.,  alt.  9100  ft.,  Ripley  and 
Barneby  4979. 

Through  the  kindness  of  Mr.  Ira  W.  Clokey  it  has  been  possible  to  compare 
.this  collection  with  Train  2141  from  the  same  locality  which  was  cited  in  a 
recent  paper  on  the  Astragali  of  the  Charlestons  (Clokey,  Madrono  6:  217. 
1942)  as  A.  artipes  A.  Gray.  Study  of  the  more  ample  material  now  available 
makes  it  clear  that  the  Charleston  plant  is  more  justly  referred  to  A.  oophorus, 
but  the  original  determination,  which  was  made  by  the  Bureau  of  Plant  In- 
dustry and  based  on  meager  and  somewhat  atypical  specimens,  is  readily 
understandable  when  it  is  realized  how  variable  and  how  closely  allied  the 
species  are,  and  how  incorrectly  they  have  been  interpreted  in  the  past. 

In  fact  there  seems  to  be  little  ground  on  which  A.  oophorus  and  A.  artipes 
can  be  separated  specifically,  for,  while  the  typical  phases  occupy  distinct, 
contiguous  areas,  there  occur,  particularly  toward  the  southern  limit  of  A. 
oophorus,  specimens  which  are  to  some  degree  intermediate.  A.  artipes  of 


Vol.  XXV]     BAENEBY:  PUGILLUS  ASIBAGALORUM  ALTER  153 

northern  Arizona,  middle  Utah  and  adjacent  Colorado  is  distinguished  by 
the  cream-colored  flowers  and  by  the  oblique  legume  with  nearly  straight 
ventral  and  strongly  convex  dorsal  sutures.  A.  uophorus,  on  the  other  hand, 
a  common  species  throughout  central  Nevada  which  extends  into  transmon- 
tane  California  along  Owens  Valley  to  the  Panamints,  has  flowers  with  pur- 
ple banner  and  keel,  and  a  regularly  ellipsoid  pod  with  either  suture  about 
equally  convex.  In  southern  Nevada  and  in  the  Panamints,  however,  purple- 
flowered  plants  are  sometimes  found  with  somewhat  oblique  legumes,  in  which 
case  the  only  absolutely  reliable  criterion  is  the  color  of  the  flowers.  This  raises 
a  serious  difficulty,  since  Watson,  in  the  original  publication,  described  the 
flowers  of  A.  oophorus  as  ochroleucous.  This  I  believe  to  have  been  an  error. 
Not  only  is  the  pigment  in  the  petals  of  A.  oophorus  extremely  fugacious  in 
the  herbarium,  but  among  the  numerous  collections  of  the  species  made  in 
Nevada  in  recent  times  not  one  with  ochroleucous  flowers  has  been  detected. 
I  have  myself  collected  the  species  to  the  north,  south  and  east  of  the  type- 
locality  (at  Austin,  in  the  Toyabe  Range  and  in  Montgomery  Pass),  yet 
among  thousands  of  individuals  none  was  observed  without  purple  keel  and 
banner.  Apart  from  the  value  of  the  flower  color  as  a  taxonomic  criterion,* 
the  matter  is  of  special  importance  in  so  far  as  it  has  affected  the  interpreta- 
tion of  A.  oophorus  by  Jones  and  Rydberg.  Evidently  it  was  a  factor  in  Jones' 
treatment  of  A.  artipes  from  middle  Utah  as  A.  oophorus  (and  simultaneously 
of  the  purple-flowered  A.  oophorus  as  A.  artipes),  while  it  also  prompted  Ryd- 
berg to  recognize  in  A.  oophorus  two  species  of  Phaca,  P.  oophora  and  P. 
jucunda,  separated  mainly  by  this  very  character. 

For  many  years  Jones  seems  to  have  considered  these  two  names  synony- 
mous, and  in  the  Index  to  the  Revision  of  Astragalus  he  actually  stated  that 
"A.  artipes  ...  is  oophorus."  Nevertheless  his  real  intentions  at  this  time 
seem  to  have  been  otherwise,  for  on  Plate  22  of  the  same  work  appears  a  figure 
under  the  name  "var.  artipes"  and  the  combination  is  repeated  in  the  Index 
to  Plates.  By  a  curious  misprint,  however,  it  was  not  validly  published.  On 
page  121  of  the  Revision,  immediately  following  a  detailed  description  of 
A.  oophorus  (in  which  the  pod  described  is  clearly  that  of  A.  artipes)  are 
printed  two  short  paragraphs,  both  under  the  heading  A.  oophorus  var. 
caulescens.  There  seems  to  be  no  doubt  that  the  first  line  of  the  upper  para- 
graph should  have  read  "A.  oophorus  var.  artipes  (Gray,  Proc.  Amer.  Acad. 
13.  370"  which  would  join  on  comprehensively  with  "(1878)  as  species)"  of 
the  second  line.  But  Jones'  confusion  went  deeper  than  a  mere  typographical 
lapsus,  for  the  plant  figured  as  var.  artipes  is  plainly  nothing  more  than 
A.  oophorus  in  the  strictest  sense,  while  the  true  A.  artipes  of  Gray  is  repre- 
sented by  the  A.  oophorus  var.  caulescens  of  the  second  paragraph  and  of  the 

*  An  exact  parallel  is  found  in  the  case  of  A.  Bedcwithii  T.  and  G.  and  its  vicariant  var. 
purpureus  M.  E.  Jones  (A.  ariemisiarum  M.  E.  Jones).  The  former,  with  ochroleucous 
floAvers,  so  common  in  western  Utah,  passes  westward,  almost  exactly  at  the  Nevada  line, 
into  a  form  with  purple  flowers. 


154  CALIFOBXIA  ACADEMY  OF  SCIENCES        [Proc.  4th  ser. 

corresponding  figure  on  Plate  22.  This  point  has  been  verified  by  study  of 
the  typical  material  of  A.  megacai'pus.  var.  caulescens  ]\I.  E.  Jones  (PO). 
It  should  also  be  noted  that  the  figures  of  what  Jones  regarded  as  typical 
A.  oophorus  on  Plate  21  contain  a  mixture :  true  A.  oopliorus  is  there  rep- 
resented only  by  the  upper  pod  and  section,  while  the  remainder  (marked 
"a,"  from  Mammoth,  Utah)  are  quite  characteristic  of  A.  artipes  and  in  no 
waj'  separable  from  the  figure  of  var.  caulescens  on  Plate  22.  A  touch  of  fan- 
tasj^  is  added  to  this  labyrinth  of  confusion  by  the  fact  that  the  material  of  A. 
artipes  gathered  by  Lemmon,  Rusby  and  Tourney  in  the  Flagstaff  region  of 
northern  Arizona  are  referred  by  Jones  (1.  c.  120)  to  A.  megacarpus  var. 
Parryi  A.  Gray,  although  he  noted  that  they  were  abnormal  in  having  ochro- 
leueous  flowers. 

Phaca  jucunda  Jeps.  and  Rydb.  has  since  publication  been  rightly  reduced 
by  Dr.  Jepson  himself  (Fl.  Calif.  2:  348.  1936)  to  A.  oophorus.  Apart  from 
the  criterion  of  color,  which  I  believe  to  be  chimerical,  Rydberg  advanced 
only  the  shape  of  leaflets  to  distinguish  the  segregate  P.  jucunda  from  P. 
oophora.  A  mere  glance  at  any  considerable  suite  of  specimens,  however, 
reveals  that  the  shape  of  leaflets  is  in  no  way  correlated  with  flower-color, 
with  shape  of  pods  or  with  distribution,  for  they  may  vary  throughout  the 
range  of  the  species  from  crowded,  suborbicular  and  refuse  through  many 
intermediate  stages  to  remote,  narrowly  elliptic  and  acute  (as  in  the  Charles- 
ton material)  and  exactly  similar  variation  occurs  in  A.  artipes.  Considerable 
variation  occurs  also  in  the  length  of  the  stipe  and  in  the  size  of  the  corolla. 

I  am  disposed,  therefore,  to  regard  the  Avhole  complex  as  a  single  species, 
of  wlych  A.  artipes  may  be  regarded  as  a  well-marked  variety,  under  the  com- 
bination A.  oophorus  var.  caidescens  Jones.  The  pertinent  literature  is  sum- 
marized in  the  following  synonymy. 

Astragalus  oophorus  S.  "Wats.  var.  typicus  Barneby  nom.  nov. 

A.  oophorus  S.  Wats.,  Bot.  King  73.  1871.  A.  oophorus  sensu  M.  E.  Jones,  Rev.  Astrag. 
120,  pro  parte,  PI.  21  (as  to  upper  figures).  A.  oophorus  var.  caulescens  M.  E.  Jones, 
op.  cif.  121  lin.  32  (lapsu),  non  ibid.  lin.  38.  A.  oophorus  vnr.  artipes  M.  E.  Jones,  op. 
cit.,  Index  to  Plates  and  PI.  23,  nomen  nudum,  non  A.  artipes  A.  Gray.  Phaca  oophora 
(A.  Gray)  Rydb.,  N.  Amer.  Fl.  24:338.  1929.  P.  jucunda  Jeps.  and  Rydb.,  in  Rydb., 
op.  cit.  339.  Astragalus  jucundus  Jeps.  and  Rydb.  ex  Peck,  Man.  PI.  Oreg.  446.  1940, 
nomen  midum. 

Astragalus  oophorus  S.  Wats.  var.  caulescens  (M.  E.  Jones)  M.  E.  Jones 

A.  oophorus  S.  Wats.  var.  caulescens  (M.  E.  Jones)  M.  E.  Jones,  Rev.  Astrag.  121,  lin.  38, 
PI.  22,  1923,  non  ibid.,  lin.  31.  A.  megacarpus  var.  caulescens  M.  E.  Jones,  Proo.  Calif. 
Acad.  Sci.  II,  5:043.  1895.  A.  artipes  A.  Gray,  Proc.  Amer.  Acad.  13:370.  1878.  A. 
oophorus  sensu  M.  E.  Jones,  Rev.  Astrag.  120,  pro  ma.r.  parte,  PI.  21,  fig.  a,  non  S.  Wats. 
riifira  arlipes  (A.  Gray)  Rydb.,  Bull.  Toir.  CI.  32:(i()4.  190G;  N.  Amer.  Fl.  I.e. 


VOL.XXV]     BABNEBY:  PUGILLUS  ASTBAGALOEUM  A  LT  E  B  155 

Astragalus  straturensis  M.  E.  Jones 

Atelophragma  straturense  (M.  E.  Jones)  Rydb.  Hamosa  atratiforniis  Rydb. 

Nevada:  dry  slopes  among  junipers  and  pinyons,  12  miles  east  of  Panaca,  Lincoln 
Co.,  alt.  7050  ft.,  9  May,  flor.  Bipley  and  Barneby  4386.  Also  ibid.,  11  June,  fruct. 
No.  4971. 

A  new  record  for  Nevada.  The  flowers  are  bright  violet-blue  when  fresh, 
with  a  pale,  striate  diamond  in  the  fold  of  the  banner. 

This  will  almost  certainly  prove  to  be  the  same  plant  as  that  collected  by 
Vernon  Bailey  {Death  Valley  Exped.  No.  1975)  "about  30  km.  east  of  Panaca" 
in  Utah  and  determined  by  Sheldon  (ap.  Gov.,  Contrib.  U.  S.  Nat.  Herb.  4  :  87. 
1893)  as  A.  ohscurus  S.  Wats.  The  latter  species  is  not  known  south  of  the 
valley  of  the  Humboldt  River  and  has  ochroleucous,  not  "purplish,"  flowers. 

Astragalus  eremiticus  Sheld.  var.  typicus  Barneby  nom.  no  v. 

A.  eremiticus  Sheld.,  Minn.  Bot.  Stud.  1:161.  1896,  sensu  strict o. 

Utah:  Gunlock,  Washington  Co.,  with  ochroleucous  flowers.  BipJey  and  Barneby  4298. 
Arizona:  Mokiak  Pass,  Mohave  Co.,  with  purple  flowers.  No.  4313.  Nevada:  Pah- 
ranagat  Mts.,  west  of  Crystal  Springs,  Lincoln  Co.,  flowers  purjile.  No.  4408. 

Astragalus  eremiticus  Sheld.  var.  spencianus  M.  E.  Jones, 
Contrib.  W.  Bot.  10  :  60. 1902 
A.  boiseanus  A.  Nels.,  Bot.  Gaz.  53:223.  1912.  A.  arrectus  var.  Kelseyi  M.  E.  Jones,  Eev. 
Astrag.  161.  1923,  non  A.  Kelseyi  Rydb.  Tium  eremiticum  (Sheld.)  Rydb.,  N.  Amer.  Fl. 
24:390.  1929;  Bull.  Torr.  CI.  57:400.  1930,  pro  parte. 

Nevada:  10  miles  west  of  Elko,  Elko  Co.,  in  sagebrush.  Bipley  and  Barneby  4588. 

The  type  of  this  variety,  though  not  specified  by  Jones,  is  evidently  his 
collection  from  Spencemont  ( ?  Lander  County),  Nevada,  in  1891  (PO),  and 
is  exactly'  similar  to  authentic  material  of  A.  hoisea^ius  from  the  neighborhood 
of  Boise,  Idaho.  Later  Jones  referred  it  as  a  variety  to  A.  arrectus  A.  Gray,  but 
mistakenly  identified  it  with  A.  Kelseyi  Rydb.  which  is  A.  atropuhescens 
Coult.  and  Fish,  and  not  certainly  distinct  from  A.  arrectus.  Rydberg,  in  his 
review  of  Tium  (Z.  c,  1930),  recognized  the  differences  which  exist  between 
typical  A.  eremiticus  and  the  var.  spencianus,  but  was  unwilling  to  separate 
them.  Nonetheless  the  morphological  criteria  enumerated  by  him,  when  con- 
sidered in  the  light  of  factors  of  distribution  and  habitat,  appear  amply  suf- 
ficient to  support  a  variety.  The  var.  typicus  is  endemic  to  the  arid,  subdesertic 
ranges  about  the  common  boundary-point  of  Nevada,  Arizona  and  Utah, 
alwaj's  in  the  drainage  of  the  Colorado  River  or  its  immediate  tributaries 
such  as  the  Muddy  River  or  the  Virgin,  where  it  occurs  on  dry,  stony  slopes 
in  the  juniper-pinyon  association,  sometimes  in  depauperate  sagebrush.  The 
var.  spencianus,  on  the  other  hand,  occupies  the  interior  basin  region  of  north- 
eastern Nevada  and  adjacent  Idaho  and  Oregon,  in  the  drainage  of  the 
Owyhee,  Snake  and  Humboldt  rivers,  where  it  is  a  characteristic  element  of 
the  flora  of  sagebrush  plains  and  valleys.  It  is  not  known  to  extend  south  of 
White  Pine  County,  Nevada,  and  no  intergrades  have  been  seen. 


156  CALIFOBNIA  ACADEMY  OF  SCIEXCES        [Proc.  4th  ser. 

Astragalus  Bryantii  Barneby  spec,  no  v. 
(Plate  17,  figs.  10-18) 

Astra ffaliis  Bryantii  Barneby  spec.  nov.  inter  Palantia  generis  Tii  Medic,  sensi.  Rydb. 
adnumeranda,  habitu  pube  etc.  A.  mol-iacensi  A.  Gray  forsan  proxinie  affinis,  sed  ab 
eo  consociisque  omnibus  leguminis  angnstioris  valde  compressi  sutura  ventrali  acuta 
necnon  pericarpio  tenui  diversissima  et  his  notis  Havinsae  Tricarinatas  Eydb.,  a  quibus 
imprimis  tubo  calycino  tubuloso  nee  campanulato  longius  distat,  simulans. 

Herba  verosimiliter  pcrennis,  'tripedalis'  teste  cl.  Bryant,  pube  e  pilis  brevibus  adscenden- 
tibus  candidis  constituta  praeter  foliolorum  paginam  superiorem  glabratam  undique  stri- 
gosa,  partibus  junioribus  villosulo-eanescentibus :  caulibus  ut  videtur  adscendentibus  (sed 
imis  ignotis)  plus  minusve  flexuosis  striatis  purpuraseentibus:  stipulis  deltoidco-acuminatis 
extus  strigoais  herbaceis  vcl  anguste  scarioso-niarginatis  ca.  5  mm.  longis,  mox  reflexis: 
foliis  patulis  subsossilibus  vel  breviter  petiolatis,  8-12  cm.  longis,  racM  subtereti  strigoso 
petiolum  multoties  superanti:  foliolis  14-20,  petiolulo  0.5-0.75  mm.  longo  gestis.  ovatis, 
late  ellipticis  vel  obovatis,  5-14  mm.  longis,  3-8  mm.  latis,  inferne  strigoso-eanescentibus: 
pedunculis  firmis  arcuato-adscendentibus  striatis  4-5  cm.  longis,  in  racemum  laxum  ca. 
15-florum  fructiferum  vix  elongatum  nee  folio  suffulcranti  multo  longiorem  abeuntibus: 
bracteis  lanceolato-attenuatis  ca.  2  mm.  longis:  floribus  violaceis  15-17  mm.  longis  pedicello 
arcuato  1.0-2.5  mm.  longo  saepissime  atropiloso  suffultis:  ealycis  membranacei  pilis  albis 
nigrisque  commixtis  strigosi  tubo  late  cylindrico,  5  mm.  longo,  postice  gibbo,  ovario  in- 
tumescenti  mox  rupto,  dentibus  anguste  subulatis  acutis  1.5-1.75  mm.  longis  coronato: 
vexillo  obovato  16  mm.  longo,  ca.  8  mm.  lato,  ad  medium  retrorsus  arcuato  sed  apice  pro- 
funde  emarginato  declinato,  marginibus  reflexis:  alls  ca.  14  mm.  longis,  lamina  lunata 
obtusa  1.5-1.75  mm.  lata,  auriculo  basali  conspicuo  incluso  7  mm.  longa :  carinae  petalis 
ca.  13  mm.  longis  laminis  latius  lunatis  saltem  7  mm.  longis,  2.5  mm.  latis,  marginibus 
superioribus  profundiuscule  concavis,  inferioribus  (connatis)  inferne  reetis  ad  medium 
abrupte  in  apicem  latum  obtusissimum  sursum  arcuatis:  legumine  horizontaliter  patulo  vel 
leviter  adscendenti,  arete  sessili,  2-loculari,  subreeto  vel  saepius  paullum  arcuato,  ambitu 
lineari  vel  lineari-lanceolato,  2.0-2.5  (3)  cm.  longo,  ad  basin  rotundato  supra  medium 
sensim  attenuate,  valde  compresso,  sulco  0.75-1.5  mm.  alto  impresso  dorsaliter  peicurso, 
sutura  ventrali  per  longitudinem  totam  proniinula,  dorsali  ad  ventralem  usque  introflexa 
septum  subcompletum  2.0-2.5  mm.  altum  fere  in  leguminis  apicem  productum  efformanti, 
valvulis  chartaceis  tenuiter  reticulatis  parce  strigulosis  vel  demum  glabratis  nonnumquam 
minute  purpureo-guttulatis,  sectione  transversali  anguste  deltoideo-obcordata,  3.0-4.5  mm. 
alta,  1.5-3.0  mm.  lata:  seminibus  oblique  reniformibus,  2  mm.  latis,  brunneis.  M 

Arizona:  at  the  head  of  Phantom  Canyon  in  the  Grand  Canyon  of  the  Colorado 
River,  Coconino  Co.,  15  Dec.  1939.  Collected  by  Dr.  11.  C.  Bryant  of  the  National 
Park  Service  in  whose  honor  it  is  named. 

Type  in  Herb.  Calif.  Acad.  Sci.  No.  293940.  Also  collected  in  sand  at  the  mouth  of 
Hermit  Creek  in  the  Grand  Canyon,  10  April  1917.  Eastwood  5991  (G,  F). 

The  type  of  A.  Bryantii  is  apparently  part  of  the  collection  referred  to  by 
Kearney  and  Peebles  (Fl.  PI.  Ariz.  486. 1942)  as  representing  an  undescribed 
species  allied  to  A.  palans  M.  E.  Jones,  but,  although  a  real  affinity  may  exist 
between  the  two,  it  has  been  found  convenient  for  purposes  of  diagnosis  to 
compare  it  rather  with  A.  luokiacotsis  A.  Gray,  with  which  it  agrees  more 
closely  in  the  quality  of  the  pubescence,  in  the  violet,  not  reddish  purple 
coloration  of  the  corolhi,  and  in  the  less  marked  curvature  of  the  legume.  From 
A.  mokiacensis,  known  only  from  a  few  localities  on  either  side  of  the  Colo- 


VoL.XXV]     BARNEBT:  PUGILLUS  ASTBAGALOEUM  ALTER  157 

rado  River  immediately  below  the  Grand  Canyon,  A.  Bryaniii  differs  in  the 
strongly  compressed,  more  slender  and  sparingly  strigose  pods  which  are  of 
thinner  texture  and  spread  horizontally  from  the  axis  of  the  raceme.  Indeed 
the  texture  and  compression  of  the  pod,  with  its  acute  ventral  suture,  led  me 
at  first  to  regard  the  species  as  more  intimatelj^  connected  with  forms  of  A. 
arrectus  A.  Gray  {Tiuni  ser.  Arrecta  Rydb.)  or  with  Hamosa  ser.  Tricarinatae 
Rydb.,  but  the  complete  absence  of  stipe  and  the  cylindric  calyx-tube  make 
such  an  alignment  improbable,  and  the  species  is  more  naturally  associated 
with  A.  mokiacensis. 

As  far  as  can  be  seen  from  the  limited  material  at  hand,  A.  Bryaniii  is  very 
constant  in  vegetative  characters,  but  the  shape  of  the  pod  is  evidently 
variable.  Normally,  as  always  in  Eastwood  5991,  the  pod  is  narrowly  lance- 
attenuate  in  outline,  the  cross-section  at  the  middle  being  about  3  mm.  high. 
Attached  to  Dr.  Brj^ant's  specimen,  however,  there  are  several  loose  pods, 
essentially  similar  in  structure  and  pubescence  but  half  as  broad  again,  the 
cross-section  being  4.5  mm.  high  and  the  outline  lanceolate,  more  abruptly 
acute  rather  than  attenuate  at  apex.  Although  the  plant  from  which  these 
pods  have  fallen  has  not  been  studied,  their  measurements  are  included  in 
the  description  of  the  species  and  they  have  been  treated  as  an  integral  part 
of  the  type.  A  similar  latitude  of  variation  has  been  found  by  the  writer  to 
occur  in  a  number  of  species  in  the  genus. 

Astragalus  Pattersonii  A.  Gray,  sens,  strict. 

Jonesiella  Pattersonii  Eydb. 

Arizona:  red  clay  hills  2  miles  east  of  Fredonia,  Coconino  Co.,  alt.  4900  ft.  Ripley  and 
Barneiy  4361. 

The  specimens  are  almost  exactly  typical,  with  narrowly  elliptic,  subsessile, 
strictly  erect  pods.  This  form  of  the  extremely  polymorphic  species  was  known 
to  Rydberg  only  from  Colorado,  and  has  not  been  reliably  reported  outside 
of  that  state,  though  a  closely  allied  form  with  spreading  pods  is  common  in 
the  Navajo  Basin.  The  A.  Pattersonii  of  Tidestrom  and  Kittell  (Fl.  Ariz,  and 
N.  Mex.  214.  1941),  if  one  may  judge  from  the  synonymy,  is  an  aggregate 
containing  a  number  of  diverse  forms. 

Astragalus  ensiformis  M.  E.  Jones  var.  typicus  Barneby  nom.  nov. 

A.  ensiformis  M.  E.  Jones,  Rev.  Astrag.  226.  1923.  A.  ursinus  M.  E.  Jones,  Proc.  Calif. 
Acad.  Sci.  II,  5:658.  1895,  non  A.  Gray.  Hamosa  ensiformis  Eydb. 

Arizona:  gentle  stony  slopes  among  junipers  in  Mokiak  Pass,  north  of  Wolf  Hole, 
Mohave  Co.,  alt.  5300-5500  ft.,  4  May,  fl.  and  fr.  Ripley  and  Barneby  4311. 

The  type-locality  of  A.  ensiformis,  "4  miles  above  Pagumpa,"  is  very  close 
to  our  station  and  the  species  has  been  collected  elsewhere  only  by  Peebles  and 
Fulton  in  northern  Navajo  County  (cf.  Kearney  and  Peebles,  Fl.  PI.  Ariz. 
488.  1942).  The  name  is  universally  cited  as  having  been  published  in  1895, 
but  this  is  incorrect,  Jones,  it  is  true,  described  the  species  quite  minutely 


158  CALIFORNIA  ACADEMY  OF  SCIENCES        [Proc.  4th  ser. 

in  that  year  (I.  c),  but  under  the  name  A.  ursinus  A.  Gray,  merely  suggesting 
A.  ensiformis  as  an  alternative  should  the  plant  prove  as  distinct  as  he  sus- 
pected, and  the  combination  was  not  validly  applied  until  the  Revision  of 
Astragalus  in  1923.  Kydberg,  who  did  not  see  the  flowers,  associated  A.  ensi- 
formis with  A.  Congdonii  S.  Wats,  and  A.  drepanololus  A.  Gray,  whereas, 
with  its  tubular  calyx  and  enlarged,  scarious  stipules,  it  clearly  belongs  to 
his  series  Malacae  of  Hamosa  and  is  closely  related  to  A.  Minthorniae  (Rydb.) 
Jeps.  In  fact  a  collection  from  southern  Utah,  described  below  as  a  new 
variety,  forms  a  passage  between  the  two. 

Astragalus  ensiformis  M.  E.  Jones  var.  gracilior  P.arneby  var.  nov. 

(Plate  17,  fig.  28) 
Astragalus  ensiformis  M.  E.  Jones  var.  gracilior  Barneby  var.  nov.  a  var.  typico  nob.  caule 
internodiis  extensis  duplo  elatiori,  floribus  post  anthesin  baud  deflexis,  racemo  fruetifero 
minime  secundo  et  praesertim  legumine  graciliori  minus  compresso  patule  erecto  eviaen- 
tius  hirsiitulo  noc  appresse  striguloso  diversa.  Ab  aflfini  A.  Minthorniae  (Rydb.)  Jcps., 
cujus  liabitum  in  memoriam  nonnihil  ledigit,  imprimis  legumine  magis  arcuato  baud 
dense  sericeo-villoso  facile  separanda. 

Utah:  in  stiff  soil  among  sagebrusb,  5  miles  south  of  Veyo,  Washington  Co.,  iilt. 
4900  ft.  Ripley  and  Barneby  4951.  Type  in  Herb.  Calif.  Acad.  Sci.  No.  300413. 

So  different  was  the  superficial  appearance  of  this  variety  from  that  of 
A.  ensiformis  var.  typicus  that  it  was  thought  in  the  field  to  represent  an 
undescribed  species,  but  subsequent  study  has  shown  that  the  differences, 
though  striking,  are  not  fundamental,  and  it  cannot  be  specifically  separated. 
In  the  typical  form  the  decumbent  stem,  with  congested  internodes,  does  not 
exceed  15  cm.  in  length,  the  fruiting  raceme  is  strongly  secund,  the  fiexuous 
pedicels  are  arcuate-reflexed  and  the  sparingly  strigose  pods,  though  arched 
upwards,  are  essentially  pendulous.  The  stems  of  the  var.  gracilior,  on  the 
other  hand,  are  upright  and  slender,  reaching  a  height  of  3  dm.,  the  raceme 
is  not  at  all  secund,  while  the  more  finely  and  densely  pubescent  pods  are 
held  erect  or  somewhat  spreading  on  straight  and  rigid  pedicels.  There  is 
no  appreciable  difference  in  the  form  of  the  flowers,  though  in  the  var.  iypicus 
they  are  reflexed  after  fertilization. 

Astragalus  chamaemeniscus  Barneby 

Nevada:  in  sand  under  sagebrush  near  the  Geyser  Ranch,  N.  Lincoln  Co.,  alt.  0050  ft., 
11  May,  flor.  Ripley  and  Barneby  4416. 

This  species  was  known  previously  only  from  extreme  northeast  Nye  and 
adjacent  White  Pine  counties  in  the  valley  of  the  White  River. 

Astragalus  Arthu-Schottii  A.  Gray,  sensu  restricto 

A.  ArtJiu-Seliottii  A.  Gray,  Proc.  Amer.  Acad.  G:209.  1863,  pro  max.  parte,  noii  op.  cit.  7: 
337.  1867.  A.  Coultcri  sensu  S.  Wats.,  Bot.  King,  66,  435.  1871;  A.  Gray,  Bot.  Calif. 
1:146.  1880;  Sheld.,  Minn.  Bot.  Stud.  1:140.  1894  et  auct.  mult,  pro  parte,  quoad  syn., 
non  Bth.  A.  lenfigiriosus  var.  borreganus  M.  E.  Jones,  Contrib.  W.  Bot.  8:3.  1898; 
Rev.  Astrag.  126,  tab.  25.  1923,  A.  lentiginosiis  var.  Coulteri  sensu  Jones,  Rev.  Astrag. 


Vol.  XXV]     BARNEBY:  PUGILLUS  ASTEAGALOBUM  ALTER  159 

127.  quoad  syn. ;  Jeps.,  F1.  Calif.  2:357.  1936,  quoad  syn.  et  pi.  cit.  Schott.  A.  agninus 
Jeps.,  Mau.  Calif.  577.  1925;  Fl.  Calif.  2:355.  1936;  Harrison  and  Kearney,  Journ. 
Wash.  Acad.  Sci.  22:227.  1932;  Munz,  Man.  S.  Calif.  Bot.  270.  1935;  Kearney  and 
Peebles,  Fl.  PI.  Ariz.  487.  1942.  Cysiium  Arthu-Schottii  (A.  Gray)  Rydb.,  N.  Amer. 
Fl.  24:407.  1923.  C.  agninum  (Jeps.)  Rydb.,  loc.  cit.  408. 

Study  of  the  somewhat  elaborate  synonymy  and  literature  cited  will  show 
that  4-  Arthu-Schottii  is  believed  to  be  the  oldest  valid  name  for  the  plant 
which  has  been  described  and  treated  in  comparatively  recent  years  as  A. 
lentiginosus  var.  horreganus  Jones  and  A.  agninus  Jeps.,  and,  while  reasons 
might  be  advanced  for  discarding  Dr.  Gray's  name  as  a  permanent  source  of 
error  and  confusion,  a  reexamination  of  the  facts  tends  to  support  its  revival 
into  current  use.  A.  Arthu-Schottii  was  first  encountered  by  Schott  during 
the  exj^loration  of  the  Mexican  boundary  in  1855,  but  his  collections  were 
not  cited  by  Gray  in  the  Botany  of  the  Survey,  doubtless  because  the  speci- 
mens available  to  him  were  without  fruit.  Several  years  later,  having  received 
from  Fremont  additional  material  of  what  he  took  to  be  the  same.  Gray  de- 
cided to  publish  the  species,  including  in  his  concept  a  plant  collected  in 
northern  Sonora  by  Capt.  Smith.  Thus  the  original  A.  Arthu-Schottii  was 
founded  on  three  elements,  cited  by  Gray  (I.  c.)  in  the  following  words:  "on 
the  ]\Iohave  River,  Fremont,  23  April.  Diluvial  banks  of  the  Colorado  near 
its  mouth,  12  March,  Schott.  Boca  Grande,  Capt.  E.  K.  Smith.",  and  each  of 
which  belonged  to  a  different  species.  The  Fremont  specimen  (NY)  is  a  very 
depauperate,  first-year  individual  of  that  form  of  A.  Coulteri  Bth.  with  small 
corollas  which  is  common  on  the  Mohave  Desert  of  California  and  which  is 
variously  treated  in  modern  literature  as  A.  lentiginosus  var.  Coulteri  or 
var.  Frctnontii:  the  Schott  element,  as  will  be  established  below,  belongs  to 
A.  agninus  Jeps. :  while  the  material  from  Boca  Grande,  a  fragmentary  speci- 
men with  leaflets  glabrous  on  the  upper  surface,  though  not  susceptible  of 
certain  determination,  is  probably  a  form  of  A.  lentiginosus  var.  yuccanus 
Jones  (certainly  not  Cystium  eremicum  as  annotated  by  Rydb.).  In  settling 
on  the  type  of  A.  Arthu-Schottii  the  latter  may  be  quickly  discarded :  not 
only  was  it  cited  last  by  Gray,  but  the  only  reference  to  it  in  the  description 
seems  to  be  "foliolis  .  .  .  pagina  superiore  interdum  gUibrata,"  and  it  must 
have  formed  a  quite  unimportant  part  of  the  species  that  the  author  had  in 
mind.  The  choice  between  the  Fremont  and  Schott  elements,  however,  is  more 
difficult,  for  the  description  fits  both  with  almost  equal  exactness.  Neverthe- 
less the  phrases  dealing  with  the  calyx — "calycis  dentihus  suhulatis  tuba 
ohlongo-campanulato  vix  dimidio  longiorihus" — and  with  the  keel — "carina 
majuscula  suhrecta  vexillo  paullo  breviore" — are  matched  more  closely  by  the 
corresponding  organs  of  Schott's  plants  than  by  those  of  Fremont's  and  the 
description  seems  to  have  been  prepared  primarily  from  the  former,  the  first 
collected  and  more  copious  specimens.  For  this  reason,  as  well  as  to  avoid 
the  anomaly  of  excluding  from  the  species  named  in  honor  of  Schott  the  very 
elements  which  he  himself  collected,  it  seems  proper  to  regard  the  Boundary- 
Survey  Xo.  256  in  herb.  Gray  as  the  type. 


160  CALIFORNIA  ACADEMY  OF  SCIENCES        [Proc.  4th  ser. 

As  stated  above,  the  material  of  A.  Arihu-Schottii  available  to  Gray  was 
without  frnit,  and  this  is  true  both  of  the  sheet  now  in  herb.  Gray  as  well  as 
of  tlie  two  in  the  Torrey  herbarium  (NY).  In  the  Field  Museum,  however, 
there  are  three  additional  isotypic  sheets,  apparently  from  Schott's  private 
herbarium,  one  of  which  has  nearly  mature  legumes  and  belongs,  quite  incon- 
testably,  to  A.  agninus  Jeps.  Even  without  this  further  evidence  the  identity 
of  Schott's  plant  would  be  almost  certain  from  locality  alone.  The  six  sheets 
seen  by  the  writer  are  variously  annotated  "Colorado  Desert  near  Fort  Yuma," 
"Colorado  Desert,  Sonora"  and  "Between  San  Diego  and  the  Colorado,"  and 
whether  or  not  these  represent  one  or  more  separate  gatherings  it  is  clear  that 
all  came  either  from  extreme  southern  Imperial  County,  California,  or  from 
closely  adjacent  Sonora,  a  territorj^  from  M^hich  A.  Coultcri  is  totally  un- 
known yet  w'here  A.  agninus  has  been  frequently  collected.  Nevertheless,  start- 
ing with  Watson  in  1871,  the  Schott  collection  has  been  regailarly  determined 
by  all  students  of  the  genus  as  A.  Coiilteri  and  is  so  annotated  by  Sheldon 
(NY)  and  cited  by  Jones  and  Jepson.  The  error  is  perhaps  traceable  back 
to  Gray's  publication,  as  early  as  1867,  (/.  c),  of  a  revised  description  of 
A.  Arthu-ScJiottii  taken  from  fruiting  material  communicated  by  Cooper 
from  the  Mohave  Desert  ("Camp  Cad^v  and  elsewhere,"  G.)  W'hicli  is  con- 
specific  with  the  early  Fremont  collection  but  not  with  that  of  Schott,  i.  e. 
A.  Coulteri  var.  It  should  be  borne  in  mind  that,  at  the  time  when  Gray  pro- 
posed A.  Arthu-Schottii,  A.  Coulteri  was  known  to  him  only  from  Bentham's 
description,  and  it  is  not  surprising  therefore  that,  when  A.  Coulteri  was 
finally  identified  with  the  Cooper  plant  in  fruit,  the  whole  oiA.  Arthu-Schottii 
was  also  thought  to  belong  there. 

In  the  North  American  Flora  (/.  c.)  Rydberg  abandoned  the  accepted  inter- 
pretation of  A.  Arthu-Schottii,  recognizing  it  as  a  species  in  the  genus  Cyn- 
tium,  but  the  effect  is  somewhat  marred  by  the  fact  that  he  recognized  Cystium 
agninum  and  C.  Coulteri  as  well.  If  one  may  judge  from  specimens  in  the  New 
York  Botanical  Garden  so  annotated  by  Rydberg  himself,  his  C.  Arthu- 
Schottii  contained,  in  addition  to  the  original  Schott  collection  (which  he  also 
regarded  as  typical),  a  number  of  plants  from  the  Mohave  Desert  such  as 
Parish  4957,  Elmer  3641,  etc.,  which  are  inseparable  from  the  earlj'  Fremont 
collection,  although  that  actual  specimen  he  referred  to  C.  Coulteri  (Bth.) 
Rydb. !  It  should  also  be  remarked  that  in  the  same  work  {I.e.,  in  syn.)  Rydberg 
incorrectly  cited  A.  lentiginosus  var.  horreganus  as  having  been  published  by 
Jones  in  the  Revision  of  Astragalus,  whereas  it  actually  had  appeared  as 
early  as  1898  (cf.  supra)  and  in  addition  failed  to  note  Jones'  excellent  figure. 

A.  Arthu-Schottii  is  a  plant  of  rather  restricted  distribution  in  the  south- 
w^estern  deserts  from  interior  San  Diego  County,  California,  at  Borrego  Wells, 
Orcutt,  type  of  var.  horreganus  (PO),  Jepson  8883,  type  of  A.  agninus  (NY, 
fragments),  to  Yuma,  Arizona,  Beckett  &  Nobles  8695  (G,  F),  southward 
just  into  Sonora  about  San  Luis,  Harrison  &  Kearney  8434  (F),  and  north 
along  the  Colorado  River  to  eastern  San  Bernardino  County,  California,  at 


VOL.XXV]     BAENEBY:  PUGILLUS  ASTEAGALORUM  ALTEE  161 

Silver  Lake,  J.  T.  Howell  3586  (CAS)  and  Nipton  on  the  borders  of  Nevada, 
Ripley  d-  Barnehy  3360.  The  species  is  very  closely  allied  to  A.  Coulteri,  which 
it  replaces  south  of  the  Coachella  Valley,  but  differs  in  the  annual  root,  lax 
fruiting  racemes  and  scarcely  inflated  pod. 

Astragalus  lentiginosus  Dougl.  var.  csesariatus  Barneby  var.  nov. 

(Plate  17,  figs.  29-30) 

Astragalu.s  lc7)figinosus  Dougl.  var.  ccesariaUts  Barneby  var.  nov.  inter  Cysfia  neogea  Eyd- 
bergiana  sive  Astragali  Diphysos  A.  Gray  adnumeranda,  caulibus  prostratis,  floribus 
congestis  et  legumine  coriaceo  A.  tehatchapiensi  (Eydb.)  Tidestr.  proxime  affinis,  sed 
ab  eo  floribus  maximis  ca.  19  mm.  longis  saturate  violaceis,  dentibus  calycinis  longio- 
ribus  acutis,  legumine  aliruptius  acuminato  necnon  racemorum  axi  calycibus  bracteisque 
pube  e  pilis  nigris  laxius  appressis  constituta  crebre  indutis,  satis  superque  diversa.  A. 
lentiginosus  var.  nigricalycis  M.  E.  Jones,  qui  etiam  in  iisdem  finibus  passim  oceurrit, 
caulibus  paucis  elatis,  raceme  laxiori,  floribus  minoribus  saepissime  ochroleucis,  pube 
crispa  implexa  et  legumine  omnino  albo-villosulo  nee  partim  atro-strigoso  a  nostra  dis- 
tantius  recedit. 

California  :  grassy  slopes  of  the  Temblor  Range  above  McKittrick,  Kern  Co.,  alt.  2900 
ft.,  13  April,  fl.  and  fr.  Eipley  and  Barneby  3243.  Type  in  Herb.  Calif.  Acad.  Sci. 
No.  289572. 

The  proposed  variety  is  here  referred  to  A.  lentigiiwsus — sensu  latissimo — 
in  order  to  fit  into  Dr.  Jepson's  treatment  of  this  complex  group  in  the  Flora 
of  California,  but  the  disposition  is  probably  not  final.  The  writer  has  spent 
many  months  in  study  of  A.  lentiginosus  and  its  allies  and  it  is  now  believed 
that  a  taxonomic  arrangement  based  on  a  specific  concept  intermediate  be- 
tween the  extremes  of  Rydberg,  who  recognized  33  species  in  Cystium,  and 
of  Jones,  who  regarded  all  the  known  races  as  varieties  of  A.  lentiginosus, 
is  not  impossible.  In  spite  of  the  fact  that  the  specific  limits  would  be  some- 
what blurred,  it  would  appear  feasible  to  recognize  at  least  three  major 
species,  A.  lentiginosus,  A.  Coulteri  Bth.  and  A.  diphysus  A.  Gray  (approxi- 
mately corresponding  with  Rydberg's  three  series  Lentiginosa,  Coulteriana 
and  Diphysa  of  Cystium),  under  which  the  remainder,  with  several  as  yet 
undescribed,  might  reasonably  be  classified  as  varieties.  In  this  case  the  var. 
ccesariatus  would  be  referred  to  A.  diphysus  and  associated  closely  with 
A.  tehatchapiensis  (Rydb.)  Tidestr.  (Proc.  Biol.  Soc.  Wash.  50:21.  1937,  as 
tehachapiensis)  yet  easily  separable  by  the  larger,  brilliantly  colored  corolla, 
acute  calyx-lobes,  sinuous  stems,  and  by  the  pubescence  of  loosely  appressed 
black  hairs  which  invest  the  w^hole  axis  of  the  raceme,  the  pedicels  and  calyces, 
and  which  recur,  mixed  with  white  villi  of  similar  structure,  on  the  valves  of 
the  more  abruptly  cuspidate  pod.  A.  lentiginosus  var.  nigricalycisM.  E.  Jones, 
the  common  race  of  the  San  Joaquin  Vallej^  differs  from  ours  in  its  taller 
stems,  more  lax  racemes  of  smaller  ochroleucous  flowers,  and  by  the  tangled, 
curly  pubescence  of  the  stem,  herbage  and  legume,  the  valves  of  which  are 
larger  and  of  thinner  texture. 


162  CALIFOBXIA  ACADEMY  OF  SCIENCES        [Proc.  4th  see. 

A.  lentiginosus  var.  ccesariatus  is  not  uncommon  on  the  eastern  slope  of  the 
Temblor  Range,  where  it  forms  large  clumps  among  the  ephemeral  spring 
grasses,  man}'  annuals  characteristic  of  the  flora  of  the  Great  Valley  foothills, 
and  such  perennial  herbs  as  Senecio  Breweri  Davy  and  Delphiniiim  hesperium 
var.  recurvatum  Jeps.  The  stems,  arising  from  a  heavy  root  and  multicipital 
caudex,  are  very  numerous,  reclinate  below  but  with  ascending  tips,  and  the 
subcapitate  or  short  and  dense  racemes  of  large  purple  flowers  with  their 
contrasting  striate  eye  on  the  banner  make  the  variety  one  of  the  showiest  of 
California  Astragali.  The  specific  epithet  is  derived  from  the  Latin  'ccesaries' 
and  refers  to  the  abundance  of  dark  hairs  in  the  inflorescence. 

Astragalus  striatiflorus  M.  E.  Jones,  Proc.  Calif.  Acad. 
Sei.II,  5:643. 1895 

(Plate  17,  figs.  19-24) 

Utah:  locally  abundant  in  white  sand  and  on  rock-ledges  beneath  sandstone  cliff>s,  10 
miles  north  of  Johnson  Ranch,  east  of  Kanab,  Kane  Co.,  alt.  5600  ft.,  6  May,  flor. 
Eipley  and  Barnehy  4358.  Ibid.,  5  June,  fl.  and  fr.  No.  4815.  Also  beneath  the  white 
cliffs  5  miles  south  of  Zion  Park  Junction,  Kane  Co.,  alt.  6100  ft.  No.  4365. 

This  exceedingly  obscure  species  lias  been  known  up  to  the  present  only 
from  an  inadequate  fragment,  devoid  of  pods,  which  was  gathered  by  Jones 
*'in  red  sand,  above  Springdale"  (the  western  entrance  to  Zion  Park,  Wash- 
ington County,  Utah)  in  September,  1894.  The  type  collection,  Jones  6080k 
(PO) ,  consists  of  a  single  very  mature  plant  from  which  even  the  flowers  have 
now  fallen.  The  characteristic  habit,  however,  the  pubescence  and  curiously 
connate  stipules  still  visible  in  the  dried  material,  when  taken  in  conjunction 
with  the  description  of  the  flower  and  its  attenuate  keel,  leave  little  doubt  that 
our  recent  collections  are  determined  correctly.  Jones  was  apologetic  about  de- 
scribing a  species  of  Astragalus  wdthout  knowledge  of  the  fruit,  but,  con- 
vinced by  the  floral  and  other  peculiarities  of  his  material  that  he  had  before 
him  a  species  "certainly  belonging  to  the  hiflati  and  probably  near  to  A.  ser- 
pens" (Z.  c. ) ,  he  was  unable  to  resist  giving  it  a  name.  That  his  proposition  was 
valid  is  now  obvious,  but  in  referring  A.  striatiflorus  to  the  Inflati  (approxi- 
matel}^  the  genus  Phaca  as  defined  by  Rydberg)  Jones  was  far  from  the  mark : 
actually  the  pod,  while  inflated,  is  bilocular  and  the  affinity  of  the  plant  is 
probably  with  A.  platytropis  A.  Gray.  In  the  text  of  the  Revision  of  Astraga- 
lus the  species  is  not  mentioned,  but  in  the  Index  (which  is  not  paginated)  it 
is  reduced  without  comment  to  A.  Silerauus  M.  E.  Jones,  a  true  member  of 
the  Inflati  with  discrete  stipules,  quite  different  flowers  and  elongated  stems. 
Rydberg,  to  whom  the  species  was  entirely  unknown,  cited  A.  striatiflorus  in 
his  list  of  omitted  synonyms  in  Astragalus  (N.  Amer.  Fl.  24  :455.  1929),  and 
could  do  little  else  but  follow  Jones'  earlier  reduction.  As  the  species  remains 
virtually  undescribed  to  this  day,  it  has  been  thought  advisable  to  prepare 
a  revised  description  with  emphasis  on  the  floral  and  carpological  characters. 


VOL.XXV]     BARNEBY:  PUGILLUS  ASTEAGALOEUM  ALTER  163 

Astragalus  striatiflorus  M.  E.  Jones,  char,  fruct. 

A  low  cespitose  perennial  herb  with  a  fleshy  taproot  and  loosely  branched  subterranean 
caudex,  the  stems  and  herbage  densely  and  softly  villous-caneseent  with  white  spreading 
and  somewhat  curly  hairs.  Stems  very  short,  decumbent,  clothed  below  with  loosely  imbri- 
cated stipules,  the  upper  2  or  3  internodes  about  1  cm.  long.  Stipules  very  broadly  obovate, 
truncate  and  often  mucronate,  2-3  mm.  long,  scarious  and  many-nerved  (the  nerves  more 
or  less  anastomosing),  adnate  neither  to  the  stem  nor  to  the  petiole  but  connate  around 
both  to  form  a  cup,  strigose-villous  without.  Leaves  2-3  cm.  long,  the  slender  rachis  and 
petiole  subequal.  Leaflets  11-13,  petiolulate,  obovate  to  oblanceolate,  obtuse,  truncate  and 
mucronulate  or  rarely  a  little  emarginate,  often  conduplicate,  contiguous,  the  terminal 
smallest.  Racemes  short,  loosely  3-5-flowered,  on  peduncles  about  equalling  the  leaves. 
Bracts  ovate,  acute  or  acuminate,  scarious,  longer  or  shorter  than  the  spreading  pedicels. 
Calyx  purplish,  villous-hirsute  with  white  or  some  black  hairs,  the  tube  campanulate  or 
obscurely  obconic,  3-4  mm.  long,  oblique  at  the  orifice,  the  acute  teeth  2-3  mm.  long,  hetero- 
morphous,  the  ventral  pair  deltoid-acuminate,  the  lateral  pair  and  the  dorsal  one  linear- 
subulate.  Corolla  9-12  mm.  long,  ochroleucous  or  suffused  with  lavender,  the  banner  striate 
with  purple  lines  and  the  keel  deeply  purple-tipped.  Banner  with  suborbicular  blade  8-9  mm. 
broad,  narrowed  into  a  very  short  claw,  arcuate  much  below  the  middle  nearly  to  a  right- 
angle,  the  margins  spreading  but  not  reflexed.  Wings  8-9  mm.  long,  the  blades  hmate 
obtuse,  about  2  mm.  broad,  including  the  large  basal  auricle  6  mm.  long.  Keel-petals  equal- 
ing the  wings,  the  blades  6-7  mm.  long,  obliquely  lanceolate  and  arcuate  upwards,  strongly 
gibbous  above  the  claw  and  there  attached  to  the  wings,  the  upper  margins  concave  at  the 
middle,  the  lower  margins  straight  below,  toward  the  middle  abruptly  convex  and  arched 
upward  to  the  slightly  declined  apex  of  the  long-attenuate  but  obtuse  beak,  free  for  1 
mm.  below  the  summit.  Style  filiform,  at  full  anthesis  exserted  about  1  mm.  from  the  keel, 
geniculate  below  the  capitate  stigma.  Pod  bladdery-inflated,  bilocular,  12-1.5  mm.  long, 
spreading,  sessile  but  seated  on  a  minute  boss  within  the  calyx  and  readily  deciduous  there- 
from, in  outline  broadly  ellipsoid,  rounded  and  somewhat  oblique  at  base,  abruptly  acute 
and  apiculate  at  the  apex  which  is  crowned  by  the  coiled  persistent  style,  shallowly  sulcate 
along  either  suture,  the  ventral  suture  nearly  straight,  filiform,  the  dorsal  introflexed  to 
form  a  thin  double  partition  3  mm.  high  produced  across  the  entire  width  of  the  cavity,  the 
valves  membraneous,  densely  villous  with  white  curled  spreading  hairs,  mottled  with  purple, 
the  cross-section  suborbicular  6-7  mm.  broad  and  a  little  less  high,  emarginate  at  the  sulcate 
sutures.  Seeds  about  8  in  either  locule,  obliquely  reniform,  compressed,  alwut  2  mm.  long, 
olive-green. 

A.  striatiflorus  is  set  off  from  all  known  species  of  Astragalus  by  a  combina- 
tion of  characters  which  excludes  it  from  a  place  in  any  natural  group  hitherto 
defined.  An  attempt  to  determine  the  species  from  Kydberg's  monograph 
leads  the  enquirer  directly  to  Cystium  platy trope  (A.  Gray)  Rydb.,  or,  more 
deviously,  to  A.  anisus  M.  E.  Jones,  but  the  similarity  between  these  plants 
is,  I  believe,  rather  superficial  and  based  entirely  on  the  structure  of  the  pod. 
In  the  present  paper  it  is  not  proposed  to  discuss  the  taxonomic  position  of 
A.  anisus,  which  differs  {inter  alia)  from  A.  platytropis  and  A.  striatiflorus 
in  the  elongate  flower  and  cylindric  calyx-tube  and  which  is  probably  cor- 
rectly placed  by  Jones  and  Rydberg  as  an  aberrant  relative  of  tlie  Mollissimi 
(A.  mollissimus  Torr.  et  spp.  affin.).  But  the  differences  between  the  two  last 
and  the  relationship  of  each  to  A.  lentiginosus  sens,  latiss.,  i.  e.  Cystium  of 
Rydberg',  will  be  examined  in  more  detail. 

It  has  already  been  implied  that  a  similarity  between  the  pods  of  two  given 


164  CALIFOBXIA  ACADEMY  OF  SCIEXCES        [Pkoc.  4th  ser. 

species  of  Astragalus  is  not  necessarily  a  criterion  of  affinity,  but  this  assertion 
must  be  amplified,  for,  thus  baldly  stated,  it  stands  in  direct  contradiction  to 
Rydberg's  most  important  principle  of  classification.  All  of  Rydberg's  later 
work  on  the  genus,  which  culminated  in  his  monographic  revision  for  the 
North  American  Flora,  is  based  on  the  primary  assumption  that  carpological 
characters,  even  when  of  the  most  minute  and  quantitative  nature,  are  of 
supreme  value  in  the  definition  of  groups  or,  in  his  terminology,  of  genera. 
Diversity  in  structure  of  the  flower,  stipules  or  pubescence  he  discarded  as 
of  quite  secondary  importance.  By  logical  application  of  this  principle,  Ryd- 
berg  was  led  to  assume  an  affinity  between  species,  scattered  perhaps  from  the 
Levant  to  the  Andes,  which  happened  to  have  a  similar  width  of  septum  in 
the  pod,  yet  simultaneously  to  refer  to  widely  separated  groups  two  species 
intimately  connected  by  ties  of  morphology  and  geography  in  which  the  sep- 
tum was  found  to  be  strongly  or  weakly  developed.  Of  course  this  method  was 
satisfactory  in  parts,  for  many  allied  species  do,  as  one  would  expect,  bear 
similar  pods,  but  in  some  cases  it  produced  polyphyletic  groups  of  great  arti- 
ficiality. Had  Rydberg  himself  admitted  this,  it  could  be  easily  forgiven.  But 
interior  evidence  from  his  published  writings  leaves  little  doubt  that  he  pro- 
posed his  genera  as  genuine  phylae,  as  evolved  expressions  of  a  single  heredity 
marked  by  a  carpological  structure  common  to  all  included  members.  And 
implicit  in  that  belief  was  the  conclusion  that  any  two  members  of  one  genus 
are  more  closely  related  to  each  other  than  either  can  be  to  a  member  of  another 
genus.  With  this  principle  in  mind  let  us  take  an  example  from  Rydberg's 
own  work. 

Tiv))i  Wilsonii  (Greene)  Rydb.  is  a  species  of  Astragalus  endemic  to  the 
high  plateaux  of  northern  Arizona,  described  by  Greene  as  related  to  A.  diphy- 
sus  A.  Gray  (Cystium  diphysum  Rydb.)  and  reduced  by  Jones  in  synonymy 
to  a  variety  of  A.  lentiginosus  Dougl.  The  pod  is  lanceolate,  leathery,  slightly 
but  not  conspicuously  inflated,  with  the  valves  introverted  dorsally  to  form 
a  partition  and  thus  partially  bilocular.  The  septum  does  not  quite  reach  the 
ventral  suture,  but  the  gap  is  not  above  one  millimeter  in  width.  With  this 
we  may  compare  Cystium  agninum  (Jeps.)  Rydb.,  discussed  elsewhere  in  this 
paper.  Here  we  have  a  pod  of  similar  outline,  little  inflated,  but  the  septum 
traverses  the  whole  width  of  the  cavity  (sometimes  less :  cf .  Jeps.  ex  char. : 
".  .  .  extending  half  way  or  nearly  to  the  ventral  suture")  and  the  gap  between 
the  sutures  is  as  a  rule  closed.  Rydberg  rightly  disregarded  the  lack  of  infla- 
tion in  the  fruit  of  C.  agninum,  otlierwise  anomalous  in  his  genus  Cystium, 
yet  attached  sufficient  importance  to  the  equally  quantitative  character  of 
the  septum  to  refer  A.  Wilsonii  to  Tinm,  and  we  are  led  to  believe  that  despite 
their  manifest  similarity  in  all  other  respects  to  A.  coulteri  and  A.  diphysns, 
both  Rydbergian  Cystia,  these  two  species  of  the  southwestern  states  are  more 
closely  related  to,  respectively,  the  old-world  A.  Cicer  L.,  genotype  of  Cystium 
and  A.  sulcatus  L.,  genotype  of  Tium,  than  tliey  are  to  one  another. 

This  relationship  is  not.  of  course,  ipso  facto,  impossible.  But  once  tlie  rela- 


Vol.  XXV]     BAENEBY:  PUGILLUS  ASTRAGALOEUM  ALTEE 


165 


tionship  is  admitted  we  must  also  accept  that  the  evolution  of  the  septum 
stopped  abruptly  at  an  incalculably  distant  date  and  has  since  been  station- 
ary, while  the  differentiation  of  species  has  continued  along  other  lines. 
However,  when  the  marvelously  diverse  carpology  of  Astragalus  is  consid- 
ered as  a  whole,  and  when  one  recollects  the  manner  in  which  the  pods  of 
species  accepted  by  Rydberg  himself  as  closely  allied  may  differ  in  size  and 
shape,  in  length  of  stipe,  in  the  degree  of  compression  and  inflation,  and  even 
(cf.  Tium  platycarjnim  and  T.  racemosum)  in  the  very  width  of  septum,  it 
becomes  clear  that  all  parts  of  the  fruit  have  been  equally  subject  to  evolu- 
tionary progress  and  recession  in  comparatively  recent  times.  Indeed  there 
seems  to  be  no  reason  why  the  similarity  of  structure  between  the  pods  of  Old 
and  New  World  species,  remarked  by  Rydberg  and  other  taxonomists  before 
him,  cannot  be  explained  as  well  by  parallel  development  as  by  a  common 
heredity.  In  this  case  no  single  carpological  character,  nor  even  general  agree- 
ment in  structure  of  the  fruit,  can  be  accepted  as  a  reliable  basis  of  a  natural 
classification  throughout  the  genus  unless  supported  by  other  criteria  of 
proved  value  :  and  the  names  of  generic  segregates  such  as  Cysiium  and  Tium, 
founded  on  European  or  Asiatic  species,  have  a  very  questionable  place, 
whether  applied  to  genera,  subgenera  or  sections,  in  the  taxonomy  of  Ameri- 
can Astragalus. 

From  this  statement  of  generalities  we  may  well  return  to  the  particular 
subject  of  the  present  note,  A.  striatiflorus  and  its  immediate  allies.  In  the 
interests  of  clarity  the  major  characters  of  the  species  are  contrasted  in  the 
following  table  with  A.  platytropis  and  A.  lentiginosus. 

A.  lentiginosus 
strongly  caulescent 
strictly  appressed 
free  or  obscurely 

adnate  to  petiole 
lunate  obtuse 

shorter  than  wings 
included 

with  septum  formed 
as  in  A.  striatiflo- 
rus, the  ventral 
suture  sometimes 
(especially  in  allied 
species)  very  nar- 
rowly introflexed 
but  never  below  in- 
sertion of  funicles 

It  will  be  seen  at  once  that  A!  striatiflorus  differs  from  its  supposed  allies 
(or,  more  precisely,  from  other  species  with  bilocular,  inflated  pods)  in  the 
attenuate  keel,  exserted  style,  stipules  and  pubescence,  and  additionally  from 
A.  platytropis,  of  which  it  has  nearly  the  habit,  in  the  structure  of  the  septum. 
A.  platytropis  (plate  26,  figs.  25-28),  on  the  other  hand,  is  unique  in  the 
concretion  of  the  stipules,  in  the  septum  and  broadly  truncate  keel.  In  the 


A  striatiflorus 

A.  platytropis 

Stems 

shortly  caulescent 

subacaulescent 

Pubescence 

villous 

loosely  strigose 

Stipules 

connate  around  stem 

connate  abaxially 

and  petiole 

about  the  petiole 

Keel 

lance-attenuate 

oblong-truncate 

equalling  wings 

equalling  wings 

Style 

exserted 

included 

Pod,  bilocular 

with   septum   formed 

with   septum   formed 

and  inflated 

by  intrusion   of 

by  a  narrow  inflex- 

dorsal   suture,    the 

ion    of   the    dorsal 

ventral  filiform 

suture  and  a  deep- 
ly   produced    wing 
from    the    ventral 
which   extends   be- 
low   the    insertion 
of  the  funicles 

](i(i  CALIFOBXIA  ACADEMY  OF  SCIENCES        [Pkoc.  4th  ser. 

opinion  of  the  writer  the  sum  of  characters  peculiar  to  each  isolates  these 
species  so  fundamentally  from  the  rest  of  Astragalus  as  to  entitle  them  to 
rank  as  representative  of  sections  in  the  genus.  The  following  names  are  there- 
fore proposed. 

Astragalus  sect.  Orocystium  Barneby  sect.  iiov.  Cystium  ser.  Platytropia  Rydb..  X.  Ainer. 
Fl.  24:405.  1929. 

Legumen  vesicario-inflatum,  siitura  dorsali  anguste  introplicata  ventralique  intus  pio- 
ducta  dimidiam  loculi  altitudinem  superanti  biloculare:  carina  oblongo-truncata : 
stipulae  circa  petiolum  abaxialiter  concretae,  altrinsecus  liberae.  Species  imica : 
A.  platytropis  A.  Gray. 

Astragalus  sect.  CystieUa  Barneby  sect.  nov. 

Legumen  vesicario-inflatum,  sutura  dorsali  introflexa  biloculare:  stylus  ad  antliesin 
exsertus:  carina  attenuata:  stipulae  multinerviae  utrinque  concretae  cupuliformes. 
Species  unica  :  A.  striatiflorus  M.  E.  Jones. 

Following  the  conclusion  reached  above  that  the  New  World  species  re- 
ferred to  Cystium  by  Rydberg  are  probably  not  allied  to  the  original  elements 
of  Steven,  and  as  this  group  has  no  acceptable  sectional  name  in  the  form  of 
a  substantive,  I  also  propose  : 

Astragalus  sect.  Diplocystium  Barneby  nom.  et.  stat.  nov.  Astragalus  ser.  Astragalus  sect. 
Diphysi  A.  Gray,  Proc.  Amer.  Acad.  6:192.  1863.  Astragalus  ser.  Eu-asiragalus  sect. 
Lentiginosus  Slield.,  Minn.  Bot.  Stud.  1 :168.  1894,  exclus.  spp.  plur.  Cystium  Rydb.,  Bull. 
Torr.  CI.  32:659.  1905;  N.  Amer.  Fl.  24:405,  sequ.  1929,  exclus.  C.  platytropide ;  Amer. 
Journ.  Bot.  17:232.  1930,  non  Cystium  Stev.,  nee  Astragalus  subgen.  Cercidothrix  sect. 
Cystium  Bge.  Astragalus  sect.  Inflati  M.  E.  Jones,  Rev.  Astrag.  123,  sequ.  1923,  quoad  sp. 
ultimam  p.  max.  p.  Astragalus  subgen.  Uiphysus  Tidestr.,  Contrib.  Nat.  Herb.  25:3]5. 
1925.  Species  typica:  A.  lentiginosus  Dougl.  ex  Hook. 

While  some  pains  have  been  taken  to  point  out  what  are  believed  to  be 
fundamental  differences  between  these  three  sections,  the  converse  and,  I 
suppose,  more  valuable  task  of  tracing  their  origin  and  real  place  in  the  genus 
cannot  be  attempted  here.  In  the  absence  of  exact  cytological  data  such  specu- 
lation is  apt  to  be  valueless  and  in  any  case  is  not  susceptible  of  proof.  I  am  in- 
clined to  suspect,  nevertheless,  that  Diplocystium  is  derived  from  a  common 
source  with  the  Mollissimi:  Orocystium  and  CystieUa  seem  to  be  more  closely 
connected  with  each  other  than  with  any  known  species,  but,  in  associating 
A.  platytropis  with  A.  cohrensis  A.  Gray,  Jones  may  have  hit  on  a  real  line 
of  affinity. 


EXPLANATION  OF  PLATE 


PLATE  17 

Figs.  1-9.  Astragalus  musimonum  Bavuel^y,  spec.  nov.  (1)  Pod  x  2.  (2) 
Cross-section  of  pod  x  3.  (3)  Flower  x  2.  (4)  Calyx  x  2.  (5)  Banner  x  2.  (6) 
Wing-petal  X  2.  (7)  Keel  x  2.  (8)  Leaf  x  1.  (9)  Stipules  x  2. 

Figs.  10-18.  Astragalus  Bryantii  Barueby,  spec.  nov.  (10)  Podxl.  (11) 
longitudinal  cross-section  of  same  x  1.  (12)  transverse  section  of  same  x  3.  (13) 
Pod  of  broader  type  x  1.  (14)  transverse  section  of  same  x  3.  (15)  Flower  x  2. 
(16)  Wing-petal  x  2.  (17)  Keel  X  2.  (18)  Leaf  X  1. 

Figs.  19-24.  Astragalus  striatiflorus  M.  E.  Jones.  (19)  Pod  x  2.  (20)  Cross- 
section  of  pod  x  2.  (21)  Flower  X  2.  (22)  Keel  x  2.  (23)  Seed  x  4.  (24)  Stip- 
ules X  3. 

Figs.  25-28.  Astragalus  platytropis  A.  Gray.  (25)  Cross-section  of  pod  x  1. 
(26)  Keel  X  2.  (27)  Stipules  x  3. 

Fig.  28.  Astragalus  en-siformis  var.  gracilior  Barneby,  var.  nov.  (28)  pod  x 
11/2. 

Figs.  29-30.  Astragalus  lentiginosus  var.  ccesariatus  Barneby,  var.  nov.  (29) 
Pod  x  11/2.  (30)  Flower  X  11/2. 

Fig.  31.  Astragalus  Whitedii  fma.  speirocarpoides  Barneby  fma.  nov.  (31) 
Pod  X  2. 


[168] 


PROC.   CALIF.    ACAD.    SCI..    4TH    SERIES.    VOL.    XXV.    NO.    3 


[BARNEBY]  PLATE  17 


[  169  J 


PROCEEDINGS 

OF  THE 

CALIFORNIA  ACADEMY  OF  SCIENCES 

Fourth  Series 
Vol.  XXV,  No.  4,  pp.  171-176  Jiiiie  1,  1944 


ALICE  EASTWOOD 
SEMI-CENTENNIAL  PUBLICATIONS 

No.  4 

SUBLITTORAL  MARINE  ALGAE  OF  THE 
MONTEREY  PENINSULA 

BY 

GILBERT  M.  SMITH 

Professor  of  Biology  (Botany),  Stanford  University 


TV  TUMEROUS  INVESTIGATIONS  liBve  been  made  on  the  marine  algae  of  the  Pacific 
1\|  Coast  of  Nortli  America  but  attention  has  been  directed  almost  exclu- 
sively to  those  collected  in  the  intertidal  zone.  Statements  concerning  sub- 
littoral  algae  (those  growing  below  extreme  low  tide  level)  have  been  based 
upon  the  assumption  that  algae  found  cast  ashore  on  the  beach  and  not  known 
to  grow  locally  in  the  intertidal  zone  must  have  come  from  deep  water.  Here 
and  there  in  the  literature  is  mention  of  a  few  algae  dredged  along  the  western 
shores  of  North  America.  The  only  exception  is  the  region  in  the  vicinity  of 
Friday  Harbor,  Washington,  where  there  has  been  an  intensive  exploration 
by  dredging.  Of  the  126  species  of  red  algae  that  Kylin  (1925)  records  from 
this  area,  75  were  found  only  in  the  intertidal  zone,  33  were  found  only  by 
dredging,  and  18  were  found  both  in  the  intertidal  zone  and  in  deep  water 
(5-10  fathoms). 

For  several  years  I  have  been  engaged  upon  a  survey  of  the  marine  algae 
of  the  Monterey  Peninsula,  California.  In  addition  to  collecting  intensively 
in  the  intertidal  zone,  numerous  collections  have  been  made  from  a  depth  of 
15-60  feet  below  the  extreme  low  tide  level.  At  the  time  this  investigation  was 
commenced  the  only  definite  records  of  sublittoral  algae  from  the  Monterey 
Peninsula  were  the  following:  Dictyo7ieuropsis  reticuhifa  (Saund.)  G.  M. 
Smith  by  Saunders  (1895)  ;  Myriogramme  spectahilis  (Eaton)  Kylin  by  Nott 
(1900);  Macrocystis  pyrifera  (L.)  C.  A.  Ag.  and  Nereocysfis  Luetkeana 
(Mert.)  Post,  and  Rupr.  by  McFarland  (1912)  ;  and  Lithothamnion  mon- 
feveyicnm  Foslie  by  Foslie  (1906). 

[171] 


172  CALIFORXIA  ACADEMY  OF  SCIENCES        [Proc.  4th  sek. 

The  first  collections  from  tlie  sublittoral  by  the  writer  were  made  with  the 
assistance  of  three  students  (Vernon  Brock,  Charles  Danforth,  and  Herbert 
Armstrong)  in  the  summer  of  1934.  By  means  of  a  diving  helmet,  several  algae 
were  collected  from  a  depth  of  15-35  feet  in  the  vicinity  of  Mussel  Pt.  Since 
1934  all  collections  from  deep  water  have  been  by  dredging  from  a  small  boat 
equipped  with  a  20-horsepower  engine.  The  only  really  favorable  place  for 
dredging  is  the  submerged  shale  reef  lying  at  a  depth  of  30-35  feet  a  quarter 
mile  east  of  the  municipal  wharf  at  Monterey.  Here  the  rock  is  so  soft  that 
large  pieces  of  rock  with  algae  attached  to  them  are  broken  loose  by  the  dredge. 
Elsewhere  on  the  Monterey  Peninsula  the  rock  on  the  ocean  bottom  is  granitic 
and  it  is  extremely  difficult  to  break  off  pieces  with  the  dredge.  Here,  also, 
the  ocean  bottom  is  so  rough  that  the  dredge  is  continually  catching  between 
the  rocky  crags,  and  on  any  dredging  expedition  one  can  expect  to  spend  at 
least  half  the  time  trying  to  dislodge  the  dredge.  In  spite  of  these  difficulties 
the  sublittoral  region  between  Mussel  Pt.  and  Pt.  Pinos  has  been  rather  thor- 
oughly explored.  So,  also,  has  the  land-locked  cove  (Stillwater  Cove)  adjoin- 
ing Pebble  Beach.  No  attempt  has  been  made  to  dredge  between  Pt.  Pinos 
and  Cypress  Pt.  because  of  the  danger  in  taking  a  small  boat  close  to  the  shore 
in  this  wave-swept  portion  of  the  coast  line. 

When  blades  of  algae  are  found  loose  in  the  dredge  there  is  always  the  prob- 
lem of  distinguishing  between  those  from  plants  growing  in  the  sublittoral 
and  those  broken  off  from  intertidal  plants  and  lying  unattached  on  the  ocean 
bottom.  For  this  reason  every  effort  was  made  to  obtain  intact  plants  attached 
to  pieces  of  rock  and  thus  know  with  certainty  that  the  alga  is  from  the  sub- 
littoral. In  the  following  list  of  algae  from  the  sublittoral  one  or  more  speci- 
mens of  all  the  species  have  been  found  attached  to  rock  dredged  from  the 
ocean  bottom.  Some  of  the  algae  brought  up  by  the  dredge  are  not  found  in 
the  intertidal  zone  of  the  Monterey  Peninsula,  or  are  known  only  from  isolated 
individuals  collected  there.  These  are  considered  true  sublittoral  algae.  Other 
■of  the  algae  are  regularly  found  in  the  intertidal  zone.  These  are  interpreted 
as  algae  the  upper  limit  of  whose  vertical  distribution  is  approximately  the 
mean  low  tide  level.  For  the  sake  of  comparison  the  occurrence  or  non- 
occurrence in  the  intertidal  zone  of  the  Monterey  Peninsula  is  given  for  each 
of  the  species,  as  is  also  the  intertidal  range  with  respect  to  mean  low  tide  level. 

PHAEOPHYTA 

ECTOCARPALES 

Ectocarpus  granulosus  (J.  E.  Smith)  C.  A.  Ag.  At  30-35  feet  near  munici- 
pal wharf,  Monterey;  at  15-20  feet,  Stillwater  Cove.  Common  between  the 
0.5  and  -1.5  foot  tide  levels. 

SPHACELARIALES 

Sphacelaria  didichotoma  Saund.  At  15-20  feet,  Stillwater  Cove.  Known 
from  the  intertidal  zone. 


VOL.XXV]  SMITH:  SUBLITTOEAL  ilAEINE  ALGAE  173 

DESMARESTIALES 

Desmarestia  linearis  Gardner,  mss.  At  60  feet  near  Monterey.  Not  known 
from  the  intertidal  zone. 

Desmarestia  herhacea  (Turn.)  Lmx.  At  30-35  feet  near  municipal  wharf, 
Monterey.  Common  between  the  0.5  and  -1.5  foot  tide  levels. 

PUNCTARIALES 

Stictyosiphon  tortilis  (Rupr.)  Reinke.  At  30-35  feet  near  municipal  wharf, 
Monterey.  Not  known  from  the  intertidal  zone. 

LAMINARIALES 

Dictyoneurum  calif  or  nicum  Rupr.  At  25  feet  near  Mussel  Pt. ;  at  15-20 
feet,  Stillwater  Cove.  Common  between  the  0.5  and  -1.5  foot  tide  levels. 

Dictyoneuropsis  reticulata  (Saund.)  G.  M.  Smith.  At  30-35  feet  near  mu- 
nicipal wharf,  Monterey;  at  20-25  feet  near  Pt.  Aulon.  Not  known  from  the 
intertidal  zone. 

Nereocystis  Luetkeana  (Mert.)  Post,  and  Rupr.  Numerous  beds  growing 
at  a  depth  of  30-50  feet.  Isolated  plants  found  between  the  0.0  and  -1.5  foot 
tide  levels. 

Macrocystis  pyrifera  (L.)  C.  A.  Ag.  Numerous  beds  growing  at  a  depth  of 
30-50  feet.  Not  known  from  the  intertidal  zone. 

Pterygophora  calif ornica  Rupr.  Growing  in  extensive  stands  at  a  depth  of 
20-60  feet  between  Cypress  Pt.  and  Pescadero  Pt.  Isolated  plants  found  be- 
tween the  0.0  and  -1.5  foot  tide  levels. 

FUCALES 

Cystoseira  osmundacea  (Menzies)  C.  A.  Ag.  At  20-35  feet  near  Mussel  Pt. 
and  near  Pt.  Aulon.  Common  between  the  0.0  and  -1.5  foot  tide  levels. 

RHODOPHYTA 

BANGIALES 

Go7iiotrichum  elegans  (Cliauv.)  Zanard.  At  35-40  feet  near  municipal 
wharf,  Monterey ;  at  25-30  feet  near  Mussel  Pt. ;  at  20-25  feet  near  Pescadero 
Rock.  Isolated  plants  collected  at  mean  low  tide  level. 

Goniotrichopsis  siiblitt oralis  G.  M.  Smith.  At  30-35  feet  near  municipal 
wharf,  Monterey.  Not  known  from  the  intertidal  zone. 

Erythrotrichia  carnea  (Dillw.)  J.  G.  Ag.  At  30-35  feet  near  municipal 
wharf,  Monterey,  and  near  Pt.  Aulon.  Common  between  the  2.0  and  -1.5  foot 
tide  levels. 

NEMALIONALES 

Rhodochorton  concrescens  Drew.  At  25-30  feet  near  Pt.  Aulon;  at  35-40 
feet  near  Pt.  Pinos.  Known  from  the  mean  low  tide  level. 

Rhodochorton  Amphiroae  Drew.  At  25-30  feet  near  Pt.  Aulon.  Known  from 
the  mean  low  tide  level. 


174  CALIFOENIA  ACADEMY  OF  SCIENCES        [Proc.  4th  ser. 

CRYPTONEMIALES 

Pikea  pinnata  Setch.  At  30-35  feet  near  municipal  wliai-f ,  ]\Ionterey ;  at 
35-40  feet  near  Mussel  Pt.  Isolated  plants  found  between  the  0.5  and  -1.5 
foot  tide  levels. 

Pikea  californica  Harv.  At  20-25  feet  near  Pt.  Anion.  Widespread  but 
scarce  between  the  0.0  and  -1.5  foot  tide  levels. 

Peyssonnelia  pacifica  Kylin..At  30-35  feet  near  municipal  wharf.  ]\Ion- 
terey;  at  20-25  feet,  Stillwater  Cove.  Fairly  common  between  the  0.0  and 
-1.5  foot  tide  levels. 

Lithothamnion  montereyicum  Foslie.  At  70  feet  near  Monterey.  Not  known 
from  the  intertidal  zone. 

Bossea  Orhigniana  (Dene.)  Manza.  At  30-35  feet  near  municipal  wharf, 
Monterey ;  at  20-25  feet  near  Pt.  Anion.  Not  known  from  the  intertidal  zone. 

Calliarthron  Setchelliae  Manza.  At  20-25  feet  near  Pt.  Aulon;  at  30-35  feet 
near  Pt.  Pinos.  Common  between  the  0.0  and  -1.5  foot  tide  levels. 

Halymenia  californica  Smith  and  Hollenb.  At  30-35  feet  near  munici]ial 
wharf,  Monterey.  Not  known  from  the  intertidal  zone. 

Prionitis  lanceolata  Harv.  At  40-60  feet  near  Mussel  Pt. ;  at  20-25  feet  near 
Pt.  Aulon  and  at  Stillwater  Cove.  Abundant  between  the  1.0  and  -1.5  foot 
tide  levels. 

Callophyllis  marginifructa  Setch.  and  Swezy.  At  30-35  feet  near  municipal 
wharf,  Monterey  ;  at  12-15  feet  near  Pt.  Anion;  at  15-20  feet,  Stillwater  Cove. 
Scarce  between  the  0.0  and  -1.5  foot  tide  levels. 

CallophyUis  megalocarpa  Setch.  and  Swezy.  At  15-20  feet  near  Mussel  Pt. 
and  at  Stillwater  Cove.  Scarce  between  the  0.0  and  -1.5  foot  tide  levels. 

Callocolax  neglectus  Schmitz.  At  30-35  feet  near  municipal  wharf.  ]Mon- 
terey.  Not  known  from  the  intertidal  zone. 

GIGARTINALES 

Schizymenia  pacifica  Kylin.  At  30-35  feet  near  municipal  wharf,  Monterey. 
Common  between  the  0.0  and  -1.5  foot  tide  levels. 

Agardhiella  Coulteri  (Harv.)  Setch.  At  20  feet  near  Mussel  Pt.  and  at 
Stillwater  Cove.  Abundant  between  the  1.0  and  -1.5  foot  tide  levels. 

Plocamivm  pacificum  Kylin.  At  20-25  feet  near  Pt.  Aulon  and  near  Pesea- 
dero  Rock.  Common  between  the  0.0  and  -1.5  foot  tide  levels. 

Gracilaria  Sjoestcdfii  Kylin.  At  30-35  feet  near  municipal  wharf,  Monterey 
at  40-60  feet  near  Mussel  Pt.  Abundant  between  the  1.0  and  -1.5  foot  tide 
levels. 

Gracilann  rohusta  Setch.  At  15-25  feet  near  Pt.  Aulon.  Isolated  plants 
found  between  the  -0.5  and  -1.5  foot  tide  levels. 

Ahnfeltia  gigartinoides  J.  A.  Ag.  At  12-15  feet  near  Pt.  Aulon.  Not  known 
from  the  intertidal  zone. 

Stenogramma  californica  Harv.  At  30-35  feet  near  municipal  wharf.  ]\Ion- 
terey ;  at  20-25  feet  near  Pt.  Anion.  "Relatively  scarce  between  the  0.5  and  -1.5 
foot  tide  levels. 


Vol.  XXV]  SMITH:  SVBLIITOBAL  MABIXE  ALGAE  175 

Gigartina  Boryi  Setch,  and  Gardii.  At  12-15  feet  near  Pt.  Anion.  Common 
between  the  1.0  and  -1.5  foot  tide  levels. 

Gigartina  corymhifera  (Ktz.)  J.  G.  Ag.  At  20-25  feet  near  Mussel  Pt.  and 
at  Stillwater  Cove.  Common  between  the  0.0  and  -1.5  foot  tide  levels. 

Gigartina  Harveyana  (Ktz.)  Seteh.  and  Gardn.  At  25  feet  near  Pt.  Anion. 
Common  between  the  1.0  and  -1.5  foot  tide  levels. 

RHODYMENIALES 

Rhodymenia  calif ornica  Kylin.  At  20-25  feet  near  Pt.  Anion  and  at  Still- 
water Cove.  Common  between  the  0.0  and  -1.5  foot  tide  levels. 

Rhodymenia  yaciflca  Kylin.  At  60-75  feet  near  Mussel  Pt. ;  at  15-20  feet, 
Stillwater  Cove.  Common  between  the  0.0  and  -1.5  foot  tide  levels. 

Gastroclo7iium  Coulteri  (Harv.)  Kylin.  At  20-25  feet  near  Mussel  Pt.;  at 
15-20  feet,  Stillwater  Cove.  Abundant  between  the  2.0  and  -1.5  foot  tide  levels. 

Coeloseira  compressa  Hollenb.  At  20-30  feet  near  Pt.  Anion.  Not  known 
from  the  intertidal  zone. 

CERAMIALES 

Afitithamnion  defectum  Kylin.  At  30-35  feet  near  Pt.  Pinos.  Not  known 
from  the  intertidal  zone. 

Antithamnion  dendroideum  G.  M.  Smith.  At  30-35  feet  near  municipal 
wharf,  Monterey.  Not  known  from  the  intertidal  zone. 

Antithamnion  glandidiferum  Kylin.  At  30-35  feet  near  municipal  wharf, 
Monterey.  Rare  between  the  0.0  and  -1.5  foot  tide  levels. 

Platythamnion  heteromorphum  J.  G.  Ag-.  At  30-35  feet  near  municipal 
wharf,  Monterey.  Not  known  from  the  intertidal  zone. 

Platythamnion  villosum  Kylin.  At  30-35  feet  near  municipal  wharf,  Mon- 
terey ;  at  25-30  feet  near  Pt.  Anion.  Rare  between  the  0.0  and  -1.5  foot  tide 
levels. 

Platythamnion  pectinatum  Kylin.  At  30-35  feet  near  municipal  wharf, 
Monterey.  Rare  between  the  0.0  and  -1.5  foot  tide  levels. 

Pleonosporium  dasyoides  (J.  G.  Ag.)  DeToni.  At  35-50  feet  near  Pt.  Anion; 
at  15-20  feet,  Stillwater  Cove.  Scarce  between  the  0.0  and  -1.5  foot  tide  levels. 

Pleonosporium  vancouverianum  J.  G.  Ag.  At  30-35  feet  near  municipal 
wharf,  Monterey.  Not  known  from  the  intertidal  zone. 

Spermothamnion  Snyderae  Farl.  At  30-35  feet  near  municipal  wharf,  Mon- 
terey ;  at  35-40  feet  near  Mussel  Pt. ;  at  30-35  feet  near  Pt.  Pinos ;  at  20-25 
feet,  Stillwater  Cove.  Common  between  the  0.5  and  -1.5  foot  tide  levels. 

Griffithsia  pacifica  Kylin.  At  25-30  feet  near  Pt.  Anion  ;  at  30-35  feet  near 
Pt.  Pinos;  at  20-25  feet  near  Pescadero  Rock.  Scarce  between  the  0.5  and  -1.5 
foot  tide  levels. 

Microcladia  Coidteri  Harv.  At  20-25  feet  near  Mussel  Pt.  and  at  Stillwater 
Cove.  Abundant  between  the  1.0  and  -1.5  foot  tide  levels. 

Branchioglossum  Woodii  (J.  G.  Ag.)  Kylin.  At  30-35  feet  near  municipal 
wharf,  Monterey,  and  near  Pt.  Pinos.  Very  rare  between  the  -0.5  and  -1.5 
foot  tide  levels. 


17G  CALIFORNIA  ACADEMY  OF  SCIEXCES        [Proc.  4th  ser. 

Polyneura  latissinia  (Harv.)  Kylin.  At  30-35  feet  near  municipal  wharf, 
Monterey,  and  near  Mussel  Pt. ;  at  12-15  feet  near  Pt.  Anion.  Common  be- 
tween the  0.0  and  -1.5  foot  tide  levels. 

Phycodrys  Setchellii  Skottsb.  At  30-35  feet  near  municipal  wharf,  Mon- 
terey; at  20-30  feet,  Stillwater  Cove.  Very  rare  between  the  0.0  and  -1.5  foot 
tide  levels. 

Niendurgia  Andersoniana  (J.  G.  Ag.)  Kylin.  At  20-25  feet  near  Pt.  Anion 
and  at  Stillwater  Cove.  Very  scarce  between  the  0.0  and  -1.5  foot  tide  levels. 

Myriogramme  spectdbUis  (Eaton)  Kylin.  At  20  feet  near  Mussel  Pt. ;  at 
70-90  feet  near  Pt.  Aulon.  Not  known  from  tlie  intertidal  zone. 

Cryptopleura  violacea  (J.  G.  Ag:)  Kylin.  At  30-35  feet  near  municipal 
wharf,  Monterey ;  at  20-25  feet  near  Pt.  Aulon ;  at  15-20  feet,  Stillwater  Cove. 
Abundant  between  the  0.5  and  -1.5  foot  tide  levels. 

Botryoglossum  Farlowianum  (J.  G.  Ag.)  DeToni.  At  30-35  feet  near  mu- 
nicipal wharf,  Monterey;  at  20-25  feet  near  Pt.  Aulon;  at  15-20  feet,  Still- 
water Cove.  Abundant  between  the  0.0  and  -1.5  foot  tide  levels. 

Dasyo'psis  densa  G.  M.  Smith.  At  30-35  feet  near  municipal  wharf,  Mon- 
terey. Not  known  from  the  intertidal  zone. 

PoJysiphonia  calif ornica  Harv.  At  20-25  feet  near  Pescadero  Rock.  Common 
between  the  1.0  and  -1.5  foot  tide  levels. 

Polysiphonia  flaccidissima  var.  Smithii  Hollenb.  At  35-40  feet  between 
Mussel  Pt.  and  Pt.  Aulon.  Not  known  from  the  intertidal  zone. 

Pterosiphonia  dendroidea  (Mont.)  Falkenb.  At  20-25  feet  near  Pt.  Aulon 
and  near  Pescadero  Rock.  Abundant  between  the  0.0  and  -1.5  foot  tide  levels. 

Pterosiphonia  Baileyi  (Harv.)  Falkenb.  At  20-25  feet  near  Mussel  Pt.  and 
near  Pt.  Aulon.  Common  between  the  0.5  and  -1.5  foot  tide  levels. 

Hcrposiplionia  rigida  Gardn.  At  30-35  feet  near  municipal  wharf,  Mon- 
terey. Rare  between  the  0.0  and  -1.0  foot  tide  levels. 

HerposipJionia  pygmnea  Plollenb.  niss.  At  20-25  feet  near  Pt.  Aulon ;  at 
35-40  feet  near  Pt.  Pinos.  Not  found  in  the  intertidal  zone. 

Laurencia  spectahilis;  Post,  and  Rupr.  At  15-20  feet,  Stillwater  Cove. 
Abundant  between  the  1.0  and  -1.5  foot  tide  levels. 

Laurencia  Gardneri  Hollenb.  At  30  feet  near  Mussel  Pt. ;  at  35-40  feet  near 
Pt.  Aulon  ;  at  30-35  feet  near  Pt.  Pinos ;  at  20-25  feet  near  Pescadero  Rock. 
Isolated  plants  found  between  the  0.0  and  -1.5  foot  tide  levels. 

BIBLIOGRAPHY 

Foslio,  M.  1906.  Algologiske  Notiser  II.  Kgl.  Norske  Vidcnsk.  ScLsk.  Skr.  1906,  No.  2:1-28. 
Kylin,  H.  1925.  The  marine  red  algae  in  the  vicinity  of  the  Ijiological  station  at  Friday 

Harbor,  Wash.  Lunds  Univ.  Irsskr.  N.  F.  21,  Nr,  9:1-87.  47  figp 
McFarland,  F.  M.  1912.  The  kelps  of  the  central  Californian  coast.  Appendix  M  (pp.  194- 

208),  Senate  Document  190,  2d.  Session,  62d.  Congress. 
Nott,  C.  P.  1900.  Nitophylla  of  California.  Proc,  Calif.  Acad.  Sci.  3  Ser.  Bot.,  2:1-62.  9  pi, 
Saunders,  DeA.  1895.  A  preliminary  paper  on  Costaria  with  description  of  a  new  species. 

Rot.  Gaz.  20:54-57.  1  pi. 


i 

I 


PROCEEDINGS 
OF  THE 

CALIFORNIA  ACADEMY  OF  SCIENCES 

Fourth  Series 
Vol.  XXW  No.  5.  j)j).  177-188,  pis.  18-19  June  1.  i()ji 


ALICE  EASTWOOD 
SEMI-CENTENNIAL  PUBLICATIONS 

No.  5 

THE  GANDER  OAK,  A  NEW  HYBRID  OA 
FROM  SAN  DIEGO  COUNTY,  CALIFORNI 

BY 

CARL  B.  WOLF 

Botanist,  Rancho  Santa  Ana  Botanic  Garden 


IX  THE  NEARLY  a  ceiitiirv  and  a  half  since  Nee  published  the  first  two  species 
of  Quercus  from  California  (Quercus  agrifolia  and  lohata)  the  list  has 
grrown  so  that  now  at  least  eig"hteen  generally  recopfnized  species  are  believed 
to  constitute  the  oak  flora  of  the  state.  In  addition  to  these  species  there  are 
several  rare  forms  which  have  evoked  considerable  interest.  One  of  the  first 
of  tliese  to  be  brought  to  the  attention  of  botanists  was  the  Oracle  Oak  ( X 
Quercus  morehus  Kellogg,  Proe.  Calif.  Acad.  Nat.  Sci.,  2  :36,  1863),  long  sus- 
pected of  being  a  hybrid  between  Quercus  Kelloggii  Newb.  and  Quercus 
Wislizenii  A.  DC,  but  only  recently  so  demonstrated  by  cultuijil  experiments 
at  Rancho  Santa  Ana  Botanic  Garden  (Rancho  Santa  Ana  Bot.  Gard..  Occas. 
Papers,  1:47-52.  1938). 

In  1935  Mr.  Frank  F.  Gander,  botanist  of  the  Museum  of  Natural  History 
at  San  Diego,  California,  discovered  another  peculiar  oak  somewhat  similar 
to  the  Oracle  Oak.  This  tree  grew  along  the  road  between  Santa  Ysabel  and 
Lake  Henshaw.  Shortly  thereafter  ]\Ir.  Gander  informed  us  of  his  find  and 
we  visited  the  area,  located  the  original  tree  and  several  others  like  it.  In  the 
fall  of  1939  we  obtained  acorns  of  this  new  oak  for  propagation  at  Rancho 
Santa  Ana  Botanic  Garden.  The  results  of  our  studies  in  the  field  and  in  the 
Garden  are  embodied  in  tlie  present  paper.  It  is  a  pleasure  to  publish  this 
oak  as  new  in  honor  of  ]\Ir.  Gander  in  the  Proceedings  of  the  California 
Academy  of  Sciences,  since  one  of  the  parents  of  this  hybrid  is  Quercus  Kel- 
loggii, a  species  honoring  Dr.  Albert  Kellogg,  one  of  tlie  founders  of  the 
Academy,  and  also  a  most  devoted  student  of  our  oaks.  Ilis  ])eautiful  draw- 

[177] 


178  CALIFOBXIA  ACADEMY  OF  SCIEXCES        [Pkoc.  4th  ser. 

ings  of  oaks  were  published  in  1889  (Illustrations  of  West  American  Oaks, 
with  the  text  by  Dr.  E.  L.  Greene).  It  has  also  seemed  appropriate  that 
this  article  should  appear  in  the  volume  dedicated  to  Miss  Alice  Eastwood 
because  she  has  long  been  interested  in  oaks  and  published  one  of  the  last  of 
the  generally  recognized  entities  for  California  {Quercus  Alvordmna  Eastw., 
Handbook  of  the  Trees  of  California,  p.  48,  pi.  27,  fig.  4, 1905) .  She  is  a  charter 
member  of  the'  Rancho  Santa  Ana  Botanic  Garden  Advisory  Board  whei'e  her 
enthusiasm  for  the  Garden's  experimental  work  has  been  a  constant  stimulus 
for  us  to  go  forward  with  the  numerous,  painstaking,  slow  and  often  dis- 
appointing cultural  projects,  many  of  which  seem  to  yield  so  little  in  pro- 
portion to  the  time  and  energy  required  for  their  completion. 

X  Quercus  Ganderi  C.  B.  Wolf,  hybr.  nov. 

{Quercus  Kelloggi'Newh.  X  Quercus  agrifolia  var.  oxyadenia  [Torr.]  J.  T. 
Howell.) 

Tree  10-16  m.  high;  6-10  m.  spread;  trunk  up  to  1  m.  in  diam.,  the  bark 
dark  brown  or  gray,  comparatively  smooth  and  essentially  like  that  of  Quercus 
Kelloggii.  In  aspect  the  tree  is  erect,  usually  is  taller  than  broad  and  more 
nearly  resembles  Q.  Kelloggii  than  it  does  Q.  agrifolia  var.  oxyadenia.  Leaves 
nearly  evergreen,  turning  yellow  in  January  and  shedding  shortly  before  the 
new  leaves  appear.  Branchlets  densely  puberulent,  but  in  the  second  year 
becoming  nearly  glabrous.  Petioles  1-2.5  cm.  long;  blades  coriaceous,  slightly 
cupped,  glossy  green  above,  but  microscopically  stellate-pubescent,  especially 
along  the  midribs  and  veins,  gray-green  beneath  and  at  first  heavily  stellate- 
pubescent,  but  in  age  becoming  much  less  so,  5-10  cm.  long,  3-6  cm.  wide, 
shalloAvly  and  irregularly  lobed  with  3  or  4  main  lobes  on  each  side,  these  in 
turn  irregularly  margined  and  bearing  1^  bristle-tipped  teeth.  Acorns  soli- 
tar}'  or  2-3  in  a  cluster,  produced  abundantly  some  years,  and  apparently 
maturing  the  first  season ;  styles  3,  about  2-3  mm.  long;  cup  12-18  mm.  high, 
15-25  mm.  broad,  the  inner  surface  hairy  at  base,  nearly  glabrous  at  margins, 
scales  thin,  light  tan  with  a  darker  border  and  mostly  with  ciliated  margins, 
well  imbricated  in  6-8  rows;  nut  with  lower  one-third  covered  by  the  cup, 
oblong,  25-35  mm.  long,  lightly  pubescent  to  nearly  glabrous  without,  densely 
pubescent  within,  rich  light  brown  at  maturity.  (The  acorns  are  scarcely 
distinguishable  from  those  of  Q.  Kelloggii,  except  that  the  nut  is  somewhat 
slenderer  pointed.)  Staminate  aments  slender,  pubescent,  borne  in  clusters  of 
several ;  each  ament  4r-6  cm.  long,  bearing  about  15-20  flowers,  each  with  6-8 
stamens. 

Arbor  erecta,  oirciter  10-16  m.  altitndine;  rajiiiilis  junioribiis  tonientellis ;  foliis  lialiitus 
Quercus  Kelloggii,  5-10  cm.  longLs,  3-6  cm.  latis;  pagiiia  superiore  sub-glabra,  inforiore 
stellata-tomentosa;  cupulis  12-18  mm.  altitudine,  squamis  planus;  glandibus  oblongis, 
25-35  mm.  longis. 

Habitat.  Lower  borders  of  the  Arid  Transition  and  the  upper  borders  of 
the  Upper  Sonoran  Life  Zones  at  elevations  of  3000  to  3200  feet,  in  association 


Vol.  XXV]  WOLF:  THE  GANDER  OAK  179 

with  Quercus  Kelloggii,  agrifolia  var.  oxyadenia  and  Engelmannii,  growing 
on  hillsides  and  near  occasional  streams  in  granitic  type  soils. 

Range.  Known  only  from  the  mountains  of  San  Diego  County,  California. 
All  of  the  known  trees  are  located  in  a  rather  restricted  area  extending  from 
the  Volcan  Indian  School,  located  north  of  Santa  Ysabel,  to  a  mile  or  two 
west  of  Lake  Henshaw  Dam  and  to  the  vicinity  of  Mesa  Grande. 

Type  Locality.  San  Diego  Count}',  California,  about  300  yards  north  of  the 
entrance  to  the  Volcan  Indian  School  on  California  State  Highway  No.  79 
between  Santa  Ysabel  and  Lake  Henshaw,  at  3100  feet  elevation,  in  decom- 
posed granite  soil,  near  an  occasional  stream  just  west  of  the  highway.  Type 
Specimen :  Carl  B.  Wolf  and  Percij  C.  Everett,  Herb.  No.  9543  {Prop.  No. 
3555),  collected  October  20,  1939,  the  tree  being  16  m.  high,  10  m.  spread,  its 
trunk  branching  into  three  main  forks  near  the  ground,  two  of  these  being 
about  46  cm.  in  diam.,  and  the  third  about  20  cm.  in  diam.  The  Type  Specimen 
is  deposited  in  Rancho  Santa  Ana  Botanic  Garden  Herbarium,  sheet  no.  23585. 
Isotypes  widely  distributed  as  indicated  below. 

Localities  and  Specimens  Examined.  All  of  the  localities  mentioned  below 
are  in  San  Diego  County,  California. 

1).  Near  the  entrance  to  the  Volcan  Indian  School  between  Santa  Ysabel 
and  Lake  Henshaw,  at  3100  feet  elevation:  Frank  F.  Gander,  Nov.  14,  1935, 
and  May  6,  1937;  C.  B.  Wolf  and  P.  C.  Everett,  Herh.  No.  9483,  January  10, 
.1939;  P.  C.  Everett,  Herh.  No.  9529,  July  28,  1939;  C.  B.  Wolf  and  P.  C. 
Everett,  Herh.  No.  9543  {Prop.  No.  3555),  Oct.  20,  1939,  Type  Collection. 
(All  of  the  above  specimens  were  made  from  the  same  tree.) 
•  2) .  Between  Santa  Ysabel  and  Lake  Henshaw,  on  the  Morelli  Ranch,  located 
about  2  miles  south  of  the  junction  of  the  road  to  Warner's  Hot  Springs  and 
Henshaw  Dam,  3000  feet  elevation :  C.  B.  Wolf,  Herh.  No.  9487,  January  24, 
1939.  from  a  tree  10  m.  high,  10  m.  spread,  and  a  trunk  60  cm.  in  diam.;  and 
from  tlie  same  tree,  C.  B.  Wolf  and  P.  C.  Everett,  Herb.  No.  9542  {Prop.  No. 
5554),  October  20,  1939. 

3).  Mesa  Grande,  M.  V.  Hood,  Nov.  24,  1940. 

4) .  One  and  eight-tenths  miles  from  Mesa  Grande  on  the  road  to  Lake  Hen- 
shaw, at  3200  feet  elevation :  C.  B.  Wolf,  Herb.  No.  9488,  January  24,  1939, 
from  a  tree  11-12  m.  high,  6-7  m.  spread,  and  a  trunk  35  cm.  in  diam. ;  C.  B. 
Wolf,  Herh.  No.  9489,  January  24,  1939,  from  a  tree  12-13  m.  high,  11-12  ra. 
spread,  and  a  trunk  nearly  1  m.  in  diam. 

5) .  Between  Mesa  Grande  and  Lake  Henshaw  f north  of  the  above  locality) . 
Several  trees  were  noted  on  January  24,  1939,  on  the  slopes  away  from  the 
road,  but  for  lack  of  time  no  collections  were  made. 

6) .  On  the  hills  to  the  south  and  west  of  Lake  Henshaw  Dam.  The  trees  noted 
in  this  area  on  January  24,  1939,  appeared  to  be  the  northwest  limit  of  dis- 
tribution for  the  Gander  Oak  in  this  region,  but  for  lack  of  time  no  specimens 
Avere  collected. 

In  its  native  habitat  the  individual  trees  of  the  Gander  Oak  are  difficult  to 


180  CALIFOEKIA  ACADEMY  OF  SCIENCES        [Proc.  4th  ser. 

detect  (luring  most  of  the  j^ear  except  upon  close  examination,  for  they  are 
mixed  with  the  Kellogg  Oak,  which  the.y  closely  resemble.  However,  in  the  late 
fall  or  early  winter  they  can  readily  be  spotted  on  the  hillsides.  At  those  times 
the  leaves  of  the  Kellogg  Oak  have  been  shed,  while  those  of  the  Gander  Oak 
are  beginning  to  turn  yellow.  In  January  of  1939  we  were  able  to  locate  be- 
tween 20  and  30  Gander  Oaks  in  the  vicinity  of  Lake  Henshaw  and  Mesa 
Grande  merely  by  driving  slowly  along  the  roads  that  traverse  the  areas. 
Exploration  of  the  adjacent  hillsides  not  visible  from  these  roads  would  likely 
result  in  increasing  this  figure  considerably.  It  is  also  probable  that  the  Gan- 
der Oak  occurs  elsewhere  in  San  Diego  County,  but  thus  far  no  additional 
localities  have  been  brought  to  our  attention.  All  of  the  trees  of  the  Gander 
Oak  which  we  have  seen  in  the  wild  appear  to  be  tirst-generation  crosses  be- 
tween the  two  parents,  despite  the  fact  that  fertile  acorns  appear  to  be  pro- 
duced somewhat  regularly.  However,  this  lack  of  second-generation  offspring 
in  the  wild  is  in  keeping  Avith  the  behavior  of  many  other  wild  hybrids. 

Wild  trees  of  X  Querciis  Ganderi  can  be  distinguished  from  Quercus  Kel- 
loggii  by  the  less  deeply  lobed  and  slightly  cupped  leaves  with  considerable 
pubescence  beneath,  and  by  the  longer  persistence  of  the  foliage.  Separa- 
tion of  X  Quercus  Ganderi  from  X  Quercus  morehus  is  also  relatively  easy, 
although  the  leaves  of  the  two  are  very  similar  in  size  and  outline.  The  latter 
species  has  nearly  plane  leaves  whose  under  surfaces  are  practically  glabrous. 
Furthermore,  X  Quercus  morehus  requires  two  seasons  to  mature  its  acorns, 
whereas,  X  Quercus  Ganderi  appears  to  mature  its  acorns  in  a  single  season 
(as  does  Querciis  agrifolia  var.  oxyadenia). 

In  October,  1941,  Mr.  John  M.  Tucker  of  the  Department  of  Botany  of  the 
University  of  California,  Berkeley,  California,  forwarded  us  an  oak  specimen 
{Tucker  No.  418)  collected  on  the  University  of  California  Hastings  Wild 
Life  Reserve  in  the  Santa  Lucia  Mountains  of  Monterey  County.  This  speci- 
men appears  to  be  a  hybrid  between  Quercus  Kelloggii  and  Quercus  agrifolia, 
but  we  have  not  been  able  to  visit  the  area  and  see  the  tree  growing  in  the  wild, 
nor  has  it  been  possible  to  obtain  acorns  for  propagation.  Tucker's  specimen 
closely  resembles  X  Quercus  Gauderi,  but  is  essentially  glabrous.  The  status  of 
this  oak  should  be  determined  by  additional  field  studies  and  by  experimental 
methods.  If  it  should  be  demonstrated  that  it  is  a  hybrid,  as  suggested  above, 
it  could  hardly  be  included  in  the  present  concept  of  XQnercus  Ganderi 
unless  that  concept  be  enlarged  and  Quercus  agrifolia  var.  oxyadenia  be  re- 
duced to  synonomy  under  Quercus  agrifolia. 

Cultural  Studies 

Because  of  the  practical  difficulties  involved  in  making  experimental  crosses 
ill  Ihe  field  we  have  not  attempted  to  produce  X  Quercus  Ganderi  by  hybridiz- 
ing Quercus  Kelloggii  and  Quercus  agrifolia  var.  oxyadenia.  We  have,  how- 
ever, been  able  to  observe  the  behavior  of  seedlings  of  X  Quercus  Ganderi 
and  feel  that  they  have  yielded  sufficient  data  to  justify  our  conclusion  that 


Vol.  XXV]  WOLF:  THE  GANDEE  OAK  181 

the  tree  is  a  hybrid  between  Querciis  Kelloggii  and  Quercus  agrifolia  var. 
oxyadenia. 

In  October,  1939,  Mr.  Everett  and  I  hand-picked  over  400  acorns  from  the 
Type  Tree  of  X  Quercus  Ganderi  (Herb.  No.  9543,  Prop.  No.  3555),  and  11 
acorns  from  a  second  tree  in  the  same  region  (Herh.  No.  9542,  Prop.  No.  3554). 
These  acorns  were  subsequently  planted  in  our  lathhouse,  where  by  April, 
1940,  there  were  311  seedlings  of  the  former  and  8  of  the  latter.  In  contrast  to 
the  seedlings  of  X  Qiiercus  morehus,  grown  previously,  these  were  remarkably 
vigorous  and  attained  average  heights  of  over  30  cm.  the  first  season. 

Plantings  of  189  seedlings  (random  selections)  of  the  Gander  Oak  {Prop. 
No.  3555)  w^ere  made  in  the  Garden  on  March  10,  1941.  Only  61  of  these  sur- 
vived to  October,  1942.  This  unusually  high  loss  can  be  accounted  for  in  part 
because  at  that  time  the  Botanic  Garden  Nursery  was  moved  and  it  was  neces- 
sary to  dig  the  seedlings  and  hold  them  for  considerable  time  before  planting. 
The  remainder  of  the  seedlings  of  this  same  lot  were  lined  out  in  nursery  rows 
where  over  50  are  still  alive.  In  the  same  nursery  rows  are  5  seedlings  of  Prop. 
No.  3554.  These  seedlings  in  the  Garden  and  in  the  nursery  rows  now  show 
the  usual  intergradation  of  characters  displayed  when  seeds  of  a  hybrid  are 
propagated.  Those  seedlings  closely  resembling  Quercus  Kelloggii  are  now 
few  in  number,  for,  under  our  conditions,  that  species  has  always  been  diffi- 
cult to  grow.  The  rest  of  the  seedlings  now  alive  represent  several  typical 
Quercus  agrifolia  var.  oxyadenia,  a  large  number  of  forms  very  much  like  the 
original  X  Quercus  Ganderi,  and  many  of  varying  characteristics.  Some  have 
the  leaf  shapes  nearly  like  Quercus  Kelloggii,  but  are  exceptionally  pubescent. 
Others  are  like  Quercus  agrifolia  var.  oxyadenia,  but  are  glabrous.  Still  others 
have  numerous  intermediate  leaf  characters  and  are  even  more  unlike  either 
of  the  parent  species  of  the  original  cross.  It  will  be  many  years  before  acorns 
are  produced  by  these  seedlings,  but  they  will  likely  show  an  almost  continu- 
ous series  of  variations  between  the  slender  form  produced  by  the  one  parent 
and  the  thick  form  by  the  other.  It  is  also  probable  that  some  of  these  seedlings 
will  require  two  seasons  to  mature  acorns  while  others  should  do  so  in  a  single 
season. 

X  Quercus  Ganderi  may  eventually  prove  of  some  merit  for  horticulture, 
for  it  has  many  of  the  fine  features  of  Quercus  Kelloggii,  plus  semi-evergreen 
foliage,  a  vigorous  growth  rate  and  hardiness  at  low  elevations.  Acorns  gath- 
ered from  some  of  the  wild  trees  could  be  gi'own  in  the  nursery  for  two  or  three 
years,  so  that  selections  of  outstanding  horticultural  types  could  be  made 
before  making  permanent  plantings. 

The  accompanying  drawings  on  Plates  18  and  19  of  Quercus  Kelloggii, 
agrifolia  var.  oxyadenia,  Ganderi  and  the  nursery  seedlings  grown  from  the 
latter  were  prepared  from  tracings  (except  the  drawings  of  the  acorns) .  It  is 
hoped  that  they  will  convey  to  the  reader  not  only  the  characteristics  of  the 
parental  species  and  the  original  hybrid,  but  also  the  great  variation  exhibited 
by  the  seedlings. 


182  CALIFORNIA  ACADEMY  OF  SCIENCES        [Proc.  4th  SER. 

DISTRIBUTION  OF  HEKBARimi  SPECBIEXS 

Specimens  referred  to  in  the  text  and  in  the  figures  are  all  on  deposit  in  Eancho  Santa 
Ana  Botanic  Garden  Herbarium  with  the  exception  of  the  two  specimens  by  F.  F.  Gander 
and  the  one  by  M.  V.  Hood,  which  are  in  the  San  Diego  Museum  of  Natural  History  Her- 
barium. Duplicates  of  our  collections  of  X  Querent  Ganderi  have  been  distributed  as  follows: 

California  Academy  of  Sciences,  San  Francisco,  California :  Nos.  9483,  9487,  9488,  9489, 
9529,9543; 

Dudley  Herbarium,  Stanford  University,  California :  Nos.  9483,  9487,  9488,  9489,  95S9, 
9542,9543; 

Field  Museum  of  Natural  History,  Chicago,  Illinois:  No.  9543 ; 

Los  Angeles  Museum,  Los  Angeles,  California:  No.  9543 ; 

Missouri  Botanical  Garden,  St.  Louis,  Missouri:  No.  9543 ; 

New  York  Botanical  Garden,  New  York  City,  N.  Y. :  No.  9543; 

Philadelphia  Academy  of  Natural  Sciences,  Philadelphia,  Penna. :  No.  9543  ; 

Pomona  College,  Claremont,  California:  No.  9543 ; 

San  Diego  Museum  of  Natural  History,  San  Diego,  California :  Nos.  9483,  9487,  9488, 
9489,  9529,  9543; 

United  States  National  Herbarium,  Washington,  D.  C:  Nos.  9483,  9488,  9543  ; 

University  of  Arizona,  Tucson,  Arizona:  No.  9543 ; 

University  of  California,  Berkeley,  California :  Nos.  9483,  9487,  9488,  9489,  9529,  9543 ; 

University  of  California,  Los  Angeles,  California:  Nos.  9483,  9543 ; 

University  of  Colorado,  Boulder,  Colorado:  No.  9543 ; 

University  of  Montana,  Missoula,  Montana  :  No.  9543  ; 

University  of  Washington,  Seattle,  Washington  :  No.  9543. 


EXPLANATION  OF  PLATES 


PLATE  18 

Figs.  1-24.  Leaves  of  seedlings  of  X  Quercus  Ganderi  C.  B.  Wolf ;  collected 
from  Botanic  Garden  plantings  and  nursery  rows,  January  13,  1943,  Prop. 
No.  3555  (grown  from  acorns  taken  from  Type  Tree). 

Figs.  1-7.  From  seedlings  most  nearly  like  Quercus  Kelloggii. 

Figs.  8-15.  From  seedlings  most  nearly  like  Quercus  agrifolia  var.  oxya- 
denia. 

Figs.  16-24.  From  seedlings  most  nearly  like  X  Quercus  Ganderi  and  inter- 
mediate forms. 


[184] 


PROC.  CALIF.  ACAD.  SCI..  4TH  SERIES,  VOL.  XXV.  NO.  5  [WOLF]  PLATE  18 


[185] 


PLATE  19 

Figs.  1-10.  Quercus  Kelloggii  Newb. 

Figs.  11-18.  Quercus  agrifoliavav.  oxyadenia  (Torr.)  J.  T.  Howell. 

Figs.  19-30.  X  Quercus  Ganderi  C.  B.  Wolf. 

Fig.  1.  C.  B.  Wolf,  Herb.  No.  2488  (wild). 

Fig.  2.  ,7.  T.  Howell,  Herb.  No.  683  (wild). 

Figs.  3  and  4.  C.  B.  Wolf,  Herb.  No.  3992  (wild). 

Fig.  5.  C.  B.  Wolf  and  B.  D.  StarTc,  Herb.  No.  5482  (wild). 

Fig.  6.  P.  C.  Everett,  Herb  No.  608  i  (nursery  seedlings  of  Prop.  No.  1951). 

Fig.  7.  C.  B.  Wolf,  Herb.  No.  5572  (nursery  seedlings  of  Prop.  No.  1951). 

Fig.  8.  P.  C.  Everett,  Herb.  No.  6078  (nursery  seedlings  of  Prop.  No.  1952). 

Fig.  9.  B.  D.  Starlc,  Herb.  No.  798  (acorn,  wild). 

Fig.  10.  C.  B.  Wolf,  Herb.  No.  2488  (acorn,  wild). 

Fig.  11.  J.  T.  Howell,  Herb.  No.  600  (wild). 

Fig.  12.  B.  D.  StarTc,  Herb.  No.  5552  (wild). 

Fig.  13. 1.  L.  Wiggins  and  J.  W.  Gillespie,  No.  4053  (wild). 

Fig.  14.  B.  D.  StarTc,  Herb.  No.  5552  (wild). 

Fig.  15. 1.  L.  Wiggins  and  J.  W.  Gillespie,  No.  4053  (acorn,  wild). 

Fig.  16.  B.  B.  StarTc,  Herb.  No.  5552  (acorn,  Avild). 

Figs.  17  and  18.  P.  C.  Everett,  Herb.  No.  6074  (nursery  seedlings  of  Prop. 
No.  2026). 

Fig.  19.  C.  B.  Wolf,  Herb.  No.  9489  (wild,  as  are  also  nos.  20-30). 

Fig.  20.  C.  B.  Wolf  and  P.  C.  Everett,  Herb.  No.  9483. 

Fig.  21.  C.  B.  Wolf  and  P.  C.  Everett,  Herb.  No.  9543  (Type). 

Figs.  22  and  23.  C.  B.  Wolf  and  P.  C.  Everett,  Herb.  No.  9543  (acorns  from 
the  Type). 

Fig.  24.  P.  C.  Everett,  Herb.  No.  9529  (young  acorn). 

Fig.  2.5.  C.  B.  Wolf,  Herb.  No.  9488. 

Fig.  20.  C.  B.  Wolf,  Herb.  No.  9489. 

Fig.  27.  C.  B.  Wolf  and  P.  C.  Everett,  Herb.  No.  9542. 

Fig.  28.  C.  B.  Wolf  and  P.  C.  Everett,  Herb.  No.  9543  (Type). 

Fig.  29.  C.  B.  Wolf  and  P.  C.-Everett,  Herb.  No.  9487. 

Fig.  30.  C.  B.  Wolf  and  P.  C.  Everett,  Herb.  No.  9542. 


[180] 


PROC.  CALIF.  ACAD.  SCI..  4TH  SERIES,  VOL.  XXV.  NO.  5  [WOLF]  PLATE  19 


[187] 


PROCEEDINGS 
OF  THE 

CALIFORNIA  ACADEMY  OF  SCIENCES 

Fourth  Series 
Vol.  XX^^  No.  6,  pp.  189-214,  pis.  20-22  J""c  1,  1944 


ALICE  EASTWOOD 
SEMI-CENTENNIAL  PUBLICATIONS 

No.  6 

THE  GENUS  DRYMARIA  IN,  AND  ADJACENT  T 

THE  SONORAN  DESERT 

BY 

IRA  L.  WIGGINS 

Professor  of  Biology  (Botany),  Stanford  University 

THE  TAXONOMiST  frequently  finds  that  it  is  necessary  to  study  not  only 
those  represenstatives  of  a  genus  that  occur  within  a  circumscribed  area  in 
which  he  is  interested,  but  numerous  other  species  from  contiguous  regions  as 
well.  This  necessity  became  apparent  at  once  when  I  attempted  to  construct  a 
key  to  the  species  of  Drymaria  known  to  occur  within  the  limits  of  the  Sonoran 
Desert,  for  the  relationships  of  several  of  the  desert  species  were  obviously 
with  forms  growing  in  the  mountains  nearby,  or  in  the  drier  parts  of  New 
Mexico,  Texas,  and  northern  Mexico.  As  these  close  relatives  were  studied,  still 
others  had  to  be  taken  into  account  until  eighteen  entities  entered  into  the 
problem.  Of  this  number,  only  four  or  five  actually  grow  within  the  desert, 
but  it  seemed  desirable  to  present  the  total  results  of  the  study  rather  than 
to  restrict  the  paper  to  a  treatment  of  those  species  which  occur  regularly  in 
the  desert. 

Willdenow  recognized  four  species,  three  of  them  new,  when  he  set  up  the 
genus  Drymaria  in  1819  (in  R.  and  S.  Syst.  Veg.  5  :  406,  1819),  and  based  the 
new  genus  on  Holosteiim  cordatiim  Linn.  Five  years  later  DeCandolle  (Prodr. 
1 :  395,  1924)  listed  five  species,  the  fifth  having  been  described  by  Humboldt, 
Bonpland,  and  Kunth  in  1823  (Nov.  Gen.  et  Sp.  6  :  17-21,  pL  515-516, 1823). 
Since  that  time  there  has  been  a  steady  increase  in  the  number  of  species  recog- 
nized in  the  genus.  At  present,  one  hundred  nine  specific  and  varietal  names 
have  been  applied  in  Drymaria.  It  is  probable  that  a  thorough  investigation 
of  the  genus  would  require  reduction  of  certain  combinations  and  the  de- 
scription of  other  forms  not  as  yet  recognized. 

Most  of  the  species  and  varieties  of  Drymaria  occur  in  Mexico  and  the  west- 

[  189  ] 


190  CALIFORNIA  ACADEMY  OF  SCIENCES        [Proc.  4th  SER. 

ern  part  of  South  America.  Sixty-four,  exclusive  of  the  new  entities  proposed 
liere,  have  been  described  from  North  America,  about  thirty  from  South 
America,  several  from  the  West  Indies  and  Central  America,  one  species  is 
nearly  cosmopolitan  in  the  tropics,  and  one  occurs  in  Australia.  Over  fifty 
species  have  been  recorded  from  Mexico. 

The  genus  includes  both  annual  and  perennial  herbs,  the  latter  sometimes 
slightly  suffrutescent  at  the  base.  Most  species  are  clearly  annual  or  perennial, 
with  little  or  no  tendency  for  the  annuals  to  persist  beyond  a  single  growing 
season.  But  occasionally  a  plant  of  an  annual  species  ma}',  under  particularly 
favorable  conditions,  become  a  biennial  or  short-lived  perennial. 

The  species  vary  in  habit  from  prostrate,  through  procumbent  and  ascend- 
ing, to  strictly  erect  plants.  The  stems  may  be  simple,  unbranched,  or  pro- 
fusely branched  from  either  the  base  or  in  the  upper  part  of  the  plant.  The 
stems  and  branches  are  slender,  often  nearly  filiform  in  such  species  as  D. 
tenella,  D.  debilis,  and  D.  carinata.  The  internodes  may  be  rather  short,  as  in 
D.  crassifolia,  with  the  leaves  exceeding  the  internodes  and  forming  dense 
mats,  or  they  may  be  greatly  elongated  and  much  surpassing  the  leaves,  as 
in  the  closely  related  D.  holostcoides.  The  herbage  is  pubescent  in  some  species, 
glabrous  or  glabrate  in  others.  The  pubescence  may  be  glandular  or  non- 
glandular  in  different  species. 

The  leaves  are  useful  in  separating  the  genus  into  two  main  groups,  one 
of  which  possesses  leaves  that  are  ovate,  cordate,  or  broadly  elliptic  and  from 
half  as  wide  as  long  to  distinctly  broader  than  long.  The  other  group  has  leaves 
that  are  linear,  filiform,  or  narrowly  oblong  or  subulate.  These  leaves  show 
little  tendency  toward  an  elliptic  outline  and  are  always  several  times  as  long 
as  broad.  A  few  species  have  fleshy  or  leathery  leaves,  but  most  of  them  have 
thin,  herbaceous  leaves  of  delicate  texture. 

The  arrangement  of  the  flowers  in  the  inflorescence  is  racemose  or  cymose. 
The  former  arrangement  is  found  in  D.  holosteoides,  C.  crassifolia,  and  D. 
pacliyphijlla.  In  these  species  the  flowers  appear  to  be  in  dense  clusters  in 
tiie  axils  of  most  leaves  and  at  or  near  the  apices  of  the  branches.  Examination 
of  this  arrangement  reveals  that  the  flowers  are  borne  in  a  short  raceme,  each 
flower  subtended  by  a  small,  scarious  bract.  The  minute  bracts  nearly  conceal 
the  rachis.  Such  racemes  are  axillary  to  the  larger  leaves  of  the  fascicles 
formed  at  each  node.  In  other  species  studied  the  inflorescence  is  cymose.  A 
schematic  representation  of  this  type  of  inflorescence  as  exhibited  by  most  of 
the  annual  species  of  Drymaria  is  shown  in  PI.  20,  fig.  1,  and  a  diagram  of 
the  inflorescence  of  D.  sperguloides  is  shown  in  fig.  2.  In  the  latter  figure,  the 
small,  solidly  inked  circles  represent  abortive  flowers. 

Floral  characters  useful  in  differentiating  species  of  Drymaria  include  the 
shape,  length,  comparative  length  and  width,  venation,  and  texture  of  the 
sepals;  the  shape,  depth  of  the  lobing,  length  as  compared  with  the  length  of 
other  floral  structures,  and  the  presence  or  absence  of  appendages  between 
the  main  lobes  of  the  petals ;  and  the  comparative  length  of  claw  and  blade 


Vol.  XXV]        If'IGGIXS:  DEYMAEIA  IX  SONURAN  DESERT  191 

of  the  petals.  Flowers  of  comparable  age  must  be  selected  in  making  these 
comparisons,  however,  for  the  petals  are  persistent  and  the  mature  capsules 
are  usually  much  longer  than  the  ovary  at  anthesis. 

The  aestivation  of  the  sepals  is,  in  all  cases  examined,  cochlear,  a  modifica- 
tion of  the  imbricate  pattern.  In  this  type  of  sepal  arrangement  one  sepal 
is  wholly  outside  the  others,  one  has  both  margins  overlapped  by  adjacent 
sepals,  and  of  the  other  three  each  has  one  margin  overlapped  by  an  adjacent 
sepal  and  the  other  margin  overlapping  that  of  its  neighbor.  The  outermost, 
exposed  sepal  is  on  the  abaxial  side  of  the  calyx,  while  the  one  with  both  mar- 
gins covered  is  on  or  near  the  adaxial  side.  The  margins  that  are  covered  by 
the  edges  of  adjacent  sepals  are  almost  always  more  broadly  scarious-winged 
than  the  exposed  margins  (PL  20,  fig.  3).  This  does  not  hold  in  the  case  of 
D.  carinaia  and  its  variety,  for  in  these  plants  the  thicker,  herbaceous  part 
of  the  sepals  is  reduced  to  a  narrow  midrib  and  the  sepals  are  nearly  or  quite 
symmetrical. 

All  species  studied  have  stamens  that  are  slightly  dilated  at  the  base  to  form 
a  shallow,  connate  cup. 

It  is  doubtful  that  there  is  any  value  in  trying  to  differentiate  between 
species  with  "stipitate"  and  "sessile"  ovaries,  for  all  species  examined  had 
the  ovaries  elevated  to  some  degree  on  a  slender,  short  stipe.  Though  short, 
this  stipe  was  distinctly  demonstrable  in  every  case. 

The  seeds  are  so  small  that  definitive  measurements  are  difficult  to  make. 
Differences  in  the  depth  of  the  notch  adjacent  to  the  hilum,  the  sculpturing 
of  the  surface,  the  over-all  pattern  of  the  surface  markings,  and  sometimes 
the  color  of  the  seeds  can  be  used  to  advantage.  The  seeds  of  D.  viscosa  are 
striking  in  that  they  are  smooth  and  pale  cream  or  pale  buff  instead  of  tes- 
sellated, muriculate,  or  tuberculate  and  dark  colored  as  the  seeds  of  most  other 
species  are. 

During  the  course  of  this  study  the  material  of  Drymaria  at  the  California 
Academy  of  Sciences,  in  the  herbarium  of  the  University  of  California,  and 
in  the  Dudley  Herbarium  at  Stanford  University  has  been  available  con- 
stantly. The  types  of  a  number  of  species  were  studied,  notes  taken,  and  photo- 
graphs of  them  made  during  a  visit  to  the  herbarium  of  the  Royal  Botanic 
Garden  at  Kew,  Surrey,  and  to  the  Gray  Herbarium  of  Harvard  University 
in  1937.  These  notes  and  photographs  were  valuable  aids  in  determining 
specific  limits.  The  type  specimens  of  Brandegee's  species,  deposited  in  the 
herbarium  of  the  University  of  California,  and  an  isotype  of  D.  depressa 
Greene,  in  the  herbarium  of  the  California  Academy  of  Sciences,  clarified  the 
status  of  several  poorly  known  species.  My  sincere  thanks  are  due,  and  gladly 
given,  to  Miss  Alice  Eastwood  for  permitting  me  to  borrow  material  from  the 
herbarium  of  the  California  Academy  of  Sciences.  Appreciation  is  hereby 
expressed,  also,  to  Dr.  Herbert  L.  Mason,  of  the  University  of  California ; 
to  Mr.  A.  D.  Cotton,  Keeper  of  the  Herbarium  of  the  Royal  Botanic  Garden, 
Kew,  England;  and  to  Dr.  M.  L.  Fernald,  of  the  Gi-av  Hei-barinm  at  Harvard 


192  CALIFOENIA  ACADEMY  OF  SCIENCES        [Proc.  4th  ser. 

University,  for  the  privilege  of  studying-  material  and  types  deposited  in  the 
herbaria  under  their  supervision. 

In  citing  specimens,  the  herbaria  in  which  the  specimens  are  deposited  are 
indicated  by  the  following  abbreviations  : 

California  Academy  of  Sciences CA 

Dudley  Herbarium,  Stanford  University D 

Gray  Herbarium,  Harvard  University G 

Royal  Botanic  Garden,  Kew,  England K 

University  of  California  at  Berkeley UC 

Taxonomic  Treatment 
Drymaria  Willd.  in  R.  and  S. 

S.yst.Veg.  5:406,  1819. 

Annual  or  perennial  plants  wdth  slender,  prostrate,  decumbent,  ascending, 
erect  or  scrambling  stems.  Branches  simple  or  profusely  branched,  sparsely 
to  densely  foliate.  Leaves  linear,  subulate,  elliptic,  ovate,  or  cordate,  thin  and 
herbaceous  or  thick  and  fleshy  or  leathery,  glabrous  or  pubescent,  glandular 
or  egiandular ;  stipules  usually  present,  subulate  or  linear,  persistent  or  fuga- 
cious. Flowers  small,  in  cymes  or  racemes,  sessile  or  pedicellate;  sepals  5, 
1-3-nerved,  similar  or  slightly  dissimilar,  the  middle  portion  herbaceous  and 
often  puberulent  or  grandular,  one  or  both  of  the  margins  scarious-winged ; 
petals  (in  ours)  deeply  cleft,  in  some  species  bearing  additional  narrow  ap- 
pendages near  base  of  the  cleft;  stamens  5,  the  filaments  slightly  dilated  and 
connate  at  the  base ;  anthers  ovate,  much  shorter  than  the  filaments.  Style 
lender,  from  much  shorter  than,  to  about  equal  to,  the  ovary;  stigma  3-lobed 
or  3-cleft,  the  lobes  shorter  or  longer  than  the  style.  Capsule  ovoid  to  sub- 
globose,  longitudinally  dehiscent,  mostly  3-valved,  1-celled.  Seeds  obovoid 
to  obovoid-reniform,  variously  sculptured,  most  dark  brown,  dull  or  slightly 
shining. 

Type  species. — Holosteum  cordafian  Linn. 

Key  to  the  Species 
Leaves  about  as  broad  as,  or  broader  than  long. 

Flowers  in  short,  crowded,  nmbel-liko  rncenies;  leaves  leathery  or  fleshy,  often  glau- 
cous; plants  mostly  prostrate  or  decumbent. 
Stamens  equaling  or  surpassing  the  ovary  at  anthesis;  style  one-fourth  as  long 
as,  or  equaling,  the  ovary ;  petals  with  4-R  appendages  near  base  of  inner 
margins  of  main  lobes. 
Herbage   somewhat   puberulent,   scarcely  glaucous;    petals   3-3.5   mm.   long, 
about  equaling  the  sepals;   style  1.4—1.6  mm.  long;   internodes  ex- 
ceeding  the   leaves 1.7).  holosteoides. 

Herbage  glabrous,  conspicuously  glaucous;   petals  2-2.2  mm.  long,  shorter 
than  the  sepals;  style  0.6-0.8  mm.  long;  internodes  mostly  shorter 

than  the  leaves 2.  D.  eras.ii folia. 

Stamens  about  half  as  long  as  the  ovary  at  anthesis;  style  less  than  one-fifth  as 
long  as  the  ovary;  petals  with  only  one  nppendage  at  base  of  each  mnin 
lobe 3.  Z>.  pachiiplnjlla. 


Vol.  XXV]       WIGGINS:  DRTMARIA  IN  HONORAN  DESERT  193 

Fln^vers  in  cymes;  leaves  thin,  not  fleshy  nor  leathery,  rarely  glaucous;  plants  mainly 
erect  or  ascending. 
Sepals  strongly  carinate,  th£  herbaceous  mid-portion  narrowly  linear. 

Claws  of  petals  half  as  long  as,  and  gradually  widening  to,  the  blade;  petals 

slightly  exceeding  the  sepals;  plant  annual 4.  D.  carinata. 

Claws  of  petals  very  short,  less  than  one-sixth  as  long  as,  and  flaring  abruptly 
to,  the  limb;  petals  often  twice  as  long  as  the  sepals;  plants  peren- 
nial  4a.  D.  carinata  var.  perennis. 

Sepals  not  carinate,  subequal,  the  herbaceous  mid-portion  elliptic. 
Petals  surpassing  the  sepals;  sepals  not  strongly  ribbed. 

Sepals  oblong-elliptic  or  narrowly  oblanceolate,  obtuse  at  the  apex;  petals 
5-6.5  mm.  long;  pubescence  of  short  hairs  0.4  mm.  long  or  less. 
Plants  perennial ;  leaves  shallowly  cordate ;  sepals  glabrous  or  spar- 
ingly hairy  near  base 5.  D.  polystachya. 

Plants  annual ;  leaves  truncate  or  broadly  cuneate  at  the  base ;  sepals 
copiously  glandular-puberulent  on  lower  two-thirds. 

5a.  D.  polystachya  var.  diffusa. 
Sepals  broadly  ovate,  acute  at  the  apex;  petals  3.8-4.5  mm.  long;  pubes- 
cence of  crinkled,  spreading  hairs  to  1  mm.  long .  .  .6.  D.  dehilis. 

Petals  shorter  than  the  sepals;  sepals  strongly  3-ribbed l.B.  Fendleri. 

Leaves  linear  or  narrowly  oblong,  several  times  as  long  as  broad. 

Herbage  heavily  viscid-puberulent  throughout;  seeds  smooth,  not  tessellate  nor  muricu- 

late 8.D.  viscosa. 

Herbage  viscid-puberulent  only  on  pedicels  and  sepals,  or  glabrous;  seeds  tessellate, 
muriculate,  or  otherwise  roughened. 
Petals  fimbriate-appendaged. 

Herbaceous  part  of  sepals  obtuse,  glabrous;  stigmas  subsessile;  petals  with 
1  lobe  at  base  of  each  main  lobe ;  leaves  pseudo-verticillate. 

9.  D.  sperguloides. 
Herbaceous  part  of  sepals  acute,  glandular-puberulent;  stigmas  not  subses- 
sile; petals  with  2-4  appendages  at  base  of  each  main  lobe;  leaves 
not  pseudo-verticillate. 
Petals  6  mm.  long,  much  exceeding  the  sepals ;  appendages  long,  slender, 
nearly  equaling  the  main  lobes ;  leaves  narrowly  elliptic. 

10.  D.  arenarioides. 
Petals  4  mm.  long,  barely  exceeding  the  sepals ;  appendages  oblong,  half 

to  two-thirds  as  long  as  the  main  lobes;  leaves  narrowly  linear. 

11.  D.  peninsidaris. 
Petals  merely  bifid,  without  appendages  along  inner  margins  of  main  lobes. 

Petals  slightly  exceeding  the  sepals ;  plants  puberulent. 

Herbaceous  part  of  sepals  ol:)tuse ;  leaves  mostly  5  mm.  long  or  less ;  claws 

of  petals  narrow,  naked 12.  D.  Johnstonii 

Herbaceous  part  of  sepals  acute;  leaves  mostly  over  10  mm.  long;  claws 

of  petals  broadly  winged,  as  wide  as  the  limb 13.  D.  effusa. 

Petals  shorter  than  the  sepals;  plants  glabrous,  or  base  of  sepals  and  upper 
parts  of  internodes  sparsely  puberulent. 
Stems  erect,  branching  above;  leaves  narrowly  linear,  1  mm.  wide  or  less; 
seeds  minutely  but  sharply  muriculate. 

Sepals  acute  ;  herbage  glabrous 14.  D.  tenella. 

Sej)als  acuminate ;   upper  parts  of  internodes  and  bases  of  sepals 

puberulent    14a.  D.  tenella  var.  nodosa. 

Stems  decumbent  or  spreading,  branched  from  the  base,  leaves  oblong,  to 
3  mm.  wide;  seeds  tessellate  with  low,  rounded  bosses. 

15.  D.  depressa. 


19-4  CALIFORNIA  ACADEMY  OF  SCIENCES        [Proc. -Ith  ser. 

1.  Drymaria  holosteoides  Benth.  Bot.  Voy.  Sulph.  16,  1844 

(Plate20,  figs.  4-12) 

Drymaria  Featchii  Curran,  Proc.  Calif.  Acad.  Sci.  (II),  1:227, 1888. 
Mollugophytum  holosteoides.  M.  E.  Jones,  Extract  from  Contr.  West.  Bot.  18 :35, 1933. 

Annual,  with  slender,  spreading,  decumbent  or  prostrate  stems  4-40  em. 
long,  some  of  them  often  stolon-like  and  forming  secondary  rosettes  at  rooted 
nodes,  internodes  2-10  cm.  long,  mostly  conspicuously  exceeding  the  leaves, 
spreadingl}^  puberulent  and  somewhat  glandular,  older  stems  glabrate ;  stip- 
ules lacking  or  represented  by  narrow  scarious  wing  between  bases  of  a  leaf- 
pair  ;  leaves  mostly  in  fascicles  of  4-6  at  each  node,  blades  spatulate-elliptic, 
3-7  mm.  wide,  4—12  mm.  long,  somewhat  fleshy  or  leathery,  faintly  3-nerved 
beneath,  glabrous  or  sparsely  puberulent  near  the  base,  lightly  glaucous,  on 
slender  petioles  2-10  mm.  long;  flowers  in  dense,  short  racemes  in  axils  of 
most  leaves,  each  flower  subtended  by  a  narrow,  ovate,  hyaline  bract  about 
1  mm.  long;  pedicels  slender,  3-9  mm.  long,  glandular-puberulent;  sepals  obo- 
vate,  obtuse  or  rounded  at  the  apex,  3-3.5  mm.  long,  the  middle  portion 
herbaceous,  glandular-puberulent  at  the  base,  glabrous  and  faintly  glaucous 
above,  the  scarious  margins  conspicuous;  petals  persistent,  about  equaling  the 
sepals,  2.8-3.2  mm.  long,  white,  bifid  about  half  way  to  the  base  and  bearing 
2—4  narrow  segments  near  the  base  of  inner  margin  of  each  main  lobe,  the  claw 
short,  broad,  somewhat  erosulate ;  stamens  about  equaling  the  petals,  nearly 
twice  as  long  as  the  ovary ;  filaments  slightly  dilated  at  the  base ;  ovary  about 
1.5  mm.  high;  style  equaling  the  ovary;  capsule  broadly  ovoid,  1.5-2  mm. 
high ;  seeds  obovoid-reniform,  0.6-0.8  mm.  long,  dark  brown  to  black,  dull, 
minutely  rounded-tuberculate  in  longitudinal  lines. 

Type  locality  :—"CsLpe  San  Lucas,"  Baja  California,  Mexico. 

Distribution: — Sandy  areas  from  the  vicinity  of  Calmalli  to  La  Paz,  and 

possibly  to  Cape  San  Lucas;  Cedros  Island;  Magdalena  Island;  Tiburon 

Island,  Mexico. 

Specimens  examined: — Magdalena  Bay,  Hinds,  1841,  (K,  type.  This  is  the  only  specimen 
of  D.  holosteoides  in  the  Kew  Herbarium  collected  on  the  Voyage  of  the  "Sulphur,"  and  in 
spite  of  the  statement  in  Bentham's  publication  to  the  effect  that  the  plant  was  obtained  at 
"Cape  San  Lucas"  the  type  locality  is  probably  Magdalena  Bay,  on  Magdalena  Island). 
Madgalena  Bay,  Brandegee,  March  1889,  (UC)  ;  Bryant,  1888,  (UC,  type  of  D.  Featchii) ; 
hills  east  of  Calmalli,  Purpus  151,  (D,  UC)  ;  rocky  hillside  4  miles  east  of  San  Ignacio, 
Wiggins  7898,  (D,  UC) ;  2  miles  west  of  San  Ignacio,  Eeed  6156,  (D) ;  Coyote  Bay,  Con- 
cepcion  Bay,  Johnston  4178,  (CA)  ;  Gentry  4059,  (D)  ;  Concepcion  Bay,  Shreve  7095.  (J))  ; 
5  miles  south  of  head  of  Concepcion  Bay,  Wiggins  5456,  (CA,  D,  UC)  ;  Mulege,  Johnston 
3690,  (CA)  ;  San  Luis  Gonzales  Bay,  Johnston  3329,  (CA)  ;  San  Francisco  Island,  John- 
ston 2949,  (CA) ;  dry  washes  between  Medano  and  Venancio,  Wiggins  5531,  (D,  UC) ;  La 
Paz,  Johnston  3048,  (CA,  UC)  ;  Brandegee,  Feb.  2,  1890,  (UC)  ;  Cedros  Island,  Stewart 
48-49,  (CA)  ;  Willard's  Point,  Tiburon  Island,  Sonora,  Johmton  42G3,  (CA,  D,  UC). 

A  long-standing  confusion  concerning  the  identity  of  this  species  and  the 
closely  related  1).  crassifolia  was  cleared  up  by  Brandegee  (Zoe  2:  66-70. 
1891). 


Vol.  XXV]       WIGGINS:  DRYMABIA  IN  SONOBAN  DESEBT  195 

The  petals  in  D.  holosteoides  are  more  pronouncedly  fimbriate  at  the  base 
of  the  sinus  than  they  are  in  D.  crassifolia,  and  the  style  is  nearly  or  quite 
as  long  as  the  ovary  in  holosteoides,  but  only  half  as  long  in  D.  crassifolia. 
The  densely  matted  habit  of  crassifolia  is  the  most  readily  observed  character 
for  separating  the  two  species. 

2.  Drymaria  crassifolia  Benth.  Bot.  Voy.  Sulph.  16,  184-1 

(Plate20,  figs.  20-26) 
Mollugophyium  crassifolium  M.  E.  Jones,  Extract  from  Contr.  West.  Bot.  18 :35.  1933. 

Prostrate,  matted,  glabrous  and  glaucous  perennial ;  stems  profusely 
branched,  3-20  cm.  long,  internodes  0.5-2.5  cm.  long,  mostly  obscured  by  the 
numerous,  fasciculate  leaves;  stipules  as  in  D.  holosteoides;  petioles  slender, 
3-9  mm.  long;  leaf-blades  broadly  ovate,  broadly  rhombic-spatulate,  or  ellip- 
tic, mostly  rounded  at  the  apex  and  broadly  cuneate  at  the  base,  2.5-8  mm. 
broad,  4-12  mm.  long,  fleshy,  glaucous,  indistinctly  3-nerved  beneath;  flowers 
numerous  in  short  cluster-like  racemes,  pedicels  5-12  mm.  long ;  sepals  slightly 
dissimilar,  broadly  elliptic,  rounded,  acutish,  or  faintly  emarginate  at  the 
tip,  1.4—1.8  mm.  wide,  3-3.4  mm.  long,  fleshy,  glaucous,  faintly  veined ;  petals 
2-3  mm.  long,  bifid  about  to  middle,  usually  with  2  short,  rounded  or  truncate 
appendages  at  base  of  inner  margin  of  each  main  lobe;  stamens  1.8-2  mm. 
long,  about  equaling  the  ovary;  style  0.4—0.6  mm.  long,  about  one-fourth  as 
long  as  the  ovary,  the  stigma-lobes  minute ;  capsule  ovoid,  about  equaling  the 
sepals ;  seeds  about  0.8  mm.  long,  obovoid-renif orm,  minutely  reticulate  in 
longitudinal  lines,  black,  dull. 

Tyi)e  locality: — Cape  San  Lucas,  Baja  California,  Mexico. 

Distribution : — Sandy  soil  and  along  washes.  Cape  Region,  Baja  California. 

Specimens  examined: — Cape  San  Lucas,  Hinds,  1841,  (K,  type)  ;  Cabo  San  Lucas, 
Shreve  7258,  (D)  ;  on  beach  sands,  between *the  town  of  Cabo  San  Lucas  and  the  end  of 
the  peninsula.  Whitehead  906,  (D)  ;  seashore,  Todos  Santos,  M.  E.  Jones  27046,  (D,  UC)  ; 
San  Jose  del  Cabo,  Purpus  265,  (UC)  ;  Anthony,  Mar.-June,  1897,  (TJC)  ;  Brandegee,  Oct. 
7,  1890,  and  Nov.,  1902,  (UC). 

In  commenting  on  D.  holosteoides,  Johnston  wrote  (Proc.  Calif.  Acad.  Sci. 
IV.  12  :  1024.  1924),  "Drymaria  crassifolia  .  .  .  is  a  very  closely  related  form 
known  only  from  San  Jose  del  Cabo,  and  with  little  more  than  its  perennial 
habit  to  distinguish  it."  It  seems  to  me  that  the  heavily  glaucous  herbage,  the 
shorter  stamens,  the  shorter  petals  with  less  strongly  fimbriate  appendages, 
the  shorter  internodes  and  matted  habit,  are  characters  of  sufficient  magni- 
tude to  warrant  retaining  D.  crassifolia  as  a  distinct  species.  Further  field 
work  may  bring  to  light  additional  localities  in  which  it  grows. 

3.  Drymaria  pachyphylla  Wooton  and  Standley,  Contr.  U.  S. 
Nat.  Herb.  16: 121, 1913 
(Plate  20,  figs.  13-19) 
Annual,  with  slender,  spreading,  prostrate  stems  3-15  cm.  long,  the  inter- 
nodes 2—12  cm.  long,  much  surpassing  the  leaves,  which  often  are  confined 
to  terminal  tufts ;  leaves  mostly  pseudoverticillate  in  4's,  the  bases  of  each 


196  CALIFORNIA  ACADEMY  OF  SCIENCES        [Pkoc.  4th  sek. 

pair  connate  in  a  narrow,  scarious  membrane  ;  petioles  slender,  5-7  mm.  long, 
blades  broadly  elliptic,  rhombic-ovate,  or  broadly  ovate,  3-11  mm.  wide, 
5-14  mm.  lony,  somewhat  fleshy,  green  or  faintly  glaucous,  indistinctly  3- 
nerved ;  flowers  in  small,  racemose  clusters,  about  3-15  flowers  on  each  short 
rachis,  each  subtended  by  a  narrowly  ovate,  hyaline  bract  1.5-2  mm.  long; 
pedicels  1-4  mm.  long;  sepals  broadly  ovate,  1.5-2  mm.  wide,  2.5-3  mm.  long, 
rounded  at  the  apex,  green  and  herbaceous,  with  membranous,  white  margins 
about  0.2  mm.  wide ;  petals  about  equaling  or  slightly  shorter  than  the  sepals, 
bifid  one-third  of  way  to  base,  each  lobe  bearing  a  broad,  blunt  appendage 
half  as  long  as  itself  near  the  base  of  the  inner  margin ;  stamens  1-1.2  mm. 
long,  half  as  long  as  the  ovary;  style  0.2  mm.  long,  the  stigmas  scarcely 
longer;  capsule  broadly  ovoid  to  subglobose,  about  equaling  the  sepals,  yel- 
lowish to  pale  brown;  seeds  0.6-0.8  mm.  hmg,  minutely  granular-tessellate. 
dark  brown,  dull. 

Type  locality: — On  the  plains  south  of  the  White  Sands,  Dona  Ana  County, 
New  Mexico. 

Distribution: — Sandy  soil,  southeastern  Arizona,  through  New  Mexico  to 
western  Texas  and  in  adjacent  Mexico. 

Specimens  examined: — Arizona:  Chiricaliua  Mts.,  M.  E.  Jones,  Sept.  22,  1931,  (D,  UC). 
New  Mexico:  plains  south  of  the  White  Sands,  Dona  Ana  Co.,  Wooion  405,  (D,  UC,  iso- 
types).  Mexico:  Torreon,  Coahuila,  Furpus  472,  1903,  (UC)  ;  Filipinas,  Coahuila,  Furpus 
4942,  (UC). 

This  is  the  species  that  Sereno  Watson  (Proc.  Am.  Acad.  17:  329.  1882) 

listed  from  Coahuila  as  Z>.  crassifolia. 

4.  Drymaria  carinata  Brandegee,  Zoe  2:70,  1891 
(PIate20,  figs.  27-31) 

Annual,  with  erect  or  ascending,  weak,  slender  stems  and  glabrous  herbage 
(sparsely  puberulent  in  the  variety)  ;  stipules  narrowly  subulate,  2-4  mm. 
long,  white,  subpersistent ;  leaves  broadly  ovate,  the  blades  14^16  mm.  long, 
5-18  mm.  wide,  apiculate,  from  broadly  cuneate  to  shallowly  cordate  at  the 
base,  thin,  finely  but  distinctly  3-veined,  bright  green  above,  slightly  paler 
beneath;  petioles  half  as  long  as  the  blades  to  equaling  them;  inflorescence 
loosely  cymose ;  peduncles  and  pedicels  filiform ;  pedicels  2-8  mm.  long,  each 
subtended  by  an  ovate,  acute,  scarious  bract  1-1.5  mm.  long;  sepals  broadly 
ovate-lanceolate,  strongly  incurved,  bluntly  carinate,  2-2.5  mm.  long,  the 
green  midvein  narrow;  petals  1.5-2.5  mm.  long,  bifid  two-thirds  of  way  to 
base,  the  lobes  laiu-eolate  to  narrowly  spatulate,  entire  or  faintly  emar- 
ginate,  devoid  o!  marginal  appendages;  the  claw  narrow;  stamens  about 
equaling  or  slightly  exceeding  the  sepals,  filaments  linear;  ovary  1-1.2  mm. 
high,  short-stipitate;  capsule  subglobose,  about  equaling  the  sepals;  seeds 
obovate-reniform,  0.4-0.6  mm.  long,  deep  amber  to  light  brown,  minutely 
tessellated. 

Type  locality: — Sieri-a  de  la  Laguna,  Baja  California,  Mexico. 

Disfrihufioii : — "Mountnins  of  southoi-n  l^>a.ia  California. 


Vol.  XXV]       WIGGINS:  DBYMAEIA  IN  SONOEAN  DESERT  197 

Specimens  examined: — Sierra  de  la  Laguna,  Brandegee  34,  Jan.  21,  1890,  (UC,  type); 
Laguiia  Mts.,  M.  E.  Jones  24193,  (CA)  ;  Sierra  de  la  Trinidad,  Brandegee,  Nov.  1902, 
(UC)  ;  Miraflores,  Brandegee,  Oct.  14,  1890,  (UC)  ;  El  Taste,  Brandegee,  Sept.  14,  1893, 
(UC)  ;  San  Felipe,  Purpus  488,  1901,  (UC)  ;  under  overhanging  rocks,  sides  and  bottoms 
(if  wash  22  miles  south  of  La  Paz,  Wiggins  5610,  (D). 

4a.  Drymaria  carinata  Brand,  var.  perennis,  Wiggins,  var.  nov. 

(Plate  20,  figs.  32-34) 

Ab  D.  carinata  typiea  radicibus  perennibus,  caulibus  basim  lignosis,  nodis 
sparse  minutissimeque  scaberulosis,  lobis  petalorum  latis  ciirtisque  differt. 

Type  specimen,  deposited  in  the  Dudley  Herbarium,  Stanford  University, 
no.  263280,  was  collected  in  humus  on  cliff  of  rocky  canyon  slope,  in  oak-pine 
forest.  La  Laguna,  Sierra  de  la  Laguna,  alt.  6000  feet,  Baja  California, 
Mexico,  H.  S.  Geniry  4415,  March  25,  1939. 

Additional  specimen  examined. — Sierra  de  San  Francisquito,  Brandegee, 
March  30, 1892,  (UC). 

There  is  a  elearcut  and  constant  difference  between  the  shape  and  size  of 
the  petals  of  the  variety  perennis  and  of  those  of  the  typical  annual  form 
(PL  20,  figs.  30,  32-34).  The  lobes  of  the  former  are  distinctly  broader  in 
comparison  with  the  length,  are  practically  the  same  width  from  base  to  tip, 
and  are  more  nearly  parallel  than  are  those  of  D.  carinata.  In  perennis,  the 
claw  is  shorter  and  broader,  and  flares  more  abruptly  to  the  blade  than  in 
the  typical  material. 

5.  Drymaria  polystachya  Brandegee,  Zoe  2:70,  1891 

(Plate21,  figs.  2-5) 

Perennial  from  a  gnarled  or  tortuous,  woody  root ;  stems  slender,  spread- 
ing to  ascending,  1-3  dm.  long,  freely  branched,  sparingly  to  moderately 
pilosulous  or  glabrate;  herbage  pubescent  and  slightly  glandular;  stipules 
linear-subulate,  1-2  mm.  long;  leaf -blades  broadly  cordate,  6-20  mm.  wide, 
6-15  mm.  long,  the  apex  abruptly  short-acuminate  or  acute,  subsessile  or  on 
petioles  2-7  mm.  long;  inflorescence  of  terminal  and  subterminal  cymules; 
peduncles  slender,  1-8  cm.  long,  a  pair  of  narrowly  ovate,  scarious  bracts 
1.6-2.2  mm.  long  at  each  node  of  the  inflorescence;  pedicels  subfiliform, 
0.6-2  cm.  long ;  sepals  subequal,  oblong-elliptic,  1.5-1.7  mm.  wide,  4.5-5.5  mm. 
long,  obtuse  or  rounded  and  erosulate  at  the  apex,  the  broad  greenish  central 
portion  glabrous  or  sparingly  puberulent,  faintly  3-nerved,  the  margins 
scarious;  petals  bifid  about  to  the  middle,  5.5-6.5  mm.  long,  devoid  of  append- 
ages, the  claws  narrow,  about  2  mm.  long,  naked ;  stamens  about  3  mm.  long ; 
ovary  ovoid,  nearly  equaling  the  stamens;  capsule  3.5-^.5  mm.  long;  seeds 
obovoid-reniform,  slightly  compressed  laterally,  brown,  finely  tessellated  in 
numerous  longitudinal  rows. 

Type  locality: — Cliffs  near  San  Jose  del  Cabo,  Baja  California,  Mexico. 

Distribution: — Among  rocks  and  on  cliffs,  southern  Baja  California. 

Specimens  examined: — San  Jose  del  Cabo,  Brandegee  35,  Mar.  14,  1890,  (UC,  type,  D, 


I 

198  CALIFORNIA  ACADEMY  OF  SCIENCES        [Pkoc.  4th  ser.  ; 

isotype)  ;  Brandegee,  March  lU,  1892,  (UC)  ;   Grabendorfer,  March,  1899,  (UC)  ;  Sieira 
de  la  Trinidad,  Brandegee,  Nov.,  1902,  (UC). 

Brandegee  described  the  leaves  as  "...  1  em.  or  less  broad,  ^2  cm.  long,  on  i 
petioles  of  nearly  the  same  length."  However,  most  of  the  leaves  are  sub- 
sessile  or  on  petioles  only  2-3  mm.  long. 

The  date  on  the  type  in  the  herbarium  of  the  University  of  California  reads 
"March  14,  1890,"  and  the  label  bears  the  number  "35."  A  specimen  in  the 
Dudley  Herbarium  also  bears  the  number  "35,"  but  is  dated,  "March  24, 
1890."  It  is  not  clear  whether  the  discrepanc}'  in  the  dates  on  the  two  speci- 
mens is  a  clerical  error  made  in  copying  the  labels,  or  two  separate  collections 
are  involved.  I  am  inclined  toward  the  former  explanation.  No  mention  of 
the  habitat  is  made  on  either  label,  so  one  must  rely  solely  upon  Brandegee's 
statement  in  the  original  publication  for  the  information  that  the  plant  grew 
on  "cliffs"  near  San  Jose  del  Cabo. 

5a.  Drymaria  polystachya  Brand,  var.  diffusa  (Rose)  Wiggins,  comb.  nov. 

(Plate  21,  figs.  6-8) 
Drymaria  diffusa  Eose,  Contr.  U.  S.  Nat.  Herb.  1 :130.  pi.  12, 1892. 

Annual,  with  weak,  spreadingiy-ascending  stems,  giandular-puberulent 
throughout;  leaves  with  very  slender  petioles  4-10  mm.  long,  the  blades 
3-10  mm.  wide,  3-8  mm.  long,  truncate  or  broadly  cuneate  at  the  base ;  sepals 
narrowly  oblanceolate,  about  3.5  mm.  long;  petals  4.5-5  mm.  long;  otherwise 
as  in  the  typical  form  of  the  species. 

Type  locality: — Carmen  Island,  Baja  California,  Mexico. 

Distrihiitio7i: — Known  only  from  Carmen  Island  and  the  mainland  of  Baja 
California  immediately  adjacent. 

Specimens  examined: — Carmen  Island,  Palmer  819,  Nov.  1-7,  1890,  (UC,  2  isotypes) ; 
shady  talus  slope  under  basaltic  cliffs,  above  Punta  Escondido,  Sierra  Giganta,  alt.  2000 
to  3000  feet,  Gentry  3751,  (UC), 

6.  Drymaria  debilis  Brandegee,  Proc.  Calif.  Acad.  Sci.  II.  2  :131,  1889 

(Plate21,  figs.  9,  10) 

Annual ;  stems  subfiliform,  weak,  spreading-ascending  or  scandent,  1-2.5 
dm.  long,  sparingly  pilosulous,  the  internodes  3—4.5  cm.  long;  stipules  linear- 
subulate,  somewhat  tortuous,  1-1.5  mm.  long ;  petioles  slender,  2-10  mm.  long, 
pilosulous;  leaf -blades  broadly  ovate,  3-10  mm.  long  and  wide,  acute  or 
abruptly  short-acuminate  at  the  apex,  abruptly  and  broadly  cuneate  or  trun- 
cate at  the  base,  sparingly  pilosulous  on  both  sides  or  glabrate  above ;  in- 
florescence loosely  cymose ;  pedicels  filiform,  4—14  mm.  long,  sparsely  pubes- 
cent; sepals  2.6-3.2  mm.  long,  pilosulous,  ovate-acuminate,  or  the  scarious 
wings  obtusish  at  the  apex,  faintly  3-nerved;  petals  3.8-4.2  mm.  long,  bifid 
about  to  the  middle,  the  lobes  broadly  spatulate,  the  claws  narrow  and  naked; 
stamens  about  equaling  the  sepals,  the  filamentas  linear,  flattened;  capsule 
ovoid,  nearly  equaling  the  sepals ;  seeds  reddish  brown,  obovoid-reniform.  fi 
0.6-0.8  mm.  long,  minutely  tessellate  in  fine,  longitudinal  lines. 


Vol.  XXV]       WIGGINS:  DEYMAEIA  IN  SONOBAN  DESERT  199 

Type  locality:  Piirisima,  Baja  California,  Mexico. 
Disirihution: — Known  only  from  the  type  locality. 
Specimens  examined: — Purisima,  Brandegee,  Feb.  13,  1889,  (UC,  type). 

In  general  appearance  D.  debilis  closely  resembles  D.  carinata,  but  differs 
from  that  species  in  having  acute,  broader  sepals  that  are  not  at  all  carinate ; 
more  pronouncedly  pilosulous  herbage,  and  somewhat  shorter  petals.  The 
crinkled  hairs  of  the  pubescence  are  often  1-1.2  mm.  long,  but  there  are  very 
few  glandular  hairs  mixed  with  the  non-glandular  ones.  The  paucity  of  glands 
helps  to  separate  D.  dehilis  from  D.  polysfachya  and  its  variety,  diffusa. 

7.  Drymaria  Fendleri  S.  Wats.  Proc.  Am.  Acad.  17:328,  1882 

(Plate21,  figs.  11-19) 
Drymaria  Fendleri  var.  perennis  M.  E.  Jones,  Extract  from  Contr.  West.  Bot.  18:65.  1933. 

Erect  or  ascending  annual  (rarely  biennial  or  perennial  under  favorable 
conditions)  with  glandular-puberulent  herbage;  stems  slender,  1-3.5  dm. 
long ;  internodes  to  12  cm.  long ;  stipules  bifid  into  linear-subulate  lobes  1.5-2 
j  mm.  long,  persistent;  leaves  thin,  broadly  ovate  or  ovate-cordate,  4r-15  mm. 
long.  6-22  mm.  wide,  acute  or  rounded  but  abruptly  apiculate  at  the  apex, 
truncate  or  shallowly  cordate  at  the  base,  on  slender  petioles  2-7  mm.  long, 
the  margins  often  crinkling  on  drying ;  inflorescence  cymose ;  peduncles 
slender,  1-6  cm.  long,  usually  strongly  glandular-puberulent,  bearing  a  pair 
of  ovate,  scarious  bracts  2-2.5  mm.  long  at  the  node ;  pedicels  slender,  1-2  mm. 
long,  or  some  flowers  subsessile  in  2-5-flowered  clusters ;  sepals  lanceolate, 
1-1.4  mm.  wide,  4-5  mm.  long,  strongly  3-ribbed,  acute  to  acuminate,  the  cen- 
tral portion  green,  glandular-puberulent,  marginal  wings  scarious ;  petals 
3.5-4  mm.  long,  bifid  about  to  the  middle,  the  lobes  linear-spatulate,  claws 
narrow  and  naked;  stamens  half  as  long  as  the  petals;  capsule  ovoid,  2-2.5 mm. 
high,  8-15-seeded;  seeds  obovoid-reniform,  0.6-0.8  mm.  long,  reddish  brown, 
minutely  tessellated  in  longitudinal  rows. 

Ty2:)e  locality: — "New  INIexico." 

Bistrihntion: — Foothills  and  mountains,  New  Mexico  and  Arizona  to  cen- 
tral Mexico,  and  the  Cape  Region  of  Baja  California. 

Specimens  examined: — Ne-\v  Mexico:  without  further  locality,  Fendler  60,  1847,  (G, 
type)  ;  Thurber  1073,  (G)  ;  C.  Wright  866,  1851,  (G)  ;  La  Cuesta,  Bigelow,  1853,  (G)  ;  (all 
of  the  foregoing  specimens  are  mounted  on  a  single  herbarium  sheet,  and  all  were  cited  by 
Watson  in  his  original  description)  ;  Silver  City,  E.  L.  Greene,  Sept.  14,  1880,  (UC)  ;  granite 
hills,  Kingston,  Sierra  County,  Metcalfe  1378,  (CA)  ;  under  junipers,  mountain  slope  near 
San  Jose,  San  Miguel  County,  Barnehy  2627,  (CA).  Arizona:  amid  big  boulders,  Prescott, 
Eastwood  16825,  (CA)  ;  near  Prescott,  Eusche,  Sept.  1-5,  1929,  (CA).  Baja  California, 
Mexico :  Cota  Eanch,  Laguna  Mts.,  14  miles  E.  of  Todos  Santos,  M.  E.  Jones  27050,  (D,  UC, 
isotypes  of  var.  perennis  Jones)  ;  Saucito,  near  Sierra  San  Francisquito,  Brandegee,  Oct.  14, 
1893,  (UC)  ;  Sierra  de  la  Laguna,  Brandegee,  Jan.  21,  1890,  (UC)  ;  Mirafiores,  Brandegee, 
Oct.  14,  1890,  (UC)  ;  Arroyo  Hondo,  Sierra  de  la  Laguna,  Brandegee,  Oct.  22,  1893,  (UC)  ; 
San  Felipe,  Purpus  469,  Jan.-March,  1901,  (LTC).  Sonora,  Mexico:  San  Bernardo,  Rio 
Mayo,  Gentry  1338,  (UC). 


200  CALIFOSXIA  ACADEMY  OF  SCIENCES        [Proc.  4th  ser.  ; 

The  specimens  from  Baja  California  are  slightly  more  robust  than  tliose 
from  Arizona  and  New  Mexico,  but  in  qualitative  characters  the  southern 
and  northern  specimens  are  identical.  A  careful  search  revealed  no  morpho- 
logical characters  accompanying  the  perennial  habit  of  Jones'  material,  so  his 
variety,  D.  Fenclleri  var.  perennis  is  reduced  to  synonomy  under  this  species. 

8.  Drymaria  viscosa  S.  Wats,  ex  Orcutt.  AVest  Am.  Sci.  2:57, 

1886,  nonien  nudum.  Proc.  Am.  Acad.  22:469,  1887 

(Plate21,  figs.  20-27) 

Annual  with  prostrate  or  spreading  branches,  dichotomously  branched 
stems  5-20  cm.  long,  and  glandular-puberulent  leaves,  pedicels,  and  sepals; 
internodes  to  6  cm.  long,  glabrous  or  sparsely  glandular-puberulent ;  stipules 
filiform-.subulate,  1-1.5  mm.  long,  glandular;  leaves  often  pseudo-verticillate, 
linear  to  linear-oblanceolate,  1.5  mm.  wide  or  less,  to  15  mm.  long,  acute  or 
obtuse  at  the  apex,  gradually  narrowing  to  a  slender  petiole  1-3  mm.  long; 
inflorescence  cymose;  peduncles  subfiliform,  to  3.5  cm.  long;  pedicels  1-3  mm. 
long  or  some  flowers  subsessile ;  sepals  narrowly  to  broadly  elliptic,  0.6-1.2 
mm.  wide,  about  2.5  mm.  long,  the  green  midrib  ending  in  an  excurrent  apicu- 
lation  slightly  below  the  apex  of  the  scarious,  glabrous  part ;  petals  bifid 
about  two-thirds  of  the  way  to  the  base,  2  mm.  long,  the  linear-spatulate  lobes 
and  narrow  claw  naked ;  stamens  about  1  mm.  long,  equaling  the  ovary  ;  style 
half  as  long  as  the  ovary;  cap.sule  broadly  ovoid,  nearly  equaling  the  sepals; 
seeds  obovoid-eircinate,  0.6  mm.  high,  pale  cream,  with  light  amber  angles 
and  dorsal  band,  smooth  or  slightly  wrinkled,  but  not  tessellate  nor  muricu- 
late,  dull. 

Tyije  locality: — Socorro,  northern  Baja  California,  Mexico.  (Misspelled 
"Socono"  in  the  original  publication). 

Distribution: — In  sandy  soil  from  about  30°  20'  N.  Lat.  (vicinity  of  San 

Quintm)  southward  to  the  Cape  Region,  Baja  California. 

Specimens  examined: — Socorro,  Orcutl  1330,  (G,  type,  D,  UC,  isotypes)  ;  Lagoon  Head, 
Palmer  767,  March,  1889,  (CA)  ;  Magdalena  Island,  Orciitf  51,  March,  1917,  (CA)  ;  Bran- 
degee,  March  13,  1889,  (UC)  ;  mainland  near  Ascension  Island,  Anthony,  April  17,  1897, 
(UC)  ;  Cahnalli,  Purpun,  Jan.-March,  1898,  (UC)  ;  along  beach  on  sand  dunes,  4  miles  south 
of  Guadalupe,  Whitehead  830,  (D)  ;  sandy  wash  27.4  miles  south  of  Pozo  Aleman,  Wiggins 
7862,  (D)  ;  San  Gregorio,  Brandegee,  Feb.  1,  1889,  (D). 

The  light  colored,  nearly  or  quite  smooth  seeds  and  the  heavily  viscid  herb- 
age of  this  species  are  distinctive.  In  general  appearance  the  plant  resembles 
Mollugo  cerviana  (L.)  Seringe. 

9.  Drymaria  sperguloides  A.  ( Jray,  Am.  Acad.  Arts  and  Sci. 

Mem.  11.4:11,  1849 
(Plate  21,  figs.  28-38) 
Mollugophytum  sperguloides  M.  E.  Jones,  Extract  from  Contr.  West.  Bot.  18:3o.  1933. 

Slender,  erect,  simple  or  dichotomously  branched,  glabrous  annual  0.5-2 
dm.  high ;  leaves  in  fascicles  of  4-6,  pseudo-verticillate,  linear,  0.5-3  cm.  long,  j 


Vol.  XXV]       WIGGIXS:  DRYMABIA  IN  SONOBAN  DESEBT  201 

the  stipules  trianoiilar-siibiilate,  1-1.5  mm.  long,  scarioiis  ;  flowers  in  terminal 
and  subterminal  cymnles,  on  peduncles  0.5-2.5  cm.  long;  pedicels  snbfiliform, 
1-5  mm.  long ;  sepals  dissimilar,  narrowly  to  broadly  elliptic,  1-1.5  mm.  wide, 
about  2.5  mm.  long,  cuculate,  1-nerved,  the  apex  rounded  or  obtuse,  central 
green  portion  terminating  slightly  below  the  apex  of  the  scarious  margins, 
glabrous;  petals  2-2.5  mm.  long,  bifid  about  to  the  middle,  2  linear  appendages 
two-thirds  as  long  as  the  main  lobes  arising  at  the  bottom  of  the  sinus,  claw 
wanged,  nearly  as  broad  as  the  blade ;  stamens  about  1  mm.  long,  three-fourths 
as  long  as  the  ovoid  ovary ;  stigma-lobes  subsessile ;  capsule  broadly  ovoid  to 
subglobose,  3-5  mm.  long,  exceeding  the  sepals ;  seeds  obovoid-circinate,  not 
compressed,  1-1.2  mm.  long,  dark  brown,  dull,  minutely  rounded-tuberculate 
in  longitudinal  rows. 

Ti/pe  locality: — "Valley  of  Santa  Fe  Creek  in  the  mountains,"  New  Mexico. 

Distrihution: — Mountain  slopes  and  meadows,  Arizona  to  western  Texas 
and  northern  Chihuahua. 

Specimens  examined: — New  Mexico:  without  furtlier  locality,  Fendler  55,  1847,  (G, 
type)  ;  C.  Wright  867,  (G,  UC)  ;  Mogollon  Mountains,  on  or  near  west  fork  of  Gila  Eiver, 
Socorro  County,  Meicalfe  349,  (D).  Texas:  near  Presidio  del  Norte,  Parry,  (G).  Arizona: 
Patagonia  Mts.,  Kearney  and  Peebles  10042,  (UC)  ;  near  San  Francisco  Mts.,  Fidion  7368, 
(CA)  ;  EastvieAv,  Eincon  Mts.,  Blumer  3583,  (D)  ;  Barfoot  Park,  Chiricahua  Mts.,  Blumer 
1433,  (D).  Chihuahua,  Mexico:  hills  and  mesas  near  Guerrero,  Pringle  1192,  (D). 

The  label  on  the  type  sheet  bears  no  locality  other  than  "Novo-Mexicanae," 
but  the  inscription  "Drymaria  sperguloides  n.  sp."  is  in  the  handwriting  of 
Asa  Gray,  and  it  is  assumed  that  the  more  detailed  locality  data  were  obtained 
by  him  from  Fendler's  notes  or  through  correspondence. 

The  large  capsules  and  seeds  are  distinguishing  characters.  The  pseudo- 
verticillate  arrangement  of  the  leaves  is  sufficient,  even  in  young  plants,  to 
separate  it  from  D.  tenella  and  D.  effusa,  both  of  wdiich  have  linear  leaves 
and  a  similar  habit. 

10.  Drymaria  arenarioides  Willd.  in  Roem.  and  Schult.  Syst. 

Veg.  5:406,  1819 

(Plate  21,  fig.  39;  Plate  22,  figs.  1-8) 
Drymaria  franlcenioides  H.  B.  K.  Nov.  Gen.  et  Sp.  6:21.  pi.  515, 1823. 

Prostrate,  glandular-puberulent,  canescent,  short-lived  perennial  with  pro- 
fusely branched  stems  3-20  cm.  long;  stipules  subulate,  1-1.5  mm.  long, 
persistent,  slightly  glandular-puberulent ;  leaves  more  or  less  fasciculate, 
linear-lanceolate  to  narrowly  elliptic,  1-2.5  mm.  wide,  5-15  mm.  long,  granu- 
lar-puberulent  and  slightly  canescent,  acute,  tapering  gradually  to  a  short, 
slender  petiole  1-2  mm.  long;  flowers  axillary  and  in  terminal,  1-3-flowered 
cymules;  pedicels  slender,  3-10  mm.  long,  often  deflexed  in  fruit;  sepals  dis- 
similar, narrowly  lanceolate  to  ovate-lanceolate,  1-2.2  mm.  wide,  5-6  mm. 
long,  acute  to  short-acuminate,  weakly  1-nerved,  the  margins  scarious  ;  petals 
5-6.5  mm.  long,  divided  into  6  lobes  about  to  the  middle,  the  main,  outer  ones 


202  CALIFORNIA  ACADEMY  OF  SCIENCES        [Proc.  4th  sek. 

1-1.2  mm.  wide,  the  inner  4  in  2  pairs  of  filiform  appendages  slightly  shorter 
than  the  main  lobes;  stamens  about  1.5  mm.  long,  equaling  the  ovary;  style 
half  as  long  as  the  ovary;  capsule  ovoid,  4—5  mm.  long;  seeds  orbieular-eir- 
cinate,  slightly  compressed  laterally,  with  a  pronounced,  broad,  dorsal  groove 
and  deep  lateral  depressions,  uniformly  and  minutely  granular-tessellate, 
dull,  brown. 

Tyjye  locality: — Pachuca,  Mexico. 

.  Distrihution : — Northern  Sonora  and  Chihuahua  and  southward  into  cen- 
tral Mexico. 

Specimens  examined: — Chihuahua:  hills  and  plains  near  Chihuahua  (City),  Pringle  715, 
(UC)  ;  Chihuahua,  Purpus  1123,  (UC).  Sonora:  gravelly  slopes  20  miles  south  of  Llano, 
Wiggins  7235,  (D)  ;  Shreve  7322,  (D)  ;  southern  edge  of  oak  belt  35  km.  south  of  Nogales, 
Ferris  8810,  (D)  ;  rocky  benches,  Nogales  road,  about  1  mile  north  of  Cumetal,  Ahrams 
13178,  (D).  Hidalgo:  Ixmiquilpan,  Purpus  483,  Feb.,  1905,  (UC).  Zacatecas:  Purpus  133, 
1903,  (UC). 

The  filiform  central  lobes  of  the  petals  are  surprisingly  uniform  in  length, 
width,  and  point  of  insertion.  There  is  a  visible  vascular  strand,  similar  to 
those  running  to  the  main  lobes,  leading  to  each  pair  of  the  appendages.  These 
bundles  depart  from  the  main  petal  bundle  just  below  the  sinus. 

The  seeds  are  distinctive  in  that  the  tessellations  are  not  arranged  in  rows, 
but  form  a  low,  close  mosaic  over  the  whole  surface.  The  broad,  dorsally 
grooved  back,  and  the  tight  coil  of  the  embryo  are  noteworthy  also. 

ll.Drymaria  peninsularis  Blake,  Journ.  "Wash.  Acad.  Sci.  14:285,  192-4 

(Plate22,  figs.  9-16) 

Prostrate,  decumbent,  or  ascending  annual,  or  sometimes  persisting  as  a 
perennial,  with  many  freely  branched  stems  0.5-2  dm.  long;  herbage  pale 
green  or  slightly  glaucescent,  glandular-hirtellous  throughout ;  internodes 
2-25  mm.  long;  stipules  scarious,  undulate,  entire,  0.6-0.8  mm.  long;  leaves 
linear,  somewhat  fleshy,  sessile,  obtuse  or  rarely  acute,  lower  ones  1-2.5  cm. 
long,  the  upper  rarely  over  4  mm.  long,  shorter  than  the  internodes ;  flowers 
solitary  in  the  axils  from  about  the  middle  of  the  stems  to  the  apex ;  pedicels 
2-12  mm.  long  (in  fruit) ,  erect  to  strongly  deflexed  ;  sepals  slightly  dissimilar, 
narrowly  to  broadly  ovate-oblong,  1-2  mm.  wide,  3.5-4  mm.  long  at  anthesis, 
4r^.5  mm.  long  in  fruit,  obtuse  or  rounded,  obscurely  1-  or  3-nerved,  the  cen- 
tral part  green,  herbaceous  and  glandular-puberulent,  the  margins  scarious, 
glabrous ;  petals  white,  3.5-4  mm.  long,  curved,  so  just  equaling  the  sepals, 
4— 5-fid  nearly  to  the  middle,  the  outer  lobes  broader  and  longer  than  the  cen- 
tral ones,  claws  slightly  dilated  and  erosulate  near  the  base,  nearly  as  broad 
as  the  limb  ;  stamens  about  two-thirds  as  long  as  the  sepals,  slightly  surpassing 
the  ovary;  style  and  stigmatic  lobes  about  equal ;  capsule  ovoid,  3-4  mm.  long ; 
seeds  round-reniform,  plump,  0.7-0.9  mm.  long,  fuscous,  finely  and  bluntly 
tessellate,  the  bosses  near  the  base  of  the  dorsal  surface  in  longitudinal  rows, 
not  aligned  elsewhere. 


Vol.  XXV]        WIGGINS:  DRYMARIA  IN  SONOBAN  DESERT  203 

Type  locality: — Cape  Region,  Baja  California,  Mexico. 
Distribution: — Known  only  from  the  Cape  Region  of  Baja  California, 
Mexico. 

Specimens  ei-amined :  Cape  Region,  Pur  pun  423,  (UC,  isotype) ;  Cabo  San  Lucas,  Bran- 
degee  30,  March  17,  1892,  (D,  UC)  ;  Brandegee,  Nov.,  1902,  (UC)  ;  Canyon  Salado,  Purpus 
337,  (UC)  ;  coast  south  of  Pescadero,  Brandegee,  Nov.,  1902,  (UC)  ;  Arroyo  Salado  (San 
Jose  del  Cabo),  Purpus  496,  (UC)  ;  El  Taste,  Brandegee,  Sept.  11,  1893,  (UC)  ;  Cabo  San 
Lucas,  Johansen  541,  (D). 

The  leaves  are  narrower  and  the  upper  ones  more  reduced  in  Z>.  peninsnlaris 
than  they  are  in  D.  arenarioides,  with  which  the  former  was  long  identified. 
The  sepals  of  D.  arenarioides  are  more  acute  than  they  are  in  D.  peninsnlaris, 
and  the  appendages  in  the  sinuses  of  the  petals  are  longer  and  narrower  in 
the  former  species  than  they  are  in  the  latter.  The  seeds  of  the  two  species 
are  quite  similar. 

12.  Drymaria  Johnstonii  Wiggins,  sp.  nov. 

(Plate22,  figs.  17-21,  36) 

Planta  annua  gracilis,  ramulis  decumbentibus  vel  ascendentibus  glaucis 
5-10  em.  longis  gianduloso-puberulentibus ;  stipulae  subulatae  0.5-0.8  mm. 
longae ;  folia  opposita  vel  f asciculata  linearia  subcarnosa  2-5  mm.  longa  ses- 
silia  apicem  obtusa  vel  rotundata ;  tlores  axillarii  solitarii,  sepalis  2.2-2.5  mm. 
longis  ellipticis  obtusis,  margine  membraneis,  petalis  semibifidis  2.5-3  mm. 
longis  albis  nudis  sepala  vix  superantibus;  staminibus  ovarium  vix  superanti- 
bus ;  ovarium  1  mm.  altum ;  capsula  anguste  ovata  2.5-3  mm.  longa  sepala 
vix  superantia;  semina  2-5  nigra  obovata  minute  granulo-muriculata  0.6  mm. 
longa. 

Annual,  with  freely  branching,  slender,  decumbent  or  spreadingly  ascend- 
ing stems  1-1.5  dm.  long  and  slightly  fleshy,  glaucescent  leaves,  puberulent 
throughout  with  stout,  short-stipitate,  whitish  glandular  and  simple  hairs; 
internodes  5-14  mm.  long;  stipules  subulate,  scarious,  0.5-0.8  mm.  long, 
persistent;  leaves  opposite  or  the  lower  ones  sometimes  fasciculate,  linear, 
slightly  fleshy,  2—5  mm.  long,  or  basal  ones  up  to  1  cm.  long,  faintly  1 -nerved, 
sessile,  rounded  or  obtuse  at  the  apex;  flowers  axillary,  solitary;  pedicels 
slender,  1-2  mm.  long  at  anthesis,  4-8  mm.  long  in  fruit ;  sepals  herbaceous- 
fleshy.  2.2-2.5  mm.  long,  elliptic,  obtuse  at  the  apex,  the  scarious,  hyaline 
margins  evident  on  the  inner  sepals  and  the  covered  edges  of  the  outer  sepals; 
petals  white,  bifid  nearly  to  the  middle,  devoid  of  appendages,  2.5-3  mm. 
long,  slightly  surpassing  the  sepals ;  stamens  equaling  or  slightly  surpassing 
the  ovary ;  style  0.2  mm.  long  or  less,  much  exceeded  by  the  stigma-lobes ; 
capsule  narrowly  ovoid,  acute,  2.5-3  mm.  long,  equaling  the  sepals  or  sligh'tly 
exceeding  them,  2-5-seeded ;  seeds  black,  0.6  mm.  long,  obovoid-reniform, 
minutely  and  uniformly  granular-muriculate,  dull. 

Tyjye  in  the  herbarium  of  the  California  Academy  of  Sciences  no.  82814, 
collected  in  crevices  of  rock  on  mesa  near  crest  of  island,  the  isthmus,  Espiritu 
Santo  Island,  Baja  California,  Mexico,  Ivan  M.  Johnston  3972,  May  31, 1921. 


204  CALIFORNIA  ACADEMY  OF  SCIENCES        [Proc.  4th  ser. 

Only  one  plant  was  seen  and  the  species  has  not  been  re-eollected.  Johnston 
(Proc.  Calif.  Acad.  Sci.  [IV],  12:1023,  1924)  determined  it,  tentatively,  as 
D.  arenarioides  and  stated  that  the  plant  seemed  undescribed.  He  also  said 
that  the  plant  on  the  mainland  of  Baja  California  which  had  been  called 
D.  arenarioides  was  different,  both  from  his  Espiritn  Island  plant  and  the 
material  called  I),  arenarioides  from  eastern  Mexico.  Subsequently  S.  F.  Blake 
described  the  peninsula  plant  as  D.  peninsularis,  but  he  did  not  have  access 
to  this  material  of  Johnston's. 

D.  Johnsfonii  closely  resembles  a  depauperate  plant  of  D.  peninsidaris,  but 
differs  from  that  species  in  having  few  leaves  over  4—5  mm.  long,  petals  en- 
tirely devoid  of  appendages,  and  in  having  seeds  that  are  less  tightly  cirei- 
nate,  ungrooved  dorsally,  and  in  being  much  more  finely  muriculate  over  the 
Avliole  surface. 

13.  Drymaria  effusa  A.  Gray,  PI.  Wright.  2:19,  1853 
Slender  annual  0.8-2  dm.  high,  dichotomously  branched  above  the  second 
or  third  nodes,  minutely  puberulent,  or  the  internodes  becoming  glabrate; 
internodes  2-4  cm.  long,  or  the  lowest  slightly  longer;  stipules  filiform- 
subulate,  1.5-2.2  mm.  long,  entire;  basal  leaves  obovate-spatulate,  1.5-3  mm. 
wide,  3-10  mm.  long,  rounded,  broadly  cuneate  at  the  base,  green,  sparsely 
puberulent,  on  petioles  1-2  mm.  long,  or  subsessile ;  cauline  leaves  filiform 
or  narrowly  linear,  sessile,  1-2.5  cm.  long,  or  the  uppermost  reduced,  faintly 
1-nerved,  minutely  puberulent ;  flowers  solitary  in  the  axils  of  the  upper 
leaves  and  of  scarious,  ovate  bracts  0.5-1.5  mm.  long;  pedicels  filiform.  2-4 
mm.  long ;  sepals  ovate,  slightly  dissimilar,  1.4-1.8  mm.  long,  obtuse  or  rounded 
at  the  apex,  inconspicuously  1-nerved,  minutely  but  closely  puberulent.  the 
winged  margins  scarious;  petals  bifid  almost  to  the  center,  2-2.8  mm.  long, 
exceeding  the  sepals,  devoid  of  appendages;  claws  narrow  and  naked;  sta- 
mens about  equaling  the  sepals ;  ovary  1  mm.  high ;  style  0.4—0.6  mm.  long, 
about  equaled  by  the  filiform  stigma-lobes;  seeds  (immature)  reddish  brown, 
muriculate. 

Tyj^e  locality: — "Mountains  east  of  Santa  Cruz,  Sonora." 
Distribution: — Mountains  in  Santa  Cruz,  Cochise,  and  Pima  counties,  Ari- 
zona, and  southward  into  adjacent  northern  Sonora. 

Specimens  examined: — Sonora  :  mountains  east  of  Santa  Cruz,  C.  Wright  869,  Sept.,  1851, 
(G,  type).  Arizona:  Babocomari,  Santa  Cruz  County,  Thiirber  995,  (G)  ;  Sonoita  Valley, 
BoihrocTc  169,  (G)  ;  "S.  Arizona,"  Lemmon  509,  1881,  (G)  ;  Huachuca  Mountains,  Price, 
Aug.  14,1895,  (D). 

14.  Drymaria  tenella  A.  Gray,  Mem.  Am.  Acad.  IT.  4 :  12,  1849 

(Plate  22,  figs.  22-27,  34) 

Erect,  slender  annual  0.5-2  dm.  high,  dichotomously  branched  above  the 
second  or  third  node,  glabrous,  or  glandular  puberulent  in  the  variety;  inter- 
nodes to  3.5  cm.  long;  basal  leaves  spatulate-obovate,  1-2.5  mm.  long  on  pedi- 
cels of  equal  length ;  cauline  leaves  linear,  sessile,  3-15  mm.  long,  acute,  in- 


Vol.  XXV]        WIGGIXS:  DBYMARIA  IN  SOXORAN  DESERT  205 

conspicuously  1-nerved,  glabrous;  stipules  white,  scarious,  linear-subulate, 

1  mm.  long-  or  less,  erect ;  flowers  sliort-peclicellate  or  subsessile  in  the  axils 
or  terminal  on  slender  peduncles  5-10  mm.  long ;  sepals  ovate-lanceolate, 
0.8-1.2  mm.  wide,  2.8-3.2  mm.  long,  acute,  strongly  3-ribbed,  the  margins 
.scarious;  petals  white,  2.3-2.8  mm.  long,  bifid  to  middle  or  slightly  below, 
the  lobes  oblong,  entire  or  minutely  emarginate,  devoid  of  appendages,  claws 
narrow  and  naked;  stamens  slender,  about  equaling  the  1-1.3  mm.  high  ovary; 
style  0.2-0.4  mm.  long,  stigmatic  lobes  about  as  long;  capsule  ovoid,  about 

2  mm.  high,  8-12-seeded ;  seeds  obovoid-reniform,  0.6-0.8  mm.  long,  minutely 
but  sharply  muriculate,  reddish  brown. 

Tyjie  locality: — Eight  miles  west  of  Las  Vegas,  New  Mexico. 

Distribution: — Mountains  of  Arizona,  New  Mexico,  and  adjacent  Chihua- 
hua and  Sonora,  and  in  northern  Baja  California,  Mexico. 

Specimens  examined: — New  Mexico:  eight  miles  west  of  Las  Vegas,  Fendler  56,  Aug., 
1847,  (G,  type)  ;  Pinos  Altos  Mts.,  E.  L.  Greene  332,  Sept.  8,  1880,  (G)  ;  Sawyer's  Peak, 
Grant  County,  Metcalfe  1428,  (CA)  ;  \vithout  definite  locality,  C.  Wright  868,  1851,  (G). 
Arizona:  Manning  Camp,  Eincon  Mts.,  Pima  County,  Bhnner  3368,  (D)  ;  rliyolite  spur  just 
N.  of  Wilgus  Ranch,  Chiricahua  Mts.,  Cochise  County,  Blumer  1639,  (D)  ;  Barfoot  Park, 
Chiricahua  Mts.,  Blumer  1411,  (D).  Chihuahua:  pine  plains,  base  of  the  Sierra  Madre, 
Pringle  1194,  (D).  Baja  California,  Mexico:  margins  of  meadow  at  La  Encantada,  Sierra 
San  Pedro  Martir,  alt.  2200  m.,  Wiggins  and  Demaree  4915,  (D). 

This  species  is  very  similar  to  B.  effiisa  in  general  appearance,  but  it  is 
separated  from  that  species  by  the  acuminate  sepals  and  the  lack  of  puberu- 
lence  on  the  herbage. 

14a.  Drymaria  tenella  A.  Gray,  var.  nodosa  (Engelm.),  Wiggins,  n.  comb, 

(Plate  22,  figs.  28-33,  37-38) 
Drymaria  nodosa  Engelm.  apud  A.  Gray,  Mem.  Am.  Acad.  II.  4:12,  1849. 

Upper  parts  of  internodes,  peduncles,  pedicels  and  herbaceous  portions  of 
the  sepals  minutely  glandular-puberulent ;  sepals  short-acuminate ;  otherwise 
as  in  typical  I),  tenella. 

Type  locality: — Cosiquiriachi,  Chihuahua,  Mexico. 

Distribution : — Northern  Chihuahua  and  northeastern  Sonora. 

Specimens  examined: — Chihuahua:  pine  plains,  base  of  the  Sierra  Madre,  Pringle  1195, 
(D)  ;  thin  soil,  rocky  hills  near  Chihuahua,  Pringle  716,  (D).  Sonora:  Los  Pinitos,  alt.  6100 
ft..  Hart  man  138,  (UC). 

There  are  only  two  minor  characters  by  which  the  variety  can  be  separated 
from  D.  tenella,  and  they  are  of  insufficient  value  to  warrant  retaining  this 
entity  as  a  distinct  species. 

15.  Drymaria  depressa  Greene,  Leafl.  Bot.  Obs.  and  Crit.  1 :  153,  1905 

(Plate22,  figs.  39-43) 

Annual,  freely  branched  plant  with  slender  branches  2-4  cm.  long,  de- 
pressed or  spreading,  glabrous,  the  internodes  0.5-2  cm.  long ;  stipules  0.5-1.2 
mm.  long,  whitish,  slightly  curled;  leaves  oblanceolate,   1.5-3  mm.  wide. 


206  CALIFORNIA  ACADEMY  OF  SCIENCES        [Proc.  4th  ser. 

5-13  mm.  long,  obtuse  at  the  apex,  graduall}'  narrowing  to  the  sessile  base, 
glabrous,  faintly  3-nerved ;  inflorescence  of  compact  cymules;  pedicels  1.5 
mm.  long  or  less,  minutely  granular-puberulent ;  sepals  oblong,  2.2-2.6  mm. 
long,  rounded  at  the  apex,  narrowly  scarious-margined,  glabrous,  distinctly 
3-nerved;  petals  narrow,  bifid  to  the  middle,  2-2.2  mm.  long,  devoid  of  append- 
ages, the  claws  narrow,  naked ;  stamens  slightly  over  half  as  long  as  the  petals, 
and  about  equaling  the  ovary  at  anthesis;  ovary  short-stipitate ;  style  nearly 
as  long  as  the  ovary,  the  stig-ma-lobes  half  as  long ;  capsule  narrowly  ovoid, 
nearly  equaling  the  sepals  at  maturity,  8-12-seeded  ;  seeds  obovoid-renif  orm, 
0.6-0.8  mm.  long,  light  buff,  uniformly  and  finely  tessellate  with  low,  rounded 
welts,  dull. 

Tyjje  local  a  y: — Sawyer's  Peak,  Grant  County,  New  Mexico. 

Distribution: — Mountains  in  eastern  Arizona  and  New  Mexico. 

Specimens  examined: — Open  glade  on  Sawyer's  Peak,  Grant  County,  New  Mexico,  alt. 
9500  feet,  Metcalfe  1430,  (CA,  isotype). 

Kearney  and  Peebles  suggest  that  this  species  may  not  be  distinct  from  D. 
effusa,  but  it  seems  to  me  that  the  broader  leaves,  compact  habit,  and  lack  or 
paucity  of  puberulence  separates  it  from  that  species.  I  have  not  seen  the  Ari- 
zona material. 


EXPLANATION  OF  PLATES 


PLATE  20 
Fig.  1.  Diagram  of  cyniose  inflorescence  common  in  Drymaria. 

Fig.  2.  Diagram  of  inflorescence  of  D.  sprrguloides;  solidly  inked  circles 
indicate  abortive  flowers. 

Fig.  3.  Diagram  of  aestivation  of  sepals  in  Drymaria. 

Figs.  4-12.  I),  holosteoides ;  figs.  4-8,  sepals,  x  6.6;  fig.  9,  petal,  x  13;  figs. 
1(1-12,  X  6.6;  fig.  10,  ovary  and  stamens;  fig.  11,  ovary;  fig.  12,  petal. 

Figs.  13-19.  D.  pachyphylla,  x  6.6;  fig.  13,  petal;  fig.  14,  ovary  and  stamens; 
figs.  15-19,  sepals. 

Figs.  20-26.  D.  crassifolia,  x  6.6;  fig.  20,  ovary  and  stamens;  fig.  21,  petal; 
figs.  22-26,  sepals. 

Figs.  27-31.  B.  carinata,  figs.  27-29,  x  6.6;  figs.  30-31,  x  13;  fig.  27,  petal; 
figs.  28-29,  sepals;  figs.  30-31,  petal  and  sepal  from  different  plant. 

Figs.  32-34.  D.  carinata  var.  pereniiis,  x  13;  fig.  32,  petal  before  anthesis; 
figs.  33-34,  petals  from  different  plants  at  anthesis. 


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PROC.  CALIF.  ACAD.   SCI..   4TH   SERIES.   VOL.   XXV.    NO.    6  [WIGGINS]  PLATE  20 


[  209  ] 


PLATE  21 

Fig.  1.  D.  crassifolia,  x  6.6  ;  arrangement  of  leaves  at  a  node,  the  dotted  lines 
indicate  a  hyaline  "sheath"  of  epidermal  and  subepidermal  cells  overlying  the 
photosynthetic  tissue. 

Figs.  2-5.  D.  polystachya,  petal  and  sepals,  x  6.6. 

Figs.  6-8.  D.  polystachya  var.  diffusa,  sepals  and  petal,  X  6.6. 

Figs.  9-10.  D.  debilis,  petal  and  sepal,  x  6.6. 

Figs.  11-19.  Z>.  Fendleri,  x6.6;  figs.  11-12,  petal  and  ovary  and  stamens 
from  a  flower  from  New  Mexico ;  fig.  13,  mature  capsule  from  same  plant;  figs. 
14-19,  sepals  and  petal  from  plant  from  Baja  California,  Mexico. 

Figs.  20-27.  D.  viscosa,  x  13;  fig.  20,  petal;  fig.  27,  ovary  and  stamens;  figs. 
21-26,  sepals  (fig.  22,  lateral  view  of  a  sepal). 

Figs.  28-38.  B.  sperguloides,  x  13 ;  figs.  28-31,  35-36,  sepals ;  figs.  32-33, 
petals ;  fig.  34,  ovary  and  stamens ;  fig.  37,  mature  capsule  and  persistent  sepals; 
fig.  38,  arrangement  of  leaves  and  branches  at  a  node. 

Fig.  39.  D.  arevarioides,  petal,  x  6.6. 


[210] 


PROC.   CALIF,  ACAD.   SCI..   4TH   SERIES.  VOL.  XXV.    NO.    6  [WIGGINS]  PLATE  21 


[211] 


PLATE  22 

rigs.  1-8.  D.  arenarioides,  x  6.6;  figs.  1-5,  sepals;  fig.  6,  ovary  and  stamens; 
figs.  7-8,  petals. 

Figs.  9-16.  D.  peninsularis,  xS.6;  figs.  9-13,  sepals;  figs.  14-10,  petals. 

Figs.  17-21,  36.  D.  Johnstonii,  x  13;  fig.  17,  flower  at  antliesis;  figs.  18-19, 
sepals;  fig.  20,  ovary  and  stamens;  figs.  21,  36,  petals  and  adjacent  stamens. 

Figs.  22-27,  34.  B.  tenella,  x  6.6;  figs.  22-26,  sepals;  fig.  27,  ovary  and  sta- 
mens ;  fig.  34,  petal. 

Figs.  28-33,  37-38.  D.  tenella  var.  nodosa,  x  6.6;  figs.  28-30,  37-38,  sepals; 
fig.  31,  ovary  and  stamens;  figs.  32-33,  petals. 

Figs.  39-43.  P.  depressa,  x  13  ;  fig.  39,  petal ;  figs.  40-43,  scpaLs. 


[212] 


PROC.  CALIF.  ACAD.  SCI..   4TH   SERIES.  VOL.  XXV,    NO.    6  [WIGGINS]  PLATE  22 


[213] 


PROCEEDINGS 


OF  THE 

CALIFORNIA  ACADEMY  OF  SCIENCES 


Fourth  Series 
Vol.  XXV,  No.  7,  pp.  215-220 


June  1,  1944 


ALICE  EASTWOOD 
SEMI-CENTENNIAL  PUBLICATIONS 

No.  7 
WATER  AND  PLANT  ANATOMY 

BY 

GEORGE  J.  PEIRCE 

Professor  of  Botany,  Emeritus 
Stanford  University,  California 


FORTUNATELY  the  processcs  of  Nature  go  on  whether  man  understands  them 
or  not ;  but  curiosity,  which  has  led  man  to  enquire  about  everything  inside 
and  outside  himself,  has  brought  not  only  the  satisfaction  sought,  but  has 
stimulated  continued  enquiry,  has  led  to  the  development  of  powers  formerly 
unsuspected,  and  to  the  recognition  of  reasons  why  the  powers  are  not  un- 
limited. It  has  been  discovered  that  living  things  require  air  to  breathe,  water 
to  absorb,  food  to  take  in ;  what  it  is  in  air,  water,  food  which  is  indispensable 
and  not  merely  stimulating;  and  some  of  the  relations  between  the  worlds 
outside  and  inside  the  living  thing.  One  of  these  relations  concerns  itself  with 
the  intake  and  outgo  of  one  or  other  of  these  things — air,  water,  food.  In  this 
paper  in  honor  of  an  associate  of  many  years  I  wish  to  consider  one  of  these 
relationships,  thereby  contributing  also  to  an  understanding,  or  at  least  a 
conjecture,  as  to  why  grass,  chaparral,  or  forest  should  be  the  natural  cover 
of  respective  areas. 

The  body  of  the  higher  land  plants  consists  of  tissues  among  which  the  vari- 
ous functions  are  divided  and  by  which  the  various  needs  are  met ;  but,  while 
there  is  division  of  labor,  each  tissue  and  each  organ  of  the  plant  may  perform 
secondary  functions,  or  be  the  seat  of  subsidiary  phenomena.  Thus  the  green 
leaf,  primarily  for  the  production  of  food,  is  the  seat  of  maximum  gas  ex- 
change and  of  water  loss.  The  green  leaf  represents  in  its  shape,  size,  dis- 
tribution, anatomy,  duration,  and  even  number,  the  adjustment  which  each 
individual  leaf,  as  well  as  the  whole  plant,  makes  among  these,  and  many 
other,  less  evident,  phenomena.  Similarly  the  root,  primarily  for  the  meehani- 

[  215  ] 


216  CALIFORNIA  ACADEMY  OF  SCIENCES        [Proc. -Ith  ser. 

cal  support  of  the  plant  and  the  intake  of  water,  must  provide  also  for  passing 
the  water  on  to  other  parts  of  the  body.  But  obviously  these  tasks  vary  accord- 
ing to  diverse  or  changing  external  conditions,  the  intake  of  water  from  moist 
soil  in  the  rainy  season  appearing  to  be  easier  and  more  understandable  than 
by  the  same  root  from  the  same  soil  in  the  dry  season.  The  facile  statements 
that  water  intake  by  roots  is  an  osmotic  phenomenon,  or  that  "work  is  done,'' 
ignore  the  fact  that,  under  conditions  very  different  from  those  of  well- 
watered  Europe,  the  relative  densities  of  the  solutions  in  root  and  in  soil  are 
also  very  different. 

In  order  to  provide  the  path  of  water  movement  from  soil  to  air  through 
the  body  of  the  plant  there  must  be  a  vascular  system,  again  so  flexible  that  it 
will  meet  the  need  of  the  plant  in  all  of  its  circumstances,  ranging  from  satu- 
rated soil  and  air  to  soil  and  air  almost  dry.  Combining  mechanical  support 
and  water  lift,  one  finds  the  vascular  tissues  variously  strengthened  and  the 
vascular  tissues  themselves  variously  composed  of  elements  as  different  as 
the  small-one-celled  tracheids  of  the  wood  of  the  conifers  and  the  large  many- 
celled  vessels  of  the  annual  herbaceous  plants  of  the  rainy  season.  Further- 
more, the  same  individual  plant  may  regularly  produce  different  conducting 
tissue  in  the  same  year  according  to  the  external  environment. 

I  should  like  now  to  cite  some  of  the  evidence  on  which  these  statements  rest, 
specifically  in  regard  to  the  conducting  tissues  of  the  vascular  land  plants. 

As  shown  in  previous  papers,'  the  quantity  and  the  physical  state  of  water 
in  the  conducting  tissues  of  plants,  and  their  parts,  varies  from  a  liquid  exert- 
ing outward  pressure  to  a  vapor  under  internal  tension  or  suction :  from 
columns  of  liquid  water  more  or  less  completely  filling  the  lumina  of  ducts 
and  tracheids  to  sheets  of  liquid  water  on  and  in  the  material  of  the  walls  of 
ducts  and  tracheids,  and  enclosing  masses  of  water  vapor  filling  the  lumina 
of  the  vascular  elements.  We  are  familiar  with  the  phenomena  of  bleeding, 
which  are  the  evidence  of  internal  pressure,  and  only  less  familiar  with  the 
phenomena  of  transpiration  or  evaporation,  which  result  in  tension  (trac- 
tion) within  the  water  mass  in  the  plant  bod5\  At  seasonal  or  other  intervals 
the  water  in  the  vascular  system  of  the  land  plant  undergoes  these  complete 
changes,  from  liquid  to  vapor,  from  pressure  to  tension  ("negative  pressure") . 
Perennial  land  plants  must  survive  these  changes  or  succumb,  either  locally 
or  extensively.  Many  annual  plants  cannot  maintain  themselves  when  the 
procession  of  the  seasons  from  wet  to  dr}^  entails  these  changes  in  the  quantity 
and  the  state  of  the  water  within  their  conducting  systems.  Examination 
reveals  that  their  vascular  anatomy  does  not  meet  the  requirements  of  ade- 
quate water  holding  and  water  lifting. 

Between  the  so-called  "water  reservoirs,'"  which  are  cells  and  tissues  evi- 
dently enlarged  and  otherwise  fitted  for  containing  water  in  their  vacuoles, 

^  Peirce,  G.  J.  The  state  of  water  in  ducts  and  tracheids.  Plant  Physiology,  11,  1936,  and 
literature  there  cited. 

'  See  Haberlandt,  G.  Physiologische  Pflanzenanatomie,  p.  371  et  seq.  Ctli  ed.  1924  or  later, 
on  Avater  storage  and  water  storing  tissues. 


Vol.  XXV]  PEIRCE:   WATER  AND  PLANT  AN  ATOMY  217 

and  the  -vvet  walls  which  not  merely  contain  but  actually  hold  water  with  great 
tenacity,  there  are  many  intermediates,  differing  from  one  another  as  do  the 
environments  in  which  the  plants  live.  These  wet  walls  vary  in  thickness  as 
they  do  in  hardness  and  stiffness.  They  may  look  as  if  they  were  mechanically 
strong  where  little  or  no  mechanical  strain  can  develop ;  but  they  may  show 
by  their  chemical  composition  that  they  fix — adsorb  and  incorporate — water, 
perhaps  in  the  manner  in  which  the  water  of  crystallization  is  incorporated 
in  blue  copper  sulphate.  It  is  recognized  that  the  crystalline  form  and  the 
crystalline  state  of  copper  sulphate  depend  upon  the  two  constituents  of  the 
crystals,  water  and  copper  sulphate.  Similarly,  but  to  a  certain  point  only, 
and  also  only  partially,  the  shape,  and  the  thickness,  of  cell  walls  depend  both 
on  the  chemical  composition  (cellulose  derivatives  and  their  accompaniments) 
of  the  cell  wall  and  water.  Collenchyma  tissues,  which  are  mechanically  weak 
though  looking  as  if  they  might  have  tensile  or  compressive  resistance,  do  store 
amounts  of  water  which  are  large  in  proportion  to  the  mass.  The  amount  of 
heat  required  to  drive  off  all  of  this  water  indicates  the  force  with  which  this 
water  is  bound :  and  the  shrinkage  shows  no  less  plainly  how  much  the  form 
depends  upon  the  constituent  water. 

Turning  now  to  specific  instances  as  illustrating  these  general  statements, 
.  one  may  consider  Montia  perfoliata  (Donn)  Howell,  "Miner's  lettuce,"  a  suc- 
culent, low,  annual  herb  which  begins  soon  after  the  first  rains  have  consid- 
erably wet  the  soil  of  natural  and  cultivated  areas,  and  which  expires  while 
the  moisture  content  of  the  air  and  of  the  soil  is  still  high.  The  appearance  of 
the  root  system  suggests  that  it  is  adequate  as  an  absorbing  organ ;  the  area 
and  composition  of  the  leaves  and  their  surface  are  such  that  transpiration 
(evaporation)  into  warm,  dry  air,  while  considerable,  need  not  be  excessive, 
provided  the  means  of  moving  water,  even  through  the  limited  distance  of  the 
height  of  these  small  plants,  be  adequate.  But  examination  of  cross  and  longi- 
tudinal sections  of  the  stem  discloses  conducting  tissues  in  vascular  bundles 
too  few  and  too  small  to  meet  the  need.  Furthermore,  the  wood  (xylem)  of 
the  vascular  bundles  consists  of  ducts  (vessels)  of  relatively  large  size  and 
thin  Avails,  and  comparatively  few  and  thin-walled  tracheids.  These  plants 
are  mechanically  weak,  but  they  successfully  carry  their  own  weight  and  the 
increasing  load  of  seeds  and  fruit,  breaking  only  when  unusual  force  is  ap- 
plied. The  supporting  tissues  of  Miner's  lettuce  are  sufficient  to  keep  it  erect : 
but  when  the  strain  of  drying  air  and  soil  is  put  upon  the  conducting  tissues, 
they  are  unequal  to  the  need  of  moving  water  rapidly  from  the  roots  which 
take  it  in  to  the  leaves  which  lose  it.  The  composition  of  the  vascular  bundles 
is  diversified  and  interesting,  but  they  are  inadequate  and  the  plants  die. 

If  one  turn  to  other  native  herbaceous  plants,  the  "tar  weeds"  (Hemizonia 
spf),  one  finds  much  longer  lived  annuals  which  start  from  seed  at  about  the 
same  time  as  Montia  but  which  persist  throughout  the  dry  season  till  the  rains 
come  again.  The  plants  form  more  or  less  perfect  rosettes  of  largish  rather 
fleshy  leaves  quite  close  to  the  ground.  These  are  the  food-manufacturing 


218  CALIFORNIA  ACADEMY  OF  SCIENCES        [Proc.  4th  ser. 

org-ans  wliieli  are  active  for  a  long  time  and  which  lose  water  comparatively 
slowly  because  of  their  position  close  to  the  gronnd,  their  hairy  covering-,  their 
water-proofing  by  cell-walls  and  secretions.  The  stalks  which  bear  the  flowers 
from  early  summer  till  late  fall  are  slender,  dry,  woody,  and  their  leaves  ai-e 
small  and  narrow.  The  surface  of  stalks  and  leaves  is  waterproofed.  The  struc- 
ture is  compact,  as  one  would  expect  in  a  plant  of  such  xerophytic  habit,  with 
adequate  mechanical  support.  The  vascular  bundles  are  composed  of  a  strik- 
ingly large  proportion  of  thick-walled  tracheids,  with  few  and  small  ducts. 
The  limited  area  of  the  surface  of  the  flowering  stalks  and  the  composition  of 
their  vascular  bundles  insures  a  sufficient  water  transport  to  offset  the  water 
loss  into  the  dry  air. 

If  one  compare  these  two  annuals  with  a  plant  which  has  a  perennial  under- 
ground root,  namely  Echinocystis  fahacea  Naud.,  common  manroot,  one  sees 
long,  slender  stems  which  climb  by  means  of  branched  tendrils  very  sensitive 
to  contact.  The  leaves  are  large  and  thin,  smooth  or  only  sparsely  spiny,  not 
thoroughly  waterproofed  nor  structurally  provided  with  water-holding  ma- 
terials. The  stems  are  mechanical^  weak,  except  that  their  tensile  strength  is 
considerable,  especially'  when  dry  :  but  their  vascular  bundles  have  long  been 
the  admiration  of  the  plant  anatomist  because  of  the  arrangement  and  the 
strikingly  large  size  of  the  duets  and  sieve-tubes.  Tracheids  make  up  only  a 
small  part  of  the  wood.  In  the  underground  roots  which,  in  older  plants,  may 
attain  the  size  of  a  half  barrel,  or  even  larger,  there  is  a  great  store  of  water ; 
but  in  spite  of  the  spectacular  vascular  anatomy,  the  leaves  and  stem  dry  and 
die  very  soon  after  the  onset" of  the  dry  season.  Furthermore,  the  leng-th  to 
which  the  leafy  stems  will  grow  seems  to  be  directly  related  to  the  humidity, 
as  well  as  the  temperature,  of  the  air — the  higher  the  humidit}'  during  the 
period  in  which  air  temperature  makes  growth  possible,  the  longer  the  stems 
become.  In  "wet  springs"  when,  because  of  occasional  rains  on  thoroughly 
watered  ground,  air  as  well  as  surface  soil  remain  damp,  the  stems  may  attain 
surprising  lengths  (30  feet)  as  they  climb  over  fences,  bushes,  and  grass.  On 
the  other  hand,  the  same  underground  roots  give  rise  to  decidedly  shorter 
stems  in  the  "dry  springs,"  when  rains  are  infrequent  or  do  not  occur  at  all. 

One  sees  the  same  thing  in  the  common  garden  sweet  pea  which,  under 
various  conditions  of  soil  and  air  temperature,  moisture,  etc.,  produces  slender 
stems  of  very  different  lengths  from  seeds  of  the  same  variety.  Assuming  the 
same  treatment  otherwise  of  soil  and  water,  one's  sweet  pea  vines  will  grow 
tall  or  remain  shorter,  they  will  continue  to  grow  in  length  or  they  will  stop, 
according  to  the  need  of  carrying  water  in  quantity  and  at  speed  through  the 
stems  from  roots  to  leaves.  Where  air  dryness  makes  transpiration  from  the 
leaves  too  rapid  for  the  vascular  bundles  of  the  stems  to  offset,  no  matter  how 
scrupulously  the  bed  may  be  ground-watered,  the  stems  promptl}^  cease  to 
elongate.  The  water-carrying  capacity  of  the  stems,  the  size  and  composition 
of  the  vascular  bundles,  seem  to  determine  the  height  of  the  plants. 

In  all  of  tlie  plants  tluis  cited,  as  well  as  in  all  the  rest,  tlie  anatomical  struc- 


Vol.  XXV]  PEIRCE:   WATER  AND  PLANT  ANATOMY  219 

ture  of  the  stems  may  be  considered  to  be  hereditary ;  but  while  the  plants 
may  vary  according  to  circumstances,  the  limits  of  variation,  as  well  as  their 
behavior  otherwise,  seem  to  be  set  by  hereditary  characters,  and  we  find  the 
sweet  pea  failing  to  grow  indefinitely  in  height  in  spite  of  the  mechanical 
support  of  frame  or  trellis.  The  reason  seems  to  be  that  the  water-carrying 
capacity  of  its  vascular  system  is  limited. 

Aesculus  calif ornica  (Spach)  Nutt.,  the  buckeye,  or  horse  chestnut,  is  a 
small,  spreading  tree  which  puts  forth  its  light  green  leaves,  and  loses  them,  the 
earliest  of  our  native  perennials.  Its  leaves  are  large  and  thin ;  one  might  call 
them  delicate,  but  their  skeleton  is  strong  enough  to  carry  a  very  considerable 
expanse.  They  fall  apparently  in  accordance  with  the  amount  of  water  avail- 
able in  the  soil,  those  buckeye  trees  becoming  bare  earliest  which  are  rooted  in 
dryest  or  thinnest  soil,  and  those  retaining  their  leaves  longest  are  rooted 
where  more  water  is  available.  However,  this  is  not  necessarily  the  case ;  young 
buckeyes  from  one  to  five  or  more  years  old,  grown  in  well-watered  gardens, 
and  even  grown  in  water  cultures,^  in  which  there  is  a  constantly  adequate 
supph"  of  water  as  w^ell  as  mineral  matters  and  air,  lose  their  leaves  before 
the  cold  of  winter  would  make  them  fall.  The  reason  for  this  behavior  is  to  be 
found,  I  believe,  in  the  anatomy  of  the  wood.  The  xylem  consists  of  strikingly 
large,  relatively  thin-walled  ducts  (vessels)  with  fewer  tracheids  or  other  thick- 
v.alled  elements  than  might  be  expected.  While  the  capacity  to  carry  large 
volumes  of  water,  if  the  water  is  abundantly  supplied  from  the  roots,  is  large, 
the  size  of  the  ducts  and  the  thinness  of  their  walls  permit  holding  only  small 
amounts  of  water.  Hence  the  large  leaves  are  insufficiently  supplied  wdth 
water  as  the  dry  season  continues,  and  they  gradually  dry  and  fall. 

If  one  considers  now  the  wood  anatomy,  the  behavior  of  the  foliage,  and 
some  of  the  other  traits  of  two  native  oaks,  Quercus  loiata  Nee,  the  Valley,  or 
White  Oak,  and  Q.  agrifolia  Nee,  the  Coast  Live  Oak,  we  shall  be  led  further. 
Q.  lohata  puts  out  its  summer  canopy  of  leaves  well  after  the  coolness  of  mid- 
winter is  past  but  with  the  soil  water  undiminished ;  and  it  retains  its  foliage 
throughout  the  dry  season,  losing  the  leaves  apparently  when  soil  dryness 
may  add  to  the  difficulty  of  maintaining  its  foliage  through  cooling  nights, 
and  days  in  which  photosynthesis  may  lag.  The  Live  Oak,  Q.  agrifolia,  on  the 
other  hand,  while  also  losing  its  leaves  once  a  year,  puts  out  its  new  crop  when 
the  days  have  already  become  warm,  and  at  least  the  surface  soil  has  begun 
to  dry.  Then  the  leaves  of  the  preceding  year  drop  off.  The  leaves  of  Live  Oak 
are  ordinarily  an  inch  or  two  long,  compact  in  form,  rather  thick  and  leathery, 
smooth  and  shiny  above,  hairy  underneath,  and  weU  waterproofed.  The  leaves 

^  For  six  years  I  have  maintained  cultures  of  buckeye  seedlings  in  one-quart  milk  bottles 
half  filled  with  Knopp's  Solution.  These  seedlings,  one  year  old  and  older,  up  to  six  years, 
are  bare  from  late  August  or  early  September  tUl  February  or  March  in  spite  of  the  nutrient 
solution :  and  they  put  out  new  leaves  in  the  spring  at  about  the  same  time  as  similar  seed- 
lings outside,  rooted  normally  in  the  soil.  The  swelling  of  winter  buds  and  the  putting  forth 
of  leaves  were  not  instigated  in  my  cultures  by  fresh  solution :  nor  do  I  see  that  there  is  any 
other  general  cause  operating  on  the  seedlings  growing  wild  out  of  doors,  as  well  as  my  cul- 
tures in  bottles,  than  the  rise  in  temperature  which  is  one  factor  in  the  phenomena  of  spring. 


220  CALIFORNIA  ACADEMY  OF  SCIENCES        [Proc.  4th  ser. 

of  White  Oak  are  much  larger,  lobed,  thin  in  proportion  to  their  area,  and 
not  especially  waterproofed.  If  one  compare  the  woods  of  these  two  species 
of  oaks  one  finds  tlie  xylem  of  AVhite  Oak  containinia:  ducts  of  fairly  large  size, 
whereas  that  of  Live  Oak  is  made  up  predominantly  of  tracheids.  The  wood 
of  the  latter  carries  enough  water  from  the  roots,  even  in  dry  soil  at  the  end . 
of  the  dry  season,  to  keep  the  leaves  from  excessive  drying,  and  from  falling 
before  the  new  crop  of  leaves  emerges  in  the  spring. 

The  wood  of  the  conifers,  as  is  well  known,  is  composed  mainly  of  tracheids 
which,  in  proportion  to  their  diameters,  have  thick  walls.  In  these  thick  walls 
much  water  is  tenaciously  held.  The  majority  of  conifers  are  also  evergreen, 
retaining  their  leaves  for  two  or  more  years.  Those  which  are  not  evergreen 
live  where  there  is  ample  moisture  in  soil  and  air.  The  evergreen  conifers  are 
distributed  from  well-watered  to  desert  areas.  Their  monotonous  wood  anatomy 
enables  them  to  meet  this  great  range  of  circumstances.  The  vascular  systems 
of  the  conifers,  all  built  upon  the  same  general  plan,  are  equal  to  the  great 
variety  of  conditions  under  which  different  species  and  individuals  live.  But 
the  vascular  systems  are  associated  with  leaves  of  limited  area,  limited  water 
loss,  and  even  limited  number.  The  vascular  systems  can  carry  enough  water 
to  supply  the  leaves  at  all  seasons.  Only  in  the  severest  of  dry  seasons,  in  years 
of  light  winter  snowfall  and  of  rainless  summers,  does  one  see  signs  of  suffer- 
ing from  lack  of  water.  In  such  seasons  the  plants  absorb  what  water  they  can 
from  dry  soil ;  they  lose  what  they  must  from  leaves  restricted  in  area  and  in 
number — many  needles  fall  during  such  summers — ;  they  make  limited 
growth  in  length  and  may  even  die  back  extensively.  In  spite  of  the  superi- 
ority of  their  vascular  systems,  one  finds  the  desert  pines  as  scattered  indi- 
viduals, not  as  close  stands  of  forest  or  even  chaparral. 

The  amount  of  precipitation  and  the  anatomy  of  the  plants  living  in  a  given 
area  are  two  of  the  factors  which  determine  the  character  of  the  vegetation, 
limiting  a  natural  area  to  grassland,  chaparral,  or  forest,  and  correspond- 
ingly limiting  what  man  can  do  in  the  area.  Attempts  to  establish  forest,  with- 
out irrigation  and  other  care,  in  regions  of  limited  rainfall  and  naturally 
supporting  only  grass  or  even  chaparral,  seem  unlikely  to  succeed  because 
the  water-carrying  capacity  of  the  wood  is  unequal  to  the  need  of  moving 
water  fast  enough  from  the  roots  in  dry  soil  to  leaves  in  dry  air  at  any  but 
limited  heights  above  ground.  The  height  to  which  a  plant  can  grow  is  the 
height  to  which  its  vascular  system  can  carrj^  water.  This  is  determined  by  its 
wood  anatomy — which  is  inherited — and  by  its  environment.  The  relation  of 
the  anatomy  to  water  is  the  dominant  factor,  in  our  environment  with  its  long 
dry  seasons,  in  the  complex  which  is  the  living  thing. 


PROCEEDINGS 

OF  THE 

CALIFORNIA  ACADEMY  OF  SCIENCES 

Fourth  Series 
\'ol.  XXV,  No.  8,  pp.  221-234,  pis.  23-24  Jiii^e  1,  1944 


ALICE  EASTWOOD 
SEMI-CENTENNIAL  PUBLICATIONS 

No.  8 

A  PLEISTOCENE  FLORA  FROM  THE  McKITTRlCl^ 
ASPHALT  DEPOSITS  OF  CALIFORNIA 

BY 

HERBERT  L.  MASON 

Associate  Professor  of  Botany  and  Curator  of  the  Herbarium 
University  of  California 

THE  KNOWN  Pleistocene  floras  of  California  are  largely  coastal  in  their  posi- 
tion and  contain  a  group  of  species  as  fossils  that  are  essentially  maritime 
in  their  occurrence  today.  Of  these  by  far  the  richest  as  to  species  content  are 
the  Tomales  flora  (1)  in  Marin  County  and  the  Carpinteria  flora  (2)  in  Santa 
Barbara  County.  Both  of  these  contain  a  Monterey  type  forest  dominated  by 
pines  and  containing  cypresses  and  live  oaks  as  well  as  an  understory  of  shrubs 
that  is  highly  diversified  as  to  genera  and  species.  The  asphalt  deposit  at 
Rancho  La  Brea  (3)  in  Los  Angeles  County,  although  near  the  coast,  contains 
a  flora  that  is  essentially  interior  in  its  nature  and  is  best  represented  today 
by  the  open  woodland  vegetation  on  the  slopes  of  the  Tehachapi  Range.  The 
occurrence  in  this  deposit  of  a  few  specimens  of  maritime  species,  however, 
suggests  that  at  least  a  part  of  the  drainage  basin  contributing  to  this  deposit 
Avas  populated  by  a  maritime  flora.  It  seems  most  likely  that  a  mountain  range 
such  as  the  Santa  Monica  Moitntains  lay  between  it  and  the  sea  and  that  the 
coastal  slope  of  these  mountains  was  covered  by  a  maritime  flora  which  ex- 
tended over  the  top  and  a  short  distance  down  the  interior  side  much  as  does 
the  coastal  pine  forest  today  on  Cedrus  Island  off  the  Lower  California  coast. 
Only  a  very  few  strictly  interior  floras  of  Pleistocene  age  in  California  are  as 
yet  known  and  these  contain  very  few  species  of  plants.  We  therefore  know 
very  little  as  to  the  nature  of  the  interior  California  floras  during  Pleistocene 
time  and  hence  welcome  any  information,  however  meager,  that  will  help  us 
to  piece  together  a  picture  of  the  vegetation  of  the  interior  for  this  period  of 
the  earth's  history. 

[221] 


222  CALIFORNIA  ACADEMY  OF  SCIENCES        [Proc.  4th  ser. 

In  the  asphalt  deposits  at  McKittriek  in  western  Kern  County,  California, 
such  a  fossil  flora  occurs.  The  jMcKittrick  deposit  is  located  at  the  southwest- 
ern edg-e  of  a  small  oil  town  of  the  same  name  in  the  southwestern  San  Joaquin 
Valle}^  just  east  of  the  base  of  the  Temblor  Range.  The  elevation  of  McKittriek 
is  300  feet  while  the  Temblor  Range  rises  to  about  4000  feet.  The  geology  of 
the  area  has  been  reported  by  Arnold  and  Johnson  (4)  and  will  not  be  further 
treated  here.  The  rich  fossil  faunas  have  been  described  and  discussed  by 
Merriam  and  Stock  (5,  6)  for  the  mammals  and  by  Miller  (7,  8)  for  the  birds. 
Summaries  of  these  latter  papers  are  presented  by  Hay  (9).  Arguments  as 
to  the  precise  position  of  these  beds  in  the  Pleistocene  have  been  presented 
and  discussed  at  length  by  these  authors.  The  plant  species  that  are  at  all 
critical  in  this  respect  all  go  back  to  the  Pliocene  both  north  and  south  of  this 
area  and  are  still  living  both  north  and  south  of  this  area  and  hence  can  throw 
very  little  light,  if  any,  upon  this  problem.  The  Pleistocene  status  of  the  flora 
is  inferred  from  the  associated  fauna  which  is  largely  made  up  of  now  extinct 
species. 

The  flora,  however,  does  have  a  contribution  to  make  as  to  the  general  eco- 
logical setting  and  can  serve  to  a  large  extent  in  forming  the  basis  for  an 
interpretation  of  the  habitat  and  the  local  climate  for  the  portion  of  the  Pleis- 
tocene represented.  It  will  fill  out  the  ecological  picture  presented  in  part  by 
the  mammalogist  and  in  part  by  the  ornithologist. 

Merriam  and  Stock  (5,  6)  report  the  occurrence  of  horses,  bison,  camels, 
antelope,  bear,  lion,  tiger,  wolf,  ground  sloth  and  mastodon,  all  of  which  are 
now  extinct.  These  are  largely  plains  or  savanna  types  of  animals.  Miller 
reports  a  list  of  thirty-five  birds  of  which  water  birds  predominate  by  two  to 
one.  Only  10  per  cent  of  the  birds  are  now  extinct. 

To  date  six  identifiable  plant  species  have  been  found.  These  consist  of  two 
gymnosperms  and  four  angiosperms.  Although  the  flora  is  meager  these  plants 
are  sufficiently  critical  as  to  their  climatic  tolerance  to  serve  as  indicators  of 
the  nature  of  the  flora  and  of  the  climate  that  prevailed  in  the  McKittriek 
region  during  Pleistocene  time.  It  may  be  that  some  future  excavations  will 
yield  a  flora  comparable  in  quantity  as  well  as  in  quality  to  other  asphalt 
deposits  in  California  but  for  the  present  we  must  be  content  with  Avhat 
there  is. 

The  plants  are  represented  in  the  deposit  by  fragments  of  poorly  preserved 
fruiting  structures  and  seeds  as  well  as  scraps  of  partially  disintegrated 
twigs.  These  are  imbedded  in  sand  that  has  been  impregnated  with  asphalt. 
The  advanced  state  of  deterioration  of  so  much  of  the  plant  material  suggests 
that  the  impregnation  of  the  asphalt  occurred  after  the  plants  were  deposited 
and  after  preliminary  decomposition  had  set  in.  There  is  not  the  evidence 
of  a  pool  of  asphalt  such  as  is  suggested  by  the  conditions  at  Ranclio  La  Brea. 
As  compared  with  equal  volume  of  matrix  from  other  asphalt  deposits  the 
plant  material  in  the  McKittriek  is  rare.  Most  of  the  species  are  represented 
by  single  specimens.  In  onlj^  one  case,  the  Atriplex,  is  there  sufficient  material 


Vol.  XXV]        MASOX:  PLEISTOCENE  McKITTEICK  FLOSA  223 

to  give  any  notion  as  to  the  range  of  morphological  variation  of  the  structure. 
Fortunately,  however,  most  of  the  material  is  of  such  a  nature  as  to  be  readily 
identifiable  and  the  following  species  may  be  recorded  : 

Pinus  monophylla  Torr. 
Juniperus  calif ornica  Carr. 
Atriplex  lentiformis  (Torr.)  Wats. 
Arctostaphylos  glauca  Lindl. 
Arctostaphylos  ptmgens  H.B.K. 
Echinocystis  fahacea  Naud. 

All  of  the  above  species  found  in  the  McKittrick  flora  are  still  living  in  the 
California  flora  today.  The  occurrence  of  all  of  them  in  a  single  Pleistocene 
flora  presents  a  harmonious  ecological  picture  that  is  not  unlike  an  assemblage 
in  the  modern  flora  of  the  eastern  end  of  the  San  Rafael  Mountains  in  north- 
eastern Santa  Barbara  County,  on  the  slopes  of  Santa  Barbara  Canyon  a  little 
above  its  entrance  upon  the  Cuyama  Valley.  (United  States  Geological  Sur- 
vey, Santa  Ynez  Quadrangle,  R  25  W,  T  9  N,  in  the  Santa  Barbara  National 
Forest.) 

The  San  Rafael  Mountains  in  this  vicinity  range  from  altitude^s  slightly  in 
excess  of  600  feet  to  altitudes  near  7000  feet.  The  great  differences  in  altitude 
and  the  roughness  of  the  terrain  are  reflected  in  great  local  diversity  of  the 
flora.  A  profile  section  of  the  San  Rafael  Range  in  this  region  in  the  direction 
of  the  storm  paths  will  help  to  explain  why  a  region  so  close  to  the  coast 
supports  such  an  arid  type  of  vegetation.  There  are  three  prominent  ridges 
separated  by  the  canyons  of  Manzana  Creek  and  the  Sisquoc  River.  The  west- 
ernmost of  these  ridges  reaches  an  altitude  of  4000  feet  at  Zaca  Peak  directly 
above  the  1500-foot  Santa  Ynez  Valley.  The  canyon  of  Manzana  Creek  drops 
precipitously  to  1700  feet.  The  ridge  between  this  canyon  and  the  canyon  of 
the  Sisquoc  River  rises  to  a  height  of  3500  feet  and  is  clothed  with  a  very 
arid  type  of  vegetation.  The  canyon  of  the  Sisquoc  drops  to  2300  feet  and  the 
third  and  highest  ridge  culminates  in  Pine  Mountain  at  6800  feet  with  a 
northward  spur  of  about  5800  feet  forming  the  west  wall  of  Santa  Barbara 
Canyon.  These  ridges  serve  to  force  the  moisture-laden  winds  from  the  ocean 
upward  and  causes  them  to  deposit  their  load  on  the  westernmost  ridge  and 
the  upper  part  of  the  easternmost  high  ridge.  This  condition  is  reflected  in 
the  vegetation  of  these  ridges.  The  western  slope  of  the  first  ridge  is  clothed 
with  a  savanna  type  of  cover  except  where  the  topography  causes  rapid  run- 
off. Here  the  vegetation  is  chaparral.  The  top  of  this  ridge  is  clothed  with  a 
forest  of  big  cone  spruce  (Pseudotsuga  macrocarpa),  Coulter  pine  {Pinus 
Coulteri)  and  western  yellow  pine  {Pinus  ponderosa),  which  runs  down  the 
canyons  and  gullies  on  the  east  side.  The  second  ridge,  as  was  pointed  out 
above,  is  clothed  with  a  very  sparse,  arid  type  of  chaparral.  The  third  ridge 
is  arid  below  the  4000-foot  level  but  above  is  clothed  with  a  very  rich  conif- 
erous forest  of  big  cone  spruce.  Coulter  pine,  western  yellow  pine  and  sugar 
pine  {Pinus  Lamheriiana).  It  is  the  lower  slopes  of  the  east  side  of  this  ridge 


224  CALIFOBNIA  ACADEMY  OF  SCIENCES        [Proc.  4th  ser. 

that  contain  the  flora  represented  in  the  ^k-Kittrick  asphalt  deposit.  Here  is 
an  arid  type  of  flora  of  scattered  stands  of  pinyon  pine  (Pinns  monophylla) 
and  California  juniper  {Juniperus  calif ornica)  and  patches  of  manzanitas 
{Arctostaplnjlos  glauca,  A.  pungens  and  A.  Parryana).  The  other  species 
represented  in  the  fossil  flora  are  abundant  on  the  lower  slopes  and  in  the 
Pleistocene  were  probably  much  closer  to  the  site  of  deposition  than  were  the 
larger  shrubs  and  trees. 

Precisely  where  the  particular  plants  that  found  their  way  to  preservation 
in  the  McKittrick  asphalt  grew  during  Pleistocene  time,  is  difficult  to  deter- 
mine. It  seems  probable,  however,  that  they  were  a  part  of  the  flora  that  occu- 
pied the  Pleistocene  Temblor  Range  and  were  borne  down  by  the  streams  to 
the  ponds  and  lakes  that  ultimately  became  silted  up  and  infiltrated  with 
asphalt.  The  pine,  the  juniper  and  the  manzanitas,  as  well  as  the  Echinocystis, 
probably  occupied  the  higher  slopes,  while  the  Atriplexocc\i]}ied  alkaline  flats 
near  the  ponds  and  marshes.  This  occurrence  would  explain  the  abundance 
of  Atriplex  and  the  relative  scarcity  of  the  other  species  in  the  deposit,  since 
these  latter  would  have  to  be  transported  by  a  none  too  rich  stream  flow. 

Climatically  the  flora  suggests  an  area  of  low  rainfall  but  perhaps  more 
than  is  received  by  the  McKittrick  region  at  the  present  time.  This  might  lead 
some  to  argue  that  the  flora  indicates  at  least  glacial  time ;  however,  in  view 
of  tlie  close  proximity  of  a  similar  modern  assemblage  of  species  it  is  more 
logical  to  assume  that  some  minor  changes  in  topography  to  the  west  would 
have  accomplished  the  same  result  during  the  Pleistocene  at  McKittrick.  If 
the  Temblor  Range  were  higher  by  1000  feet,  it  might  support  such  a  flora 
around  its  summit.  Evidence  as  to  glacial  or  interglacial  time  is  negative  so 
far  as  the  McKittrick  flora  is  concerned. 

Ecologically  the  flora  is  entirely  consistent  with  the  assemblage  of  mammals 
and  of  the  birds.  It  was  sufficiently  open  to  permit  grazing  and  browsing  and, 
although  the  presence  of  Atriplex  does  not  necessarily  mean  ponds  and  lakes, 
it  frequently  occurs  near  such  sites  today. 

Since  all  of  the  plants  represented  in  the  McKittrick  flora  are  modern 
species  the  nomenclature  here  used  is  that  of  the  modern  plants.  In  one  of 
the  plants  the  nomenclature  of  the  modern  group  is  not  as  yet  stabilized, 
owing  to  incomplete  knowledge  and  hence  uncertain  taxonomic  concepts.  In 
this  instance  it  is  felt  that  it  is  not  within  the  province  of  this  paper  to  solve 
these  problems,  nor  could  they  be  solved  on  the  basis  of  the  material  here 
presented.  The  nomenclature  for  this  species  therefore  follows  current  usage. 
The  problem  will  be  raised  in  connection  with  the  species  in  question. 

The  specimens  here  cited  are  all  deposited  in  the  Palaeobotanical  Collection 
of  the  University  of  California  at  Berkeley. 


VOL.XXV]        MASON:  PLEISTOCENE  McKITTEICK  FLOBA  223 

Pinus  monophylla  Torr. 

(Plate23,  fig.  3) 
Pinus  monophylla  Torr.  in  rremont,  Second  Eeport,  319,  1845. 

Pinus  monophylla  is  represented  by  a  cone,  of  relatively  large  size  for  this 
species,  that  is  poorly  preserved.  The  structure  is  practically  reduced  to  vas- 
cular tissue  but  enough  of  the  surface  features  of  a  few  of  the  scales  is  pre- 
served to  show  the  essential  nature  of  the  apophysis  and  of  the  umbo  as 
well  as  to  clearly  indicate  the  size  and  shape  of  the  seed  cavity.  The  cone  is 
about  8  em.  long  and  about  7  cm.  wide.  The  scales  are  terminated  by  a  some- 
what recurved,  quadrate  apophysis  bearing  a  dorsal  umbo.  The  seed  cavity  is 
ovoid,  about  12  mm.  long  and  8  mm.  wide,  indicating  a  relatively  large  seed. 

Pi7ius  monophylla  ranges  primarily  to  the  east  of  the  Sierra  Nevada.  A 
few  stations  have  been  recorded  for  its  occurrence  on  the  west  slopes  of  the 
Sierra.  It  occurs  also  in  the  Tehachapi  and  the  Mount  Pinos  region,  and 
extends  northward  in  the  coast  ranges  to  the  Cuyama  Valley  in  the  San  Rafael 
Mountains,  and  on  the  watershed  of  the  headwaters  of  the  Santa  Ynez  River. 

University  of  California  Palaeobotanical  Collections,  Plesiotype  no.  2810. 

Juniperus  californica  Carr. 

(Plate23,  fig.  1) 

Juniperus  californica  Carr.  Eev.  Hort.  352,  1854. 

Juniperus  californica  Carr.  var.  breaensis  Frost,  Univ.  Calif.  Publ.  Bot.  14:77,  1927. 

Juniperus  californica  Carr.  is  represented  in  the  fossil  flora  by  a  seed  8  mm. 
long  by  5  mm.  wide  that  in  no  way  differs  from  seed  of  the  modern  species  as 
it  occurs  in  the  Tehachapi  Mountains  today.  The  seed  coat  is  of  medium  thick- 
ness, and  agrees  with  other  Pleistocene  material  collected  from  the  asphalt 
deposits  at  Rancho  La  Brea  and  described  by  Frost  as  distinct  from  the 
modern  plants. 

The  modern  J.  californica  Carr.  is  an  aggregate  species  that  includes  with 
varietal  rank  the  desert  and  Great  Basin  form  J.  utahensis  (Engelm.)  Lem- 
mon.  These  two  forms  intergrade  in  the  region  of  the  Tehachapi  Mountains 
so  as  to  make  their  separation  as  distinct  species  impossible.  They  have  been 
separated  on  the  basis  of  the  glandular  nature  of  the  foliage  of  typical  /. 
californica.  Another  important  difference  is  the  relative  thickness  of  the  seed 
coat  in  the  extremes  of  the  two  forms.  Typical  /.  calif orriica  has  a  thin  seed 
coat  while  typical  /.  californica  var.  utahensis  has  a  very  thick  seed  coat.  Both 
the  characters  of  the  seed  coat  and  of  the  glandulosity  intergrade  in  material 
collected  by  the  writer  from  the  Tehachapi  Mountains  and  from  the  Sierra 
Liebre  in  California  (see  Mason  3737,  Herbarium  of  the  University  of  Cali- 
fornia). The  fossil  material  from  the  McKittrick  asphalt  is  of  the  intermediate 
type  represented  by  this  material.  Because  of  its  intermediate  character  it 
is  impossible  to  designate  whether  it  belongs  to  the  species  or  to  the  variety. 

In  the  fossil  record  J.  californica  has  been  reported  from  tlie  asphalt  depos- 


0Qiq^ 


.'^xrJ^^*^ 


226  CALIFOENIA  ACADEMY  OF  SCIENCES        [Proc.  4th  ser. 

its  at  Carpinteria  and  at  Rancho  La  Brea.  In  the  latter  flora,  however,  the 
material  was  described  as  new  under  the  name  J.  calif ornica  var.  hreaensis 
Frost.  J.  californica  ranges  from  Lower  California  through  the  western 
]\Iojaye  Desert  to  the  Tehachapi  Mountains  and  northward  in  discontinuous 
patches  in  the  inner  coast  ranges  to  Shasta  County  and  sparingly  on  serpen- 
tine soil  in  the  Sierra  Nevada  footliills  to  Butte  County.  It  is  of  interest  to 
the  present  paper  to  record  it  in  abundance  in  the  inner  south  coast  ranges 
that  border  the  Cuyama  Valley. 

L'niversity  of  California  Palaeobotanical  Collections,  Plesiotype  no.  2811, 

Atriplex  lentif onnis  (Torr.)  Wats. 

(Plate  24,  figs.  1,  2,  3) 
Atriplex  lentif ormis  (Torr.)  Wats.  Proc.  Am.  Acad.  9:118,  1874. 

The  species  is  represented  by  a  large  number  of  fruiting  bracts  often  enclos- 
ing fragments  of  the  seed  coats.  They  are  flattened  and  wedge-shaped.  Some  | 
of  them  possess  a  stipe  while  others  appear  to  have  been  sessile  on  the  branch. 
They  average  4  mm.  in  length  and  5  mm.  in  M'idth.  The  bracts  are  united  below] 
over  the  seed  and  are  apparently  free  above.  The  upper  margins  are  somewhat  ] 
broken  and  it  is  difficult  to  reconstruct  the  character  of  these  organs.  There  l 
is  evidence  to  indicate  a  fringed,  denticulate  margin.  The  surface  shows  aj 
coarsely  reticulate  venation  and  is  entirely  devoid  of  any  facial  processes. 

It  seems  evident  from  a  careful  comparison  of  the  specimens  that  but  a 
single  species  of  the  genus  is  represented.  The  great  range  of  variation  exhib- 
ited by  many  of  the  modern  species  shows  forms  that  are  not  unlike  the  fossils. 
How^ever,  each  species  has  a  characteristic  kind  of  fruiting  structure  and 
its  variation  centers  around  this  type.  Of  the  modern  species  now  living  in  j 
California,  A.  lentif  ormis  seems  closest  both  on  morphological  and  ecological 
grounds.  The  area  Avherein  this  flora  is  best  represented  today  is  rich  in 
species  of  Airiplex  and  any  or  all  of  them  might  well  have  been  living  in  the 
McKittrick  locality  during  the  Pleistocene. 

The  genus  Atriplex  has  been  recorded  from  the  Pleistocene  Tomales  flora] 
of  central  California  where  fossil  material  of  the  coastal  species  of  A.  hastata^ 
has  been  collected. 

University  of  California  Palaeobotanical  Collections,  Plesiotype  no.  2812. 

Arctostaphylos  glauca  Lindl. 

(Plate24,  figs.  4,  5,  6) 
Arctostaphylos  glauca  Lindl.  Bot.  Reg.  sub  t.  1791,  1835. 

This  large-fruited  species  of  manzanita  is  represented  in  the  fossil  flora  by 
frag-ments  of  fruits  one  of  which  is  12  mm.  in  diameter,  with  its  carpels  closely 
coalesced  so  as  to  present  a  very  smooth  surface.  The  material  is  identical  Avith 
similar  structures  on  the  modern  A.  glauca  and  is  clearly  referable  to  that 
species. 


VOL.XXV]        MASON:  PLEISTOCENE  McKITTRICE  FLORA  227 

The  modern  plant  is  a  characteristic  species  of  the  Upper  Sonoran  life  zone 
in  the  blue-oak-dig:ger  pine  belt,  and  ranges  from  central  California  south- 
ward to  southern  California.  It  is  frequent  in  the  Tehachapi  Mountains  and 
the  inner  south  coast  ranges  in  association  with  juniper.  Its  occurrence  in  the 
fossil  flora  suggests  conditions  similar  to  those  of  these  mountains  at  an 
altitude  of  about  3000  feet. 

Arctostaphylos  glauca  is  knowai  in  the  Pleistocene  from  both  the  Carpin- 
teria  and  the  Rancho  La  Brea  asphalt  deposits.  It  is  an  indicator  of  arid 
interior  conditions  (Chancy  and  Mason,  2) . 

University  of  California  Palaeobotanical  Collections,  Plesiotype  no.  2813. 

Arctostaphylos  pungens  H.B.K. 

Arctostaphylos  piingens  H.B.K.  Nov.  Gen.  et  Sp.  3  :278,  259,  1818. 

This  species  is  represented  in  the  fossil  flora  bj'  a  fruit  4  mm.  long  and  6 
mm.  wide  with  coalesced  carpels.  The  individual  carpels  show  unequal  devel- 
opment and  present  a  strongly  ribbed  appearance  to  one  side  of  the  fruit. 
The  specimen  is  not  unlike  many  that  are  of  frequent  occurrence  on  modern 
plants  of  Arctostaphylos  pungens  of  the  inner  south  coast  ranges. 

Arctostaphylos  pungens  is  a  species  of  the  arid  hills  bordering  on  the  upper 
part  of  the  Sonoran  and  the  lower  part  of  the  Transition  zones  from  central 
California  southward.  Its  precise  range  cannot  at  present  be  stated  because 
it  is  obvious  that  what  has  been  going  under  the  name  of  A.  pungens  in  current 
floras  of  the  region  is  not  the  same  as  the  plant  originally  described  under 
that  name  from  the  vicinity  of  Mexico  City.  Furthermore,  there  is  consider- 
able confusion  about  the  taxonomy  of  the  group  locally  and  until  this  problem 
can  be  straightened  out  by  a  monographer  of  the  genus  as  a  whole  the  precise 
name  must  stand  in  abeyance.  It  is  possible  that  A.  montana  Eastwood  and 
A.  Hookeri  Don  may  be  involved  in  relationship  here  to  such  an  extent  as  to 
invoh'e  the  nomenclature  of  the  fossils.  At  any  rate,  the  specimen  under  con- 
sideration is  of  a  type  like  the  modern  plants  of  this  group  now  growing  in 
the  inner  south  coast  ranges  of  California  and  referred  to  in  modern  floras 
as  A.  pungens. 

University  of'  California  Palaeobotanical  Collections,  Plesiotype  no.  2814. 

Echinocystis  fabacea  Naud. 

(Plate  24,  figs.  8,  9) 
Echinocystis  fahacea  Naud.  Ann.  Sci.  Nat.  ser.  IV,  12 :  154,  1859. 

Fragments  of  the  seed  coat  of  this  species  preserved  in  this  deposit  differ 
in  no  way  from  the  modern  species  that  is  so  widespread  in  the  inner  and 
outer  coast  ranges  of  California.  Identification  was  based  upon  the  histologi- 
cal nature  of  the  seed  coat,  where  there  are  tw^o  very  different  layers  of  cells 
evident,  an  inner  layer  composed  of  narrow  cylindrical  cells  and  an  outer 
layer  about  three  times  as  thick  composed  of  manv  irregularly  rounded  cells,-— r-3r—-s-. 


228  CALIFORNIA  ACADEMY  OF  SCIEXCES        [Proc.  4th  ser. 

The  inner  face  of  tlie  seed  coat  is  lined  with  a  very  soft  spongy  tissue  that 
readily  sloughs  off.  Echitiocystis  fahacea  ranges  throughout  the  inner  and 
outer  coast  ranges  of  central  California,  crossing  the  Tehachapi  Mountains  to 
the  Mojave  Desert,  where  it  has  been  reported  in  tlie  vicinity  of  Muroc.  It  is 
not  common  in  the  Sierra  Nevada  foothills,  but  is  abundant  in  the  inner  south 
coast  ranges. 

University  of  California  Palaeobotanical  Collections,  Plesiotype  no.  2815. 

LITERATURE  CITED 

1.  Mason,  H.  L.  Carnegie  Institution  of  Washington  Publication  no.  415 :  83-179.  1934. 

2.  Chanet,  R.  W.,  and  H.  L.  Mason.  Carnegie  Institution  of  Washington  Publication  no. 

415:47-79.1933. 

3.  Frost,  F.  H.  University  of  California  Publications  in  Botany  14 :  73-98.  1927. 

4.  Arnold,  R.,  and  H.  R.  Johnston.  Bull.  U.  S.  Geological  Survey  no.  406:1-225.  1910. 

5.  Merriam,  J.  C,  and  C.  Stock.  Science,  n.  s.  54:566-567.  1921. 

0. .  Carnegie  Institution  of  Washington  Publication  no.  347:11,  38—44.  1925. 

7.  Miller,  L.  II.  Condor  24 :  122-125.  1922. 

8. .  University  of  California  Department  of  Geology,  Bull.  15 :  307-326.  1925. 

9.  Hay,  O.  P.  Carnegie  Institution  of  Washington  Publication  no.  322B:  196-199.  1927. 


EXPLANATION  OF  PLATES 


PLATE  23 

Gymnosperms  in  the  McKittriek  Flora 

Fig.  1.  Juniperus  calif ornica  Carr.,  drawing  of  the  fossil,  x  6.  Univ.  Calif. 
Palaeobot.  Coll.,  Plesiotype  no.  2811. 

Fig.  2.  Seed  of  a  modern  plant  of  Juniperus  calif  ornica  Carr.,  x  1. 

Fig.  3.  Cone  of  Pinus  monophylla  Torr.  x  1.  Univ.  Calif.  Palaeobot.  Coll., 
Plesiotype  no.  2810. 


[230] 


PROC.  CALIF.  ACAD.  SCI..  4TH  SERIES.  VOL.  XXV.  NO.  8 


[MASON]  PLATE  23 


[231] 


PLATE  24 

Angiosperms  in  the  McKittrick  Flora 

Figs.  1,  2,  3.  Atriplex  lentiformis  (Torr.)  Wats.  Fruiting  bracts,  x  10.  Univ. 
Calif.  Palaeobot.  Coll.,  Plesiotype  no.  2812. 

Figs.  4,  5,  6.  Arciostaphylos  glauca  Lindl.  Fragments  of  fruit.  Figs.  4  and 
5,  X  4 ;  fig.  6,  X  5.  Univ.  Calif.  Palaeobot.  Coll.,  Plesiotype  no.  2813. 

Fig.  7.  Arciostaphylos  pungens  H.B.K.  Endocarp  of  fruit,  x  8.  Univ.  Calif. 
Palaeobot.  Coll.,  Plesiotype  no.  2814. 

Fig.  8.  Echinocystis  fabacea  Naud.  Cross  section  of  the  seed  coat  of  a  mod- 
ern specimen  showing  the  two  layers  of  the  seed  coat,  x  15. 

Fig.  9.  Echinocystis  fabacea  Naud.  Cross  section  of  the  seed  coat  of  the  fos- 
sil, X  15.  Univ.  Calif.  Palaeobot.  Coll.,  Plesiotype  no.  2815. 


[  232  ] 


PROC.  CALIF.  ACAD.  SCI..  4TH  SERIES,  VOL    XXV.  NO.  8 


[MASON]  PLATE  24 


[  233  ] 


PROCEEDINGS 

OF  THE 

CALIFORNIA  ACADEMY  OF  SCIENCES 

Fourth  Series 
\'ol.  XX\^  No.  9,  pp.  235-244,  6  text  figures  Jiine  1 .  1944 


ALICE  EASTWOOD 
SEMI-CENTENNIAL  PUBLICATIONS 

No.  9 

THE  PERENNIAL  SOUTHWESTERN  DATUR 
AND  THE  VALIDITY  OF  MATTHEW'S 
HYPOTHESIS  IN  PLANT  GEOGRAPHY 

BY 

JOSEPH  EWAN 

Instructor  in  Biology,  Curator  of  Botany,  University  of  Colorado 

EVIDENCES  OF  plivlogeiiy  among  major  plant  groups  rest  upon  two  lines  of 
observational  research,  viz.,  comparative  morphology  and  paleobotany, 
and  upon  two  methods  of  experimental  research,  viz.,  serum  diagnosis  and 
cytogenetics.  Among  species  within  a  single  genus  evidences  of  phylogeny 
rest  again  upon  comparative  morphology  in  the  broadest  sense,  along  with 
breeding  experiments  and  cytogenetics,  with  either  supporting  evidence  or 
the  lack  of  it  from  the  fields  of  ecology  and  plant  geography.  Interpretation 
of  morphology  is  thrown  into  sharper  perspective  by  subjecting  the  data  to 
Haeckel's  Biogenetic  Law,  Dollo's  Law  of  Irreversibility,  Darwin's  evolu- 
tionary theory,  Liebig's  Law  of  the  Minimum  as  applied  to  distribution  prob- 
lems. Good's  Theory  of  specific  tolerance,  Jordan's  Law  of  Geminate  Species, 
Eames'  concept  of  "phyletic  slides"  within  plant  groups,  and  what  I  have 
chosen  to  call  Matthew's  Hypothesis  of  Peripheral  Populations.  The  subject 
of  the  present  paper  is  a  consideration  of  Matthew's  Hypothesis  as  it  may 
prove  valid  in  interpreting  distribution  phenomena  in  Datura  metdoides  DC. 
In  his  now  classic  paper  entitled  "Climate  and  Evolution"  (1915)  the  Amer- 
ican vertebrate  paleontologist,  W.  D.  Matthew,  presented  a  working  hypothe- 
sis for  the  evolution  of  the  higher  vertebrates.  He  devoted  three  paragraphs 
in  the  beginning  to  his  "Principles  of  Dispersal,"  with  which  we  are  chiefly 
concerned.  His  evolutionary  theory  embraced  the  concept  that  the  evolution 
of  land  life  has  responded  in  adaptation  to  recurrent  periods  of  aridity 

[  23.5  ] 


236  CALIFORNIA  ACADEMY  OF  SCIENCES        [Proc.  4th  ser. 

through  geologic  time.  In  short,  Matthew  demonstrated  that  the  major  ad- 
vances of  vertebrate  evolution  have  coincided  with  shifts  in  megaclimates, 
especially  those  shifts  toward  uicreasing  aridity.  It  is  a  significant  corollary 
of  Matthew's  Hypothesis,  in  my  opinion,  that  the  angiosperms  of  arid  and 
semi-arid  regions  exhibit  conspicuous  evolutionary  advance  in  their  mor- 
phology over  their  relatives  of  mesic  regions. 

Matthew's  Hypothesis 

Matthew's  Hj'pothesis  holds  that, 

at  any  one  time  in  the  evolution  of  a  taxonomic  gro^ip  the  most  advanced  stages  should 
be  nearest  the  center  of  dispersal,  the  most  conservative  stages  farthest  from  it. 

Or,  to  use  the  words  of  another  vertebrate  paleontologist,  Prof.  R.  S.  Lull,  it 
may  be  stated  that, 

"the  most  ancient  members  of  a  group  are  not  to  be  found  at  the  old  center  of  evolution, 

but  rather  at  the  periphery  of  their  migratory  area."  (Lull  in  George  A.  Baitsell,  ed., 

Evolution  of  Earth  and  Man,  pp.  118-119.  1929.) 

In  his  original  paper  Matthew  points  out  that  the  evolution  of  a  race  "should 
'be  at  first  most  progressive  at  its  point  of  original  dispersal,  and  it  will  con- 
tinue this  progress  at  that  point  in  response  to  whatever  stimulus  originally 
caused  it  and  spread  out  in  successive  waves  of  migration,  each  wave  a  stage 
higher  than  the  previous  one"  (1915,  180).  Assuming  that  a  species  is  the 
product,  in  part,  of  its  environment,  and  that  environments  are  subject  to 
change,  "it  is  the  environment  itself,  biotie  as  well  as  physical,  that  migrates, 
and  primitive  species  are  those  which  have  followed  it,  while  those  which 
remained  have  had  to  adapt  themselves  to  a  new  environment  and  become 
altered  thereby."  However,  it  should  be  remembered  that  probably  "it  is 
never  the  case  that  the  environment  of  the  marginal  species  is  an  absolute 
replica  of  the  older  environment  of  the  race"  (1915,  180-181). 

The  American  plant  ecologist  Stanley  Cain  expressed  the  concept  in  1940 
as  follows :  "in  wide-ranging  species  populations  mere  distance  constitutes 
a  type  of  isolation  and  marginal  or  other  portions  of  a  population  are  fre- 
quently recognizable  .  .  .  without  the  development  of  any  conspicuous  dis- 
continuity" (19-40,  214).  Cain  points  out,  furthermore,  that  these  marginal 
populations  are  genetically  more  homozygous.  In  the  instance  of  Daiuva 
meieloides,  the  native  solanaceous  perennial  known  by  the  Indian  name 
"tolguacha,"  this  homozygosity  may  be  demonstrated  by  the  occurrence  of 
distinctive  leaf  types  of  representative  individuals  from  geographically  mar- 
ginal populations,  as  recorded  in  some  of  the  larger  herbaria. 

Materials  and  Procedure 

This  study  of  Datura  meieloides,  it  should  be  pointed  out  at  the  outset,  is 
based  Avholly  upon  herbarium  specimens  and  is  subject  to  confirmation  from 
an  even  more  statistical  approach,  especially  in  the  field  over  the  extensive 
geographic  range  of  the  species,  in  the  manner  of  Fassett's  "mass  collection" 


Vol.  XXV] 


EWAN:  DATUEA 


237 


studies  (1941).  It  may  be  of  interest  to  record  how  this  problem  arose.  De- 
Candolle's  original  description  of  Datura  meteloides  rests,  not  upon  the  usual 
preserved  herbarium  "type"  specimen  but  upon  a  Sesse  and  Mocirio  drawing, 
rather  unfortunate  for  having  been  hastily  copied  from  the  original.  The 
plant  from  which  the  drawing  was  made  was  collected  at  an  unrecorded  local- 


Fig.  1.  Maximum  natural  range  (generalized)  of  Datura  meteloides  DC. 


ity  in,  probably,  central  Mexico  by  Mocifio.  By  comparing  this  drawing  with 
modern  herbarium  collections  as  to  close  similarity  of  leaf  blade  outline  an 
attempt  was  made  to  fix,  ixi  a  general  way  at  least,  the  possible  geographic 
source,  that  is,  a  putative  "type  locality,"  of  the  material  which  served  as  the 
basis  of  what  must  be  considered,  in  lieu  of  an  actual  preserved  plant,  the  type 
of  the  species.  The  taxonomic  history  of  this  plant,  and  certain  problems  in 
the  systematics  of  it,  are  treated  elsewhere  by  the  present  author  (1944) . 


238 


CALIFORNIA  ACADEMY  OF  SCIENCES        [Proc.  4th  ser. 


This  paper,  then,  arose  from  an  examination  of  nearly  300  collections, 
primarily  from  two  large  herbaria,  the  U.  S.  National  Herbarium,  and  the 
Rocky  Mountain  Herbarium  of  the  University  of  Wyoming,  with  subsequent 
examinations  of  materials  in  the  herbaria  of  the  Academy  of  Natural  Sci- 
ences at  Philadelphia  and  the  Field  Museum.  To  the  curators  of  these  collec- 
tions I  am  grateful  for  the  privilege  of  studying  the  materials  in  their  charge. 
The  collections  were  sorted  according  to  closeness  of  fit  in  leaf  blade  config- 
uration. Four  chief  repetitions  in  leaf-form  were  encountered.  These  char- 
acteristic infraspecific  variations  in  leaf -form — including  equilateral  and 
inequilateral,  serrate  and  subentire  forms — were  then  plotted  on  outline  maps 
according  to  the  place  of  collection  as  recorded  on  the  herbarium  labels  of 
each.  ^lature  leaves  were  chosen  from  each  sheet  and  tracings  were  made  of 
representative  collections.  Almost  invariably  the  collections  were  of  flowering 
specimens,  and  the  leaves,  therefore,  associated  with  flowering  shoots.  It  was 
found  that  those  collections  showing  closely  comparable  leaf -forms  came,  for 
the  most  part,  from  geographically  limital  localities.  These  data  are  given 
below.  The  total  range  of  the  species  is  shown  in  a  generalized  way  by  the 
accompanying  map  (fig.  1). 


Data  from  Datura  meteloides 
Four  distinctive  groups  showing  close  repetition  of  leaf-form  are  listed 
below.  From  each  group  representative  collections  are  cited,  chosen  from 
widely  separated  and  geographically  peripheral  localities.  Pen  line  tracings 
and  map  positions  are  given  for  each  collection. 


Leaf-form 
inequilateral 
ovate  and 
subentire 


Leaf-form 
equilateral 
ovate  and 
serrate 


Leaf -form 
equilateral 
ovate  and 
subentire 

Leaf -form 
equilateral 
deltoid  and 
serrate 


''      Locality 
1.  San  Bernardino,  Calif 


Group  One  (fig.  2) 

Collector 


2.  Santa  Barbara,  Calif. 

3.  St.  George,  Utah 

4.  Valencia  Beach,  L.  Calif. 


G.  E.  Vasey  443 
W.  H.  Brewer  310 
M.  E.  Jones  6093 
Wiggins  and 
Gillespie  3948 

Group  Two  (fig.  3) 

Locality  Collector 

5.  Holbrook,  Ariz.  Osterhout  6767 

6.  Tarrant  Co.,  Texas  A.  Ruth  583 

7.  Fuerte,  Sinaloa,  Mex.  Eose,  Standley 

and  Eussell  13452 

Group  Three  (fig.  4) 
Collector 


< 


f        Locality 

8.  Santa  Ysabel,  Calif. 

9.  Colonia  Diaz,  Chih.,  Mex. 
in.  Buckskin  Mts.,  Ariz. 


Henshaw  68 

E.  W.  Nelson  6430 

Jones  6063n 


Group  Four  (fig.  5) 
Locality  Collector 

11.  Montezuma  Co.,  Colo.  Crandall  in  1895 

12.  Pasadena,  Calif.  G.  B.  Grant  270 

13.  Mazatlan,  Sinaloa,  Mex.        Ortega  5654 


Herb,  and  sheet  no. 
USNH  156906 
USNH  322527 
EM  14182 
USNH  1491044 


Herb,  and  sheet  no. 
EM  169488 
USNH  1523537 
USNH  636275 


Herb,  and  sheet  no. 
USNH  238095 
USNH  360146 
USNH  260638 

Herb,  and  sheet  no. 
USNH  963190 
EM  54824 
USNH  1208683 


Vol.  XXV] 


EWAN:  DATURA 


239 


Fig.  2.  Group  One  of  Datura  meteloides  leaf -forms. 


Fig.  3.  Group  Two  of  Datura  meteloides  leaf-forms. 


240 


CALIFORNIA  ACADEMY  OF  SCIENCES        [Proc.  4th  ser. 

L 


Fig.  4.  Group  Three  of  Datura  meteloides  leaf -forms. 


Fig.  5.  Group  Four  of  DaUira  meteloides  leaf -forms. 


Vol.  XXV] 


EWAN:  DATURA 


241 


Collections  from  the  more  central  portion  of  the  range  of  Datura  meteloides 
show  a  range  of  leaf-forms  well  represented  by  the  five  tracings  made  from 
Charles  Wright  526  from  Turkey  Creek,  Texas,  which  is  the  lectotype  of 
Datura  Wrightii  Kegel  (USNH  60043,  cf.  fig.  6,  tracings  14-18  inc.  all  from 
same  coll.).  Collections  closely  comparable  to  Wright  526  in  the  range  of 
variation  of  leaf -forms  have  been  made  at  the  stations  indicated  by  the  circles 


Fig.  6.  Center  of  distribution  of  Daiura  vieteloides  as  determined  by  conformity  in 
leaf -form,  with  five  leaves  (nos.  14-18)  from  single  coll.  {WrigM  526)  from  locality 
at  "14." 

in  fig.  6.  This,  then,  represents  the  biologic  center  of  the  species.  Doubtless 
the  species  will  be  found  to  be  present,  in  suitable  river  bottom  and  alluvial 
habitats,  in  northern  Mexico  as  well,  where  the  map  suggests  its  absence. 

Discussion 
The  center  of  distribution  for  Datura  meteloides  may  be  recognized  as  the 
Rio  Grande  catchment  basin  of  New  Mexico  westward  to  the  upper  Gila  River 
basin  of  Arizona.  This  conclusion  is  based  upon  a  noticeable  homogeneity  in 
the  populations  as  demonstrated  by  the  limited  range  of  variation  of  their 
leaf -forms.  The  peripheral  populations  are  considered,  therefore,  to  represent 
fragmentary  remnants  of  the  original  species  colony  persistent  today  as  a 
fringe  about  the  now  altered  population  mass  comprising  the  present  "center 
of  distribution." 


242  CALIFOBNIA  ACADEMY  OF  SCIENCES        [Proc.  4th  ser. 

Gleason  noted  the  agreement  of  marginal  members  of  a  group  about  a  center 
of  distribution  in  his  studj^  of  the  genus  Vernonia.  He  wrote,  "that  those  spe- 
cies most  nearly  alike  morphologically  .  .  .  ai-e  always  distributed  about  the 
same  center"  (1906, 152).  C.  H.  MuUer  found  .Matthew's  Hypothesis  to  obtain 
in  certain  species  of  Mexican  Quercus  (oral  communication ;  cf.  also  Am.  Midi. 
Nat.  18:  844,  1937).  In  my  studies  of  the  Calif ornian  flora  instances  which 
apparently  corroborate  Matthew's  Hypothesis  have  been  noticed  in  Monar- 
della  lanceolata  Gray,  where  small  capitulae  of  closely  comparable  form  and 
size  are  to  be  noticed  in  plants  of  Yosemite  Valley  on  the  north  and  San  Diego 
County,  California,  on  the  south.  In  the  mariposa  called  "lantern-of-the- 
fairies,"  Calochortus  alhus  DougL,  closely  parallel  small-flowered  forms  may 
be  recognized  at  almost  the  same  geographically  distant  localities  as  in  Monar- 
della,^  each  an  extreme  limital  population  of  the  species  range.  Among  the 
North  American  Delphiniums,  infraspecific  phases  have  been  detected  in  the 
Calif  ornian  Delphinium  Parry)  Gray  where  small-flowered  populations  occur, 
again,  at  limital  positions  over  the  total  range  of  the  species.  In  Delphinium 
patens  Greenei  (Eastw.)  Ewan,  a  very  restricted  subspecies  of  California, 
plants  with  identical  glandular  pedicels  occur  on  Marysville  Buttes  in  the 
northern  Great  Valley  on  the  north  and  from  Fresno  County  upper  foothills 
on  the  south,  with  non-glandular  plants  of  Delphinium  paiens  patens  occupy- 
ing the  interveniug  area. 

An  apparent  antithesis  of  Matthew's  Hypothesis  as  valid  for  Datura  mete- 
loides  occurs  in  Tradescantia  yvhere  the  peripheral  populations  have  been 
demonstrated  by  Anderson  and  Woodson  (1935)  to  be  derivitive  tetraploid 
infraspecific  races,  with  the  ancestral  diploid  races  occupying  the  center  of 
distribution.  These  infraspecific  races  are  indistinguishable  in  their  morphol- 
ogy. The  oldest  members  of  the  species  populations  are  central,  then,  and 
not,  as  in  Datura  meteloides,  marginal  to  the  mass  of  the  species.  In  Trad- 
escantia the  center  of  distribution  significantly  is  geologically  an  old  land 
mass  within  the  total  range  of  the  species.  In  Datura  meteloides  the  peripheral 
populations  occupy  extremely  diverse  regions  which  are  not  often  cognate 
geologically.  Furthermore,  these  peripheral  populations  are  not  morpho- 
logically homogeneous,  at  least  as  to  their  leaf -forms,  but  differ  within  them- 
selves from  the  much  more  uniform  plants  of  the  center  of  distribution. 
There  are,  in  fact,  four  distinctive  leaf -forms  existing  more  or  less  side  by 
side  within  this  marginal  population;  at  least  in  coastal  Southern  California 
three  of  the  four  leaf -forms  occur  together. 

In  the  European  crucifer  Biscutella  laevigata  L.  three  polyploid  infra- 
specific  races — a  diploid,  tetraploid  and  hexaploid  race — have  been  shown 
to  exist  by  Irene  Manton  (1937) .  Here  the  archaic  diploids  are  "xerothermal" 
interglacial  relicts  along  the  river  valleys  of  central  Europe  from  Germany 
to  the  Balkans,  nol'th  and  east  of  the  Alps.  The  tetraploid  plants  occupy  the 

^  "As  is  frequently  the  case  with  Lnhiatae,  the  extreme  forms  geoprraphically  are  often 
most  similar,  at  least  superficially."  (Epling,  A  Revision  of  Salvia:  Subgenus  Calosphace, 
191,1939). 


Vol.  XXV]  EWAN:  DATVEA  243 

Swiss  and  Austrian  Alps,  swept  over  by  glaciers,  as  a  subsequent  invading 
population.  Obviously  there  is  no  direct  relation  to  glaciation  in  the  distribu- 
tion of  Datura  meteloides. 

There  is,  however,  with  this  Datura  a  clear  floristic  affinitj^  to  the  Sierra 
Madrean  element  of  Southwestern  United  States  and  Mexico.  Axelrod's  map 
(1939,  59,  fig.  2B)  should  be  studied  in  this  regard.  This  map  shows  the 
"supposed  distribution  of  the  Sierra  Madrean  element  as  a  dominant"  at 
Lower  Pliocene  time.  This  extent  agrees  very  closely  with  the  extreme  geo- 
graphic range  enjoyed  by  Datura  meteloides  at  the  present  time.  Indeed,  the 
peripheral  populations  whose  leaves  exhibit  these  four  distinctive  leaf -forms 
characterized  in  the  present  paper  fall  very  nearly  upon  the  margin  of  this 
mapping  of  the  Sierra  Madrean  floristic  element  by  Axelrod.  The  only  notable 
exception  is  the  northeastern  arm  in  its  range  lying  to  the  east  of  the  Llano 
Estacado  of  Texas  (cf.  fig.  1).  This  arm  falls,  perhaps  significantly,  to  the 
east  or  coastward  of  the  Fall  Line,  passing  northward  on  the  Coastal  Plain 
sediments.  The  absence  of  Datura  meteloides  in  the  region  to  the  west,  that 
is,  on  the  Llano  Estacado,  is  believed  by  Prof.  E.  L.  Reed  to  be  due  essentially 
to  the  lack  of  proper  alluvial  or  river-bottom  habitats  in  that  region  (per- 
sonal communication,  22  Feb.  1939).'  The  occupation  of  the  Coastal  Plain, 
and  hence,  the  significance  of  the  Fall  Line  as  marking  the  upper  limit  of 
the  Coastal  Plain  or  the  "boundary  between  Triassic  and  older  formations  on 
the  inland  and  Lower  Cretaceous  (Comanchean)  and  later  formations  coast- 
ward,"  has  been  discussed  by  Pennell  very  fully  (1935,  572-579). 

It  will  be  noted  that  the  marginal  phases  of  Datura  meteloides  today  are 
usually  topographically  isolated  by  mountain  ranges  of  varying  height  and 
extent  or  by  arid  desert  expanses  with  their  playas  and  bolsons,  serving  as 
topographic  barriers.  Topographic  isolation,  to  a  degree  then,  has  maintained 
these  distinctive  leaf -forms  at  these  limital  stations. 

Summary 

The  central  populations  of  Datura  meteloides,  the  "tolguacha,"  though  vari- 
able, are  uniform  to  a  degree  and  show  in  general  an  absence  of  such  leaf-forms 
as  are  exhibited  by  the  geographically  peripheral  populations.  There  exist  on 
the  margin  of  its  range  today  four  leaf-forms ;  these  leaf -form  types  are  not 
geographically  segregated,  however,  but  sometimes  occur  together  in  the  same 
marginal  locality. 

An  explanation  may  be :  the  population  of  the  original  center  of  distribu- 
tion was  highly  heterozygous  and  consisted  of  several  infraspecific  races  (this 
needs  cytogenetic  confirmation).  Applying  the  concept  of  Matthew,  these 
several  races  moved  outward  in  irregular  waves  of  varying  intensity  and 
direction  and  for  varying  time  durations.  This  movement  may  have  been  part 
of  the  larger  general  spread  of  the  Sierra  Madrean  floristic  element  over  the 
Southwest.  The  end  result  is  a  series  of  partial-populations  now  scattered  on 

-  Prof.  B.  C.  Tharp  finds  no  coll.  from  the  Llano  Estacado  in  Univ.  Texas  Herb. 


244  CALIFORNIA  ACADEMY  OF  SCIENCES        [Proc.  4th  ser. 

the  periphery  of  the  species  range.  Each  partial-population  is  reminiscent  in 
its  leaf  architecture  of  its  descent  from  a  parent  race  which,  as  a  race,  has 
been  lost  in  the  center  of  the  species  mass  by  the  subsequent  swamping  and 
intermingling  of  races  and  the  genetic  stabilization  of  the  species. 

LITERATURE  CITED 

Anderson,  Edgar  and  Kobert  E.  "Woodson.  Species  of  Tradescantia  indigenous  to  the  United 
States.  Contr.  Arnold  Arboretum  9:1-132.  12  pis.  1935. 

Axelrod,  Daniel  I.  Miocene  flora  from  the  western  border  of  the  Mohave  Desert.  Carn.  Inst. 
Wash.  Publ.  516:1-129.  12  pis.  1939. 

Cain,  Stanley.  Some  observations  on  the  concept  of  species  senescence.  Ecol.  21:213-215. 
1940. 

Ewan,  Joseph.  Taxonomic  history  of  Tolguacha,  Datura  meteloides  DC. :  a  problem  in  the 
application  of  the  Type  Basis  Concept.  Khodora  vol.  46.  1944.  (In  press.) 

Fassett,  Norman.  Mass  collections :  Bubus  odoratus  and  B.  parviflorus.  Ann.  Mo.  Bot.  Gard. 
28:299-374.  1941. 

Gleason,  Henry  .\llan.  Eevision  of  the  North  American  Vernonieae.  Bull.  N.  Y.  Bot.  Gard. 
4:144-243.  1906. 

Manton,  Irene.  Problem  of  Biscutella  laevigata  L.  Annals  Bot.  n.  s.  1:439-462.  1937. 

Matthew,  W.  D.  Climate  and  evolution.  Ann.  N.  Y.  Acad.  Sci.  24:171-318.  1915. 

Pennell,  Francis  W.  Scrophulariaceae  of  Eastern  temperate  North  America.  Monog.  Acad. 
Nat.  Sci.  Phila.  l:i-xiv.  1-650.  1935. 


PROCEEDINGS  J^/*^  ^*^  ^^'^^ 

:^  I L I  -^  R  A  R  y 

OF  THE  '•*r.\        ;?>^f  «*•       ) 

CALIFORNIA  ACADEMY  OF  SCIENCE^^jK^^^^^^^S^ 

Fourth  Series 
Vol.  XXV,  No.  lo,  pp.  245-268,  pi.  25  November  10,  1944 


ALICE  EASTWOOD 
SEMI-CENTENNIAL  PUBLICATIONS 

No.  10 
A  REVISION  OF  SOME  ARIZONA  CACTACEAE 

BY 

LYMAN  BENSON 

Associate  Professor  of  Botany 
Pomona  College 

THE  CACTACEAE,  like  otlier  succulent  groups,  are  frequently  neglected  be- 
cause of  the  common  impression  that  specimens  are  impossible  to  prepare 
or  are  not  worth  much  after  they  are  made.  It  is  hoped  that  recently  published 
articles  at  least  have  helped  to  dispel  this  notion  (Benson,  1939 ;  Peebles, 
1942).  Mr.  Peebles  has  given  a  particularly  thorough  account  of  preparation 
of  cactus  specimens,  and  it  is  illustrated  with  excellent  photographs.  Lack  of 
adequate  series  of  herbarium  specimens  tends  to  place  the  burden  of  study 
of  the  Cactaceae  upon  examination  of  living  individuals  in  the  field.  Despite 
the  many  advantages  of  this  procedure  and  its  absolute  necessity,  it  is  im- 
possible to  make  adequate  comparisons  from  individual  to  individual  without 
intervening  lapse  of  time,  and  it  is  difficult  for  one  person  to  visit  all  of  the 
localities  in  which  a  particular  species  is  to  be  found. 

When  "The  Cacti  of  Arizona"  was  published  (Benson  et  al,  1940),  the 
writer  was  impressed  by  the  close  relationship  of  a  number  of  described 
species,  but  evidence  of  intergradation  was  not  available  in  the  plants  exam- 
ined in  the  field  or  in  the  inadequate  herbarium  material.  Continued  study 
in  the  field  and  preparation  and  acquisition  of  additional  herbarium  speci- 
mens have  resulted  in  the  following  rearrangement  of  species,  particularly  in 
the  genus  Echinocereus. 

Symbols  for  herbaria  in  which  specimens  are  deposited  are  as  follows  :  Sac, 
U.  S.  Field  Station,  Sacaton,  Arizona ;  UA,  University  of  Arizona,  Tucson ; 
B,  Herbarium  of  Lyman  Benson,  Pomona  College,  Claremont,  California; 
3Io,  Missouri  Botanical  Garden,  St.  Louis,  Missouri;  US,  United  States  Na- 

[  245  ] 


246  CALIFOBNIA  ACADEMY  OF  SCIENCES  [Proc.  4th  sek. 

tional  Herbarium,  Smithsonian  Institution.  "Washington,  D.  C. ;  NY,  New 
York  Botanical  Garden,  New  York  City;  8,  Dudley  Herbarium,  Stanford 
University,  California;  3Iich,  University  of  Michigan,  Ann  Arbor.  A  few  of 
these  specimens  have  been  examined  only  in  figures  or  photographs.  The 
herbarium  of  the  writer  is  now  a  part  of  the  Pomona  College  Herbarium. 

The  vegetation  types  referred  to  are  as  follows  (ef.  Benson,  1942;  Shreve, 
1936,  1942;  Shantz  and  Zon,  1924)  : 

Circumboreal  Flora 

Eockj  Mountain  Forests  (spruce-fir  and  yellow  pine) 
Eastern  Forest  Flora 
Sierra  Madrean  Flora 

Southwestern  Oak  Woodland  and  Chaparral 

Southwestern  Coniferous  Woodland  (juniper-pinyon) 

Northern  Desert  (sagebrush) 
Central  Prairie  Flora 

Short  Grass  Prairie  (Great  Plains  grassland) 

Desert  Grassland 
Mexican  Desert  Flora  (creosote  bush) 

Mojavean  Desert 

Sonoran  Desert 

Chihuahuan  Desert 

OPUNTIA 

Opuntia  acanthocarpa  Engelm.  and  Bigel. 
Opuntia  acanthocarpa  Engelm.  and  Bigel.,  1856,  Pac.  R.  E.  Eept.,  4:  51,  pi.  18,  f.  1-3. 

Typically  an  erect,  arborescent  shrub  1-2  m.  high  but  sometimes  lower 
or  even  sprawling;  main  trunk  very  short,  usually  less  than  one-fifth  the 
height  of  the  plant,  the  crown  of  long  branches  not  dense;  branches  cylin- 
drical, the  joints  1.5-3  dm.  long  or  longer,  mostly  about  2-3  cm.  in  diameter, 
the  surface  bearing  areoles  about  18-30  mm.  long  and  about  6  mm.  broad, 
laterally  compressed;  spines  about  10-12  at  an  areole,  the  larger  ones  2.5-3.7 
cm.  long,  rather  dense  on  the  branches  and  more  or  less  obscuring  the  surface 
of  the  stem,  straw-colored,  the  sheaths  conspicuous  and  persistent ;  gloehids 
minute ;  petals  remarkably  variable  in  color,  most  frequently  red  or  yellow, 
the  flower  2.5-3.7  cm.  in  diameter;  fruit  dry  and  more  or  less  shrivelled  at 
maturity,  spiny,  not  giving  rise  to  flowers  at  the  areoles  (not  proliferous)  and 
not  persistent. 

Abundant  on  sandy  flats  and  in  washes  in  the  IMojavean  and  Sonoran 
Deserts  at  2,000  to  4,000  feet  elevation.  California  in  the  southern  and  eastern 
parts  of  the  Mojave  Desert  and  in  the  western  part  of  the  Colorado  Desert; 
southern  Nevada ;  southwestern  Utah ;  Arizona  from  western  IMohave  County 
and  northern  Yuma  County  to  the  Verde  River  and  the  Tucson  Mountains 
and  southward  to  Sonora. 

Type  collection :  "On  the  mountains  of  Cactus  Pass,  about  500  miles  west  of  Santa  Fe." 
Bigelow,  Mo. 


Vol.  XXV]         BENSON:  REVISION  OF  ARIZONA  CACTACEAE  247 

Opuntia  acanthocarpa  var.  Thornberi  (Thornber  and  Bonker)  L.  Benson, 

comb.  nov. 

Opuntia  Thornberi  Thornber  and  Bonker,  1932,  Fantastic  Clan,  133,  148,  pi.  opposite  134, 
upper  fig. 

Bushy  cliolla  occurring  in  patches  1  ni.  or  more  in  diameter ;  joints  of  the 
main  stem  4  dm.  or  more  in  length ;  tubercles  3-5  cm.  long,  about  6  mm.  broad 
and  6  mm.  high ;  spines  7-10,  not  forming  a  dense  mass  on  the  stem,  much  less 
conspicuous  than  the  surface  of  the  stem,  1.5-2.2  cm.  long. 

Gravelly  soil  of  washes,  flats,  and  alluvial  fans  in  the  Mojavean  and  Sonoran 
deserts  at  1,500  to  3,500  feet  elevation.  Peach  Springs,  Mohave  County,  and 
southeastward  along  the  edge  of  the  desert  below  the  Mogollon  Rim  to  north- 
western Graham  County  and  northeastern  Pima  County.  Many  of  the  plants 
occurring  within  a  10  mile  radius  of  Hillside,  Yavapai  County,  are  inter- 
mediate between  this  and  the  typical  variety. 

Type  collection :  "Opuntia  Thornberi  has  been  described  only  very  recently.  .  . ."  There 
is  no  evidence  of  description  elsewhere.  No  type  is  given,  but  fortunately  the  line  drawang 
is  excellent. 

Opuntia  Stanlyi  Engelm. 

Opuntia  Stanlyi  Engelm.  in  Emory,  1848,  Notes  Mil.  Eecon.  p.  157. 

Corynopuntia  Stanlyi  F.  M.  Knuth  in  Backeb.  and  Knuth,  1935,  Kaktus  ABC,  p.  114. 

Low-growing  cholla  consisting  of  a  clump  of  short,  erect  or  ascending 
branches  1-3  dm.  high,  the  clumps  often  3-7  m.  in  diameter;  joints  narrowed 
below,  1-2  dm.  long,  usually  3-5  cm.  in  diameter,  the  surface  bearing  con- 
spicuous, mammillate  tubercules  about  2.5-4  cm.  long,  6-12  mm.  broad,  and 
about  6-8  mm.  high  or  in  an  extreme  form  12-18  mm.  high,  not  tending  to 
join  one  another  or  to  form  ridges  along  the  joint ;  spines  10-18,  those  of  a 
single  areole  1.3-5  cm.  long,  the  larger  ones  commonly  1-1.5  mm.  broad,  but 
sometimes  2-3  mm.  broad,  tan  or  brownish  or  sometimes  red,  strongly  flat- 
tened, sheaths  wanting  or  rudimentary  and  adherent  to  the  tip  of  the  spine, 
papillate-roughened;  glochids  large ;  petals  yellow,  the  flower  about  2.5-3  cm. 
in  diameter ;  fruit  markedly  spiny,  yellow. 

Rocky  or  sandy  mesas  or  arroyo  banks  in  the  Sonoran  Desert  at  2,500  to 
5,000  feet  elevation.  Arizona  on  the  San  Pedro  River  above  Winkleman  and 
from  the  eastern  edge  of  Aravaipa  Valley  to  the  upper  Gila  River  Valley 
and  Clifton,  Duncan,  and  the  San  Simon  Valley  (Graham,  Greenlee,  and  Co- 
chise counties)  ;  New  Mexico  on  the  Gila  River. 

Type  collection:  "October  22d,  1846.  Abundant  on  the  Del  Norte  and  Gila."  Type  not 
preserved.  On  page  63,  the  location  of  the  camp  of  Oct.  22,  1846,  is  given  as  Lat.  32°  38'  13" ; 
Long.  109°  07'  30".  This  is  just  south  of  the  Gila  River  on  the  boundary  between  the  present 
states  of  Arizona  and  New  Mexico. 


248  CALIFORNIA  ACADEMY  OF  SCIENCES  [Pkoc.  4th  ser. 

Opuntia  Stanlyi  var.  Kunzei  (Rose)  L.  Benson,  comb.  nov. 
Opuntia  Kunzei  Rose,  1908,  Smiths.  Misc.  Coll.,  50:  505. 

Joints  narrowed  below,  7.5-15  cm.  long,  2.5-3.5  cm.  in  diameter;  tubercles 
distinct,  about  1.5-2.5  cm.  long,  narrower  and  lower  than  in  the  typical 
variety;  fruit  (as  far  as  known  to  the  writer)  3.3-5  cm.  long,  12-14  mm.  in 
diameter. 

Plains  at  1,000  to  2,000  feet  elevation.  ]\Iaricopa  Mountains  to  Casa  Grande 
and  southwestward  to  the  Gunsight  Mountains  and  southward  to  Baboquivari 
Valley.  Perhaps  occurring  in  Sonora, 

Type  collection :  Pima  County,  Arizona,  Br.  B.  E.  Kunze. 

Opuntia  Stanlyi  var.  Wrightiana  (Baxter)  L.  Benson,  comb.  nov. 

Grusonia  WrigMiana  Baxter,  1935,  Calif.  Cactus,  p.  58. 
Opuntia  Wrightiana  Peebles,  1937,  Desert  Plant  Life,  9:  43. 

Joints  nearly  cylindrical,  1-1.5  or  2  dm.  long,  2.5-3.5  or  4  cm.  in  diameter; 
tubercles  tending  to  be  confluent  into  ribs  on  the  stem,  12-18  mm.  long,  3^ 
mm.  broad,  3-5  mm.  high ;  fruit  3.3-5  cm.  long,  about  12-14  mm.  in  diameter. 

Sand  and  clay  soils  of  valley  floors  in  the  Sonoran  Desert  at  300  to  1,500 
feet  elevation ;  restricted  to  regions  of  1-5  inches  of  annual  rainfall.  South- 
eastern edge  of  California;  Arizona  from  Detrital  Valley,  Mohave  County, 
to'Yuma  County  and  the  western  edge  of  Pima  County ;  southward  as  far  as 
St.  George's  Bay,  Sonora. 

Type  collection:  "Type  Specimen:  Three  joints,  one  with  fruit,  collected  by  Allan  B. 
Clayton,  April  15,  1934,  and  deposited  in  the  Dudley  Herbarium  of  Stanford  University. 
Type  Locality :  Petrified  forest  near  the  Colorado  Eiver,  four  miles  west  of  the  Quartzite- 
Yuma  road,  33  miles  north  of  Yuma,  Arizona." 

Opuntia  erinacea  Engelm.  and  Bigel. 
Opuntia  erinacea  Engelm.  and  Bigel.,  1856,  Pac.  E.  E.  Eept.,  4:  47,  pi.  13.  f.  8-11. 

Low-growing  prickly  pear  forming  clumps  0.3-1  m.  in  diameter,  commonly 
1-1.5  dm.  high,  the  stems  usually  2-4  jointed ;  joints  ovate  to  elliptic-oblong  or 
oblong,  flat,  5-15  or  20  cm.  long,  5-7.5  cm.  broad,  glabrous;  areoles  usually 
about  8-10  mm.  apart,  all  spine-bearing ;  spines  4-7  or  9,  the  longer  ones  about 
5  cm.  long,  more  or  less  stiff,  somewhat  flattened,  elliptic  in  cross  section,  white 
or  pale  gray ;  glochids  1.5-3  mm.  long ;  petals  yellow,  white,  deep  pink,  or  red, 
the  flower  about  5-6  cm.  in  diameter;  fruit  dry  at  maturity,  more  or  less  cylin- 
drical, about  3  cm.  long,  12-14  mm.  in  diameter,  spiny. 

Alluvial  areas  in  the  Mojavean  Desert  at  1,500  to  3,000  feet  elevation.  Cali- 
fornia in  the  southeastern  Mojave  Desert;  southern  Nevada;  southwestern 
Utah ;  northwestern  Arizona  in  Mohave  County ;  Havasupai  Canyon,  Coco- 
nino Count}' ;  near  Carrizo,  Apache  County. 

Type  collection:  "West  of  the  great  Colorado  near  the  Mojave  Creek."  Bigeloio  in 
1853-4,  Mo. 


Vol.  XXV]         BENSON:  BEVISION  OF  AEIZONA  CACTACEAE  249 

Opuntia  erinacea  var.  ursina  (Weber)  Parish. 

Opuntia  nrsina  Weber  in  Boiss.,  1898,  Diet.  Hort.,  2 :  896. 

Opiintia  erinacea  var.  ursina  Parish  in  Jepson,  1936,  Fl.  Calif.,  2 :  542. 

Similar  to  the  typical  variety;  joints  obovate  to  oblong,  1-2  em.  long, 
6-7.5  cm.  broad;  areoles  all  spine-bearing;  spines  usually  6-14,  the  longer 
ones  remarkably  flexible  and  thread-like,  white  or  pale  gray,  not  defiexed. 

Rocky  mountain  slopes  in  the  Mojavean  Desert  at  2,000  to  3,000  feet  eleva- 
tion. California  in  the  southeastern  part  of  the  Mojave  Desert;  Arizona  at 
scattered  stations  in  Mohave  County. 

Type  collection:  Ord  Mountains,  southern  Mojave  Desert  in  San  Bernardino  County, 
California. 

Opuntia  erinacea  var.  hystricina  (Engelm.  and  Bigel.)  L.  Benson 

comb.  nov. 
Opuntia  hystricina  Engelm.  and  Bigel.,  1856,  Pac.  R.  E.  Rept.,  4:  44,  pi.  15.  f.  5-6. 

Joints  obovate,  5-10  or  15  cm.  long,  3.7-7.5  or  9  cm.  broad,  areoles  all  or  all 
but  the  lowest  spine-bearing ;  spines  4  or  usually  5-8,  the  longest  ones  4.5-7.5 
or  10  cm.  long,  tending  to  be  rigid,  defiexed;  petals  red  or  yellow. 

Plains  and  mountains  in  the  Southwestern  Coniferous  Woodland,  the  Short 
Grass  Prairie,  and  the  Rocky  Mountain  Forests  at  4,500  to  7,000  feet  eleva- 
tion. Northern  Arizona  from  Coconino  County  to  Apache  County;  northern 
New  Mexico.  Flowering  in  June. 

Type  collection :  "...  Abundant  from  the  Eio  Grande  westward  to  the  San  Francisco 
mountains, . . ..  The  specimens  before  us  were  obtained  at  the  Colorado  Chiquito  and  on  the 
San  Francisco  Mountains."  Bigelow  in  1853-4,  Mo. 

Opuntia  erinacea  var.  rhodantha  (K.  Schum.)  L.  Benson,  comb.  nov. 

Opuntia  rhodantha  K.  Schum.,  1897,  La  Semaine  Hort.;  1898  Gesamtb.  Kakteen. 

735. 

Joints  obovate,  5-10  cm.  long,  3.7-7.5  cm.  broad,  more  or  less  lead-colored 
(in  the  other  varieties  green)  ;  usually  only  the  distal  areoles  of  each  joint 
spine-bearing,  those  on  at  least  the  lower  half  of  the  joint  usually  spineless; 
spines  1-6  or  8,  the  longer  ones  2.5-3.7  cm.  long,  tending  to  be  rigid,  defiexed ; 
flowers  usually  red  but  sometimes  yellow  {e.g.  McGee  Creek,  Mono  County, 
California,  L.  Benson  11212,  TJA,  B.). 

Flats  and  rocky  hills  in  the  Southwestern  Coniferous  Woodland,  the  North- 
em  Desert,  and  the  Rocky  Mountain  Forests  at  5,000  to  8,000  feet  elevation. 
California  along  the  east  side  of  the  Sierra  Nevada  in  southern  Mono  County 
and  Inyo  County  (Big  Trees  Camp,  Lake  Sabrina  Road  above  Bishop,  L.  Ben- 
son 6009,  B)  and  eastward  to  western  Nebraska;  Arizona  from  Mt.  Dellen- 
baugh,  Mohave  County,  to  Prescott,  the  Mogollon  Rim,  the  northern  end  of 
the  White  Mountains  and  northeastward. 

Type  collection:  "Colorado,  bei  200—2300  m  Hohe:  PURPUS;  bliihte  bei  SPATH  im 
Juni " 


250  CALIFOBXIA  ACADEMY  OF  SCIENCES  [Proc.  4th  ser. 

Opuntia  compressa  (Salisb.)  Macbr. 

Plate  25,  fig.  A,  3 

Cactus  Opuntia  L.  1753,  Sp.  PI.,  p.  468. 

Cactus  compressxis  Salisb.,  1796,  Prodr.,  p.  348. 

Opuntia  compressa  Macbr.,  1922,  Contr.  Gray  Herb.,  (65)  :  41. 

Op^mtia  vulgaris  of  authors,  not  of  Mill.,  1768. 

Low,  prostrate  or  nearly  prostrate  prickly  pear;  roots  fibrous;  joints  light 
green,  broadly  obovate  to  nearly  circular,  4—11  cm.  long,  4—9  cm.  broad ;  leaves 
4r-5  mm.  long,  more  or  less  appressed;  spines  none  or  solitary  in  the  distal 
areoles;  petals  pale  yellow,  the  flower  about  5  cm.  in  diameter,  "with  about 
8  petals"  ;  fruit  2-3  cm.  long. 

Dry  gravelly  or  sandy  soil  in  the  Eastern  Forest  Flora  at  low  elevations. 
Southern  New  England  and  southward  along  the  Atlantic  coastal  plain  to 
South  Carolina.  This  species  is  represented  westward  by  varieties,  and  the 
Arizona  i)lants  of  both  fibrous-rooted  and  tuberous-rooted  types  tentatively 
are  referred  to  them. 

Opuntia  compressa  (Salisb.)  Macbr.  var.  microsperma  (Engelm.) 

L.  Benson,  comb.  nov. 

Cactus  humifusus  Raf.,  1820,  Ann.  Nat.,  p.  15. 

Opuntia  humifusa  Raf.,  1830,  Med.  Fl.  U.  S.,  2 :  247. 

Opuntia  mesacantJia  Raf.,  1830,  Bull.  Bot.  Seringe,  p.  216. 

Opuntia  caespitosa  Raf.,  1830,  loc.  cit. 

Opuntia  Rafinesquei  Engelm.,  1856,  Pac.  R.  R.  Rept.,  4:  41,  pi.  11.  f.  1-3. 

Opuntia  Eafincsquei  var.  microsperma  Engelm.,  1856,  loc.  cit. 

Opuntia  cymochila  Engelm.  and  Bigel.,  1856,  Pac.  R.  R.  Rept.,  4:  43,  pi.  12.  f.  1-3. 

Opuntia  stenochila  Engelm.  and  Bigel.,  1856,  loc.  cit.,  p.  44,  f .  4-6, 

Opuntia  mesacantlia  var.  microsperma  Coult.,  1896,  Contr.  U.  S.  Nat.  Herb.,  3  :  429. 

Opuntia  nesacantha  var.  cymochila  Coult.,  1896,  loc.  cit.,  p.  430. 

Opuntia  mesacantlia  var.  stenocJdla  Coult.,  1896,  loc.  cit.,  p.  430. 

Roots  fibrous ;  joints  dark  green,  averaging  a  little  larger  than  in  the  typical 
variety,  7-12  cm.  long,  5-10  cm.  broad ;  leaves  "6-8"  mm.  long,  not  appressed; 
spines  mostly-  1-3,  one  much  larger  than  the  other  (accessory)  ones;  petals 
"10-12" ;  fruit  3-4  cm.  long.  {Fig.  A  3.) 

According  to  Peebles,  apud  Kearney  and  Peebles,  U.  S.  Dept.  Agric.  Misc. 

Publ.,  (423)  :  611,  1942,  this  plant,  designated  as  0.  Eafincsquei,  occurs  in 

northern  Arizona  from  "Apache  County  to  northern  Mohave  County,  not 

common,  5,000  to  7,000  feet .  .  .."  The  writer  has  seen  no  Arizona  specimens. 

This  wide-ranging  middle  western  variety  of  the  eastern  Opuntia  compressa  is 

exceedingly  variable.  It  includes  a  host  of  minor  forms  variously  considered 

as  species  and  varieties,  and  some  may  be  worthy  of  segregation  as  varieties. 

Type  collections:  (1.)  C.  humifusus,  "From  New  York  to  Kentucky  and  Missouri." 
(2.)  0.  mesacantlia,  Kentucky  to  Louisiana.  (3.)  0.  caespitosa,  Kentucky  and  Tennessee. 
(4.)  0.  Rafinesquei,  nom.  nov.  for  the  first  three  described  species,  and  antedated  l)y  all 
of  them.  No  type  designated  and  only  the  general  range  given.  (5.)  Var.  microsperma. 
No  type  designated  and  the  range  not  distinguished  from  the  broad  range  of  the  typical 


Vol.  XXV]         BENSON:  BETISION  OF  AEIZONA  CACTACEAE  251 

variety  (of  0.  Eafinesquci)  in  tlie  Mississippi  Valley.  (6)  0.  cymochila,  called  a  subspecies, 
but  the  name  published  as  a  binomial,  "On  the  Camanche  plains  east  of  Llaiio  Estaeado, 
near  the  100th  degree  of  longitude;  and  from  there  to  Tucumcari,  80  miles  east  of  the 
Pecos"  in  New  Mexico.  Bigeldw  in  1853-4,  Mo.  (7.)  E.  stenochila,  called  a  subspecies  but 
published  as  a  binomial,  "At  the  Canyon  of  the  Zuiii,"  Bigelow  in  1853-4,  Mo. 

Opuntia  compressa  var.  macrorhiza  (Engelm.)  L.  Benson,  comb,  no  v. 

Plate  25,  figs.  A,  1,  2 
Opuntia  macrorhiza  Engelm.,  1850,  Bost.  Jour.  Nat.  Hist.,  6 :  206. 
Opuntia  fu  si  for  mis  Engelm.  and  Bigel.,  1856,  Pac.  R.  E.  Eept.,  4:  43. 
Opuntia  mesacantha  Eaf .  var.  macrorhiza  Coult.,  1896,  Contr.  U.  S.  Nat.  Herb.,  3  :  430. 
Opuntia  plumbea  EosE,  Smiths.  Misc.  Coll.,  1908,  50 :  524. 
Opuntia  delicata  Eose,  1911,  Contr.  U.  S.  Nat.  Herb.,  13 :  310. 
Opuntia  Loomisii  Peebles,  1938,  Cact.  and  Succ.  Jour.,  9:  109. 

Similar  to  var.  microsperma;  roots  tuberous  even  in  young  plants;  spines 
commonly  2  or  more  per  areole. 

Plains  and  hills  in  the  Desert  Grassland,  the  Southwestern  Coniferous 
Woodland,  the  Southwestern  Oak  Woodland,  and  the  Rocky  Mountain  For- 
ests at  4,500  to  6,000  feet  elevation  and  (eastward)  in  the  Short  Grass  Prairie 
and  in  the  Eastern  Forest  Flora.  Arizona  from  Mohave  County  to  Apache 
County,  eastern  Pima  Countj^,  and  Santa  Cruz  County ;  eastward  to  Missouri 
and  Texas.  The  Arizona  form  is  variable.  Usually  it  is  longer-spined  and  often 
the  joints  are  larger.  Some  forms  approach  the  fibrous-rooted  species,  Opuntia 
phaeacantha. 

Type  collections:  (1.)  0.  macrorhisa,  "Naked,  sterile,  rocky  places  on  the  Upper  Guada- 
loupe."  Texas,  Lindheimer,  Mo.  (2.)  0.  fusiformis,  called  a  subspecies,  but  published  as  a 

binomial,  "Cross-timbers  longitude  97°-99° "  Bigelow  in  1853-4,  Mo.  (3.)  0.  plumbea, 

San  Carlos  Indian  Eeservation,  Arizona,  Coville  in  1894,  US.  This  form  is  apparently 
identical  with  that  on  the  Great  Plains.  A  specimen  (Pearce,  Cochise  County,  Nichol 
in  1940  [pi.  25  fig.  A,  1])  is  growing  in  the  Pomona  College  botany  greenhouse.  (4).  0. 
delicata,  "Type  U.  S.  National  Herbarium  no.  454622,  collected  by  J.  N.  Eose  at  Calabasa, 
April  30,  1908  (no.  11951)."  Arizona.  (5.)  0.  Loomisii,  "Type,  E.  F.  Loomis  No.  SF.  298, 
Prescott,  Arizona,  Aug.  10,  1930,  represented  by  a  plant  grown  under  cultivation  at  Sacaton, 
Arizona,  and  material  removed  from  the  cultivated  plant  and  deposited  in  the  U.  S.  Na- 
tional Herbarium,  Washington,  D.  C."-  This  is  the  extreme  opposite  the  type  described  by 
Eose  as  0.  plumbea,  and  it  approaches  0.  phaeacantlia. 

Opuntia  tortispina  Engelm.  and  Bigel. 

The  plant  described  previously  as  Opuntia  tortispina  is  merely  a  form  of 
Opuntia  phaeacantha  with  long,  twisted,  light-colored  spines.  The  description 
was  drawn  from  plants  collected  four  miles  south  of  Navajo  Bridge,  Coconino 
County,  by  A.  A.  Nichol  in  1940. 

ECHINOCEREUS 

Mostly  small  cacti  forming  low  clumps  1-4  dm.  or  rarely  about  1  m.  in 
diameter  and  rarely  more  than  3  (maximum  6)  dm.  high  or  the  stems  not 
infrequently  solitary,  the  length  of  a  single  branch  not  more  than  8  (or  rarely 


252  CALIFOBNIA  ACADEMY  OF  SCIEXCES  [Proc. -Ith  ser. 

in  a  decumbent  type  12)  times  the  diameter ;  stem  usually  branching  only  at 
or  near  the  base  except  after  injury,  or  sometimes  unbranched,  with  continu- 
ous ribs,  cylindrical  or  ovoid,  leafless  even  on  the  new  season's  growth;  areoles 
not  producing  glochids ;  flower  appearing  below  the  apex  of  the  branch ;  the 
flower  bud  bursting  through  the  epidermis  above  the  lower  edge  of  a  spine- 
producing  areole;  floral  tube  (hypanthium?)  long  or  at  least  well-developed 
above  the  ovary;  fruit  spiny,  the  clusters  of  spines  readily  removed  at  ma- 
turity. 

The  plants  of  this  genus  occurring  in  Arizona  are  members  of  complex 
groups.  Previously  (Benson  et  al.,  1940)  ten  described  species  were  recog- 
nized as  closely  related  entities,  although  it  was  clear  that  the  group  was  in 
need  of  reorganization  and  that  several  species  might  better  be  reduced  to 
varieties.  Further  study  in  the  fleld  and  collection  of  additional  herbarium 
and  garden  material  emphasizes  the  need  for  the  following  revision.  The  dif- 
ferentiation of  this  genus  from  Cereus  has  been  discussed  in  another  paper 
(Benson,  1941). 

Key  to  the  Species 

1.  Petals  scarlet  or  crimson   (with  plastid  pigmentation?);  areoles  bearing  white  felt  at 
maturity;  flowers  not  closing  at  night,  remaining  open  until  the  end  of  anthesis. 

1.  Ecldnocereus  trifflocliidiatits. 

1.  Petals  purple  to  lavender  (probably  pigmented  with  anthocyans)  ;  areoles  not  bearing 

white  felt  at  maturity,  the  felt  of  the  young  areoles  rarely  persistent  for  two  years; 

flowers  closing  at  night  and  reopening  in  the  morning  or  in  hot  weather  withering  at  the 

end  of  a  single  day. 

2.  Areoles  vertically  elongate;  stems  unbranched 2.  Echinocereus  pectinatus. 

2.  Areoles  circular;  stems  usually  branched  except  in  young  plants. 

3.  Central  spines  2—6,  approximately  the  same  size  or  at  least  all  well  developed,  usually 
(at  least  the  lower  deflexed  ones)  flattened  at  least  at  the  bases. 

3.  Echinocereus  Engelmannii. 
3.  Central  spine  1,  often  with  1-3  shorter  accessory  or  rudimentary  centrals  in  some  of 
the  areoles  of  the  plant,  circular  in  cross  section,  not  flattened. 

4.  Principal  central  spine  not  both  strongly  curved  and  pointing  do^^^lward ;  spines 
white,  gray,  red,  black,  bro^\^l,  or  bicolored;  flowers  5-8.5  cm.  in  diameter. 

4.  Echinocereus  Fendleri. 

4.  Principal  central  spine  strongly  curved  near  the  base,  turned  do^^^lward ;  spines 

yellow  or  straw-color,  uniformly  colored;  flowers   (in  the  specimens  available) 

3.5-5  cm.  in  diameter. 5.  Echinocereus  Ledingii 

1.  Echinocereus  triglochidiatus  Engelm. 

Echinocereus  triglochidiatus  Engelm.,  in  Wisliz.,  1848,  Mem.  Tour.  N.  Mex.,  p.  93. 
Cereus  triglochidiatus  Engelm.,  apud  A.  Gray,  1849,  Mem.  Amer.  Acad.,  II.  4:  50. 
Echinocereus  paucispinus  (Engelm.)  Engelm.,  ex.  Eiimplcr  var.  triglochidiatus  K.  Schum., 
1898,  Gesamtb.  Kakteen,  281. 

Older  plants  dense,  forming  clumps  of  few  to  many  stems  each  2^  or  6  dm. 
high,  mostly  4—8  cm.  in  diameter,  obviously  and  conspicuously  green,  not 
obscured  by  the  spines,  ribs  commonly  5-8,  somewhat  tuberculate  at  the 
areoles ;  spines  exceedingly  variable,  sometimes  as  few  as  3  or  as  many  as  16, 


Vol.  XXV]         BENSON:  REVISION  OF  ARIZONA  CACTACEAE  253 

1-3  em.  long;  central  spines  0-5  or  6,  smooth  and  ribbed,  somewhat  longer 
than  the  radials ;  radial  spines  3-10;  areole  circular,  bearing  white  felt ;  petals 
crimson  or  scarlet,  the  flower  about  3  cm.  in  diameter ;  floral  tube  2-3  cm.  long 
above  the  ovary,  the  scales  with  tufts  of  white  hair ;  fruit  more  or  less  spiny, 
subcylindrical,  about  2-3  cm.  long. 

Southern  Colorado  to  trans-Pecos  Texas.  A  few  Arizona  plants  approach 
the  typical  variety. 

Type  collection:  "On  "Wolf  Creek  [near  Santa  Pe,  New  Mexico],  in  pine  woods,  flowers 
in  June  [Wislizenius  in  1846,  Mo.] ;  Santa  Fe  (Fendler) "  Mo. 

1a.  Echinocereus  triglochidiatus  var.  gonacanthus  (Engelm.  and  Bigel.) 

Engelm.  and  Bigel.  ex  W.  T.  Marshall 

Cereus  gonacanthus  Engelm.  and  Bigel.,  1856,  Pae.  E.  E.  Eept.,  4:  33,  pi.  5.  f.  2-3. 
Echinocereus  gonacanthus  Engelm.,  ex  Eiimpler  in  Forster,  1885,  Handb.  Cacteenk.,  ed. 

2,  p.  806. 
Echinocereus  paucispinus  (Engelm.)  Engelm.,  ex  Eiimpler  var.  gonacanthus  K.  Schum., 

1898,  Gesamtb.  Kakteen,  p.  281. 
Echinocereus  triglochidiatus  var.  gonacanthus  (Engelm.  and  Bigel.)  Engelm.  and  Bigel., 

ex  W.  T.  Marshall  in  Marshall  and  Bock,  1941,  Cactaceae,  p.  117. 

A  robust  type  similar  to  the  typical  variety;  spines  mostly  6-8  per  areole, 
twisted  and  somewhat  curved ;  central  spine  usually  solitary  but  sometimes 
more  than  one,  3-5  cm.  long,  markedly  angled  wdth  usually  6  or  7  sharp  ribs ; 
radial  spines  2-3  cm.  long. 

High  plains  at  the  lower  edge  of  the  Rocky  Mountain  Forests  at  6,000  to 
8,000  feet  elevation.  Arizona  40  miles  west  of  Zuiii,  New  Mexico,  and  near  St. 
Johns,  Apache  County,  (Wyatt  W.  Jones,  the  specimen  with  3-angled  central 
spines.)  New  Mexico  near  Gallup  and  Zuni.  Reported  from  the  White  Sands. 

Type  collection:  "On  high  sand-bluffs,  covered  with  scattering  cedars,  near  the  natural 
well,  about  40  miles  west  of  Zuni,  near  the  109th  degree.  Only  seen  in  that  locality."  Prob- 
ably Apache  County,  Arizona,  Bigelow  in  1853-4,  Mo.  Zuni,  New  Mexico,  is  within  10  or 
12  miles  of  the  Arizona  line,  but  the  109th  Meridian  is  just  east  of  the  state  boundary. 

1b.  Echinocereus  triglochidiatus  var.  polyacanthus  (Engelm.)  L.  Benson, 

comb.  nov. 

Echinocereus  polyacanthus  Engelm.  in  Wisliz.,  1848,  Mem.  Tour.  N.  Mex.,  p.  104. 
Cereus  polyacanthus  Engelm.  apud  A.  Gray,  1849,  Mem.  Amer.  Acad.,  II,  4:  50. 
Echinocereus  arisonicus  Orctjtt,  1926,  Cactography,  (3) :  3. 

Clumps  of  stems  not  dense  or  dome-like,  the  stems  usually  15^5,  1.5-5  dm. 
high,  6-10  cm.  in  diameter,  elongated  and  cylindrical ;  ribs  about  9-10,  promi- 
nent, not  markedly  tuberculate  at  the  areoles;  central  spines  2-4,  turned  down- 
ward or  perpendicular  to  the  stem,  straight,  stout  or  slender,  2.5-5  cm.  long ; 
radial  spines  about  10,  appressed ;  flower  about  3.5-4  cm.  in  diameter ;  floral 
tube  about  3  cm.  long  above  the  ovary. 

Desert  Grassland  and  Southwestern  Oak  Woodland  at  3,500  to  5,500  feet 
elevation.  Arizona  in  the  Pinal  Mountains,  Pinal  County,  and  the  Galliuro 


254  CALIFORNIA  ACADEMY  OF  SCIENCES  [Proc.  iiH  ser. 

Mountains,  Graham  County,  and  southward  to  Santa  Cruz  and  Cochise  coun- 
ties. Southeastern  Arizona  to  western  New  Mexico,  Chihuahua,  and  Durango. 
Floweriiip-  late  in  ^lay  and  in  June. 

Type   collections:    (1.)    E.   poly  acanthus,   "Cosiluiiriachi,"   Chihuahua.    Wifilizenius  in 
1846-7,  Mo.  (2.)  E.  arizonicus,  Pinal  Mountains,  Pinal  County,  Arizona. 

Ic.  Echinocereus  triglochidiatus  mw.  melanacanthus  (Engelm.)  L.  Benson, 

comb.  nov. 

Echinocereus  coccineus  Engelm.  in  Wisliz.,  1848,  Mem.  Tour.  N.  Max.,  p.  93. 

Cereus  coccineiis  Engelm.  apud  A.  Gray,  1849,  Mem.  Amer.  Acad.,  II,  4:  51.  1849,  not 

Salm-Dyck  ex  DC.  in  1828. 
Cereus  coccineus  yar.  melanacanthus  Engelm.,  loc.  cit. 
Cereus  coccineus  var.  cylindricus  Engelm.,  loc.  cit. 

Cereus  majavensis  var.  suniensis  Engelm.  and  Bigel.,  1856,  Pac.  R.  R.  Rept.,  4 :  33,  pi.  4,  f .  9. 
Cereus  Bigelovii  Engelm.  var.  zuniensis  Engelm.,  apud  Engelm.  and  Bigel.,  1856,  loc.  cit. 

(on  the  plate). 
Cereus  phoeniceus  Engelm.  apud  Engelm.  and  Bigel.,  1856,  loc.  cit.,  p.  34,  f .  1-3. 
Echinocereus  phoeniceus  Engelm.,  ex  Riimpler  in  Forster,  1885,  Handb.  Cacteenk.,  ed.  2, 

p.  788. 
Echinocereus  mojavensis  var.  zuniensis  Engelm.,  ex  Riimpler,  Forster,  1885,  loc.  cit.,  p.  803. 
Echinocereus  phoeniceus  var.  inermis  K.  ScHUM.,  1896,  ]Monatsschr.  Kakteenk.,  6:  150. 
Echinocereus  coccineus  var.  inermis  3.  A.  Purpus,  1925,  Mitt.  Deutseh.  Dendr.  Ges.,  1925  :  49. 
Echinocereus  triglochidiatus  var.  coccineus  Engelm.,  ex  W.  T.  Marshall  in  ]SlARsnALL  and 

Bock,  1941,  Cactaceae,  p.  117. 
Echinocereus  canyonensis  Clover  and  Jotter,  1941,  Bull.  Torrey  Club,  (58  :  417.  f .  1  (4). 

Stems  from  a  few  to  more  than  500,  cylindrical  or  ovoid,  0.7-1.5  dm.  high, 
mostly  3-5  or  6  cm.  in  diameter ;  ribs  commonly  8  or  10,  not  markedly  tuber- 
culate  at  the  areoles ;  central  spine  1  or  sometimes  more  than  1,  perpendicular 
to  the  joint,  straight,  tapering,  1.3-3  cm.  long;  radial  spines  commonly  5-10, 
similar  to  the  central  spine  but  smaller;  flow^er  3-4.5  cm.  in  diameter;  floral 
tube  2.5-3  cm.  long  above  the  ovary. 

Rocky  hillsides  and  the  forest  floor  in  the  Southwestern  Coniferous  Wood- 
land, the  Southwestern  Oak  Woodland  and  Chaparral,  and  the  Rocky  ]\Ioun- 
tain  Forests  at  4,000  to  8,000  feet  elevation.  Arizona  in  the  mountainous 
regions  from  the  Ilualpai  Mountains  to  the  Kaibab  Forest,  and  southeastward 
to  the  Santa  Catalina  Mountains,  Pima  County,  and  to  Greenlee  County, 
thence  southward  to  Santa  Cruz  and  Cochise  counties.  Southern  Utah  to 
Colorado,  Arizona,  and  New  Mexico.  Flowering  from  March  until  June,  de- 
pending upon  altitude. 

Type  collections:  (1.)  E.  coccineus,  "With  the  foregoing,  also  about  Santa  Fe."  The 
"foregoing"  was  E.  triglochidiatus,  see  above.  Mo.  (2.)  Var.  melanacanthus,  "Among  a 
number  of  plants  of  this  family  Avhich  Mr.  Fendler  sent  from  Santa  Fe  in  a  living  state, 
but  which  unfortunately  were  all  dead  when  they  came  to  hand,  are  some  specimens  which 

appear  to  be  varieties  of  this  species "  (3.)  Var.  cylindricus,  cf .  statement  for  var. 

melanacanthus.  Mo.  (4.)  Var.  suniensis,  "It  was  found  near  Canon  Diablo,  on  the  Colorado 
Chiquitn,  about  120  miles  west  of  Zufii."  Canyon  Diablo,  near  the  Little  Colorado  River, 
Arizona,  Bigclow  in  1853-4,  Mo.  (5.)  C.  phoeniceus,  nom.  nov.  for  E.  coccinms.  (6.)  Var, 


Vol.  XXV]         BEN  SON  :  BEVISIOX  OF  ABIZON  A  CACTA  CEAE  255 

hterniis,". . .  Heir  C.  A.  Purpus  aus  Colorado  an  die  liiesige  beriihinte  Gaitueiei  von  SPATH 
schickte  . .  .."  (7.)  E.  conyonensis,  "TYPE  {Clover  and  Jotter  2317)  collected  in  a  sandy 
pocket  on  a  steep  limestone  outcrop  100  yards  from  the  [Colorado]  river,  Bass  Cable  below 
Hermit  Creek  Eapids,  Grand  Canyon,  Coconino  Co.,  Arizona.  Locally  abundant.  Living 
specimen  in  the  Botanical  Gardens,  (Gard.  16846),  University  of  Michigan,  Ann  Arbor, 
Michigan."  Specimen  now  pressed. 

Id.  Echinocereus  triglochidiatus  var.  mojavensis  (Eiigelm.)  L.  Benson, 

comb.  nov. 

Cereus  mojavensis  Engelm.  and  Bigel.,  1856,  Pac.  E.  E.  Eept.,  4 :  33.  pi.  4. 
Cereus  Bigelovii  Engelm.,  apud  Engelm.  and  Bigel.,  1856,  loc.  cit.  (on  the  plate.) 
Echinocereus  mojavensis  Engelm.,  ex.  Eiimpler  in  Forster,  1885,  Handb.  Cacteenk.,  ed.  2, 
p.  803. 

Similar  to  var.  melanacanthus ;  ribs  (of  the  material  available)  strongly  tu- 
bereiilate  at  the  areoles;  spines  curving,  tortuous,  somewhat  flexible,  Avhitish; 
floral  tube  (of  the  plants  available)  1.7-2  cm.  long. 

Rocky  hillsides  in  the  upper  Mojavean  Desert  and  the  Southwestern  Coni- 
ferous Woodland  at  3,500  to  5,000  feet  elevation.  Arizona  in  scattered  locali- 
ties in  Mohave  County  (Black  Mountains;  Grand  Wash  Cliffs;  Hualpai 
Indian  Eeservation).  Southeastern  California  to  southern  Utah  and  north- 
western Arizona. 

Type  collections:  (1.)  C.  mojavensis,  "Found  between  the  Eio  Colorado  and  Mojave 
Creek."  California  east  of  the  Mojave  River,  Bigelow  in  1853-4,  Mo.  (2.)  C.  Bigelovii, 
apparently  a  discarded  name  once  intended  to  be  proposed  for  the  same  plant. 

2.  Echinocereus  pectinatus  (Scheidw.)  Engelm. 

Echinocactus  pectinatus  Scheidw.,  1838,  Bull.  Acad.  Sci.,  Brux.,  5  :  492. 

Eehinocactus  pectiniferus  Lemaire,  1839,  Cact.  Gen.  Nov.  sp.,  p.  25. 

Ecliinopsis  pectinata  Fennel,  1843,  Allg.  Gartenz.,  11 :  282. 

Echinocereus  pectinatus  Engelm.,  in  WiSLiz.,  1848,  Mem.  Tour.  N.  Mex.,  p.  110. 

Cereus  pectinatus  Engelm.,  apud  A.  Gray,  1849,  Mem.  Amer.  Acad.  Sci.,  II,  4:  50. 

Cereus  pectiniferus  Labouret,  1853,  Monogr.  Cact.,  p.  320. 

Stem  unbranehed,  erect,  cylindrical,  1-2  dm.  high,  3-8  cm.  in  diameter ;  ribs 
about  20,  not  markedly  tuberculate  at  the  areoles,  but  transversely  grooved 
between  adjacent  areoles;  spines  white  to  red,  not  completely  obliterating  the 
stem ;  central  spines  2-5,  mostly  3,  mostly  1-3  mm.  long,  in  a  single  row ; 
radial  spines  15-20,  appressed  against  the  stem,  pectinate,  markedly  ap- 
pressed,  3-8  mm.  long,  flattened,  straight,  tapering;  areoles  vertically  elon- 
gate, the  growth  of  different  years  marked  off  by  the  color  of  the  spines 
produced  in  early  or  late  season;  flower  funnel-form;  petals  reddish-purple, 
tending  toward  lavender ;  floral  tube  about  12  mm.  long  above  the  o.vary ;  fruit 
nearly  spherical,  about  2.5  cm.  in  diameter. 

Rocky  or  gravelly  hills  in  the  Desert  Grassland  and  perhaps  the  Southwest- 
ern Oak  Woodland  at  4,000  to  5,000  feet  elevation.  Southeastern  Arizona  to 
Chihuahua.  The  plant  occurring  in  Cochise  Count}^,  Arizona,  seems  to  be 
referable  to  the  typical  variety,  but  it  has  not  been  seen  in  flower  (Perilla  Mts., 


256  CALIFORNIA  ACADEMY  OF  SCIEXCES  [Proc.  4th  ser. 

Harlan  in  1939,  Sac;  near  Douglas,  W.  T7.  Jones  TJA,  B;  16  miles  northeast 

of  Donglas,  Benson  and  Darrow  11113,  UA,  B).  This  is  one  of  several  Chi- 

huahiian  Desert  species  occurring  on  limestone  hills  in  southeastern  Arizona. 

Type  collection:  The  identity  of  the  type  is  difficult  to  establish.  According  to  Britton 
and  Rose,  Cactaceae,  3:  29-30,  1922,  "This  species  was  first  collected  by  Galeotti  who  sent 
a  collection  to  Belgium  from  the  states  of  San  Luis  Potosi  and  Guanajuato,  Mexico.  The 
type  station,  Villa  del  Pennasco,  we  have  not  located.  It  was  soon  after  figured  by  Lemaire 
(Icon.  Cact.,  pi.  14  or  15)  and  Pfeiffer  (Abbild.  Beschr.  Cact.,  2:  pi.  10),  very  likely  from 
the  type  collection."  According  to  Engelmann,  loc.  cit.,  however,  the  type  is  from  Chihuahua: 
"The  description  of  the  plant  (which  died  ^\^thout  producing  flowers)  found  in  several 
works,  as  well  as  in  the  latest  publication  on  Cactaceae,  before  me,  of  Foerster,  Leipzig, 
1846,  was  made,  as  Prince  Salm  informed -me,  from  specimens  sent  from  Chihuahua  by 
Mr.  Potts.  It  entirely  agrees  with  my  specimen  from  the  same  region." 

2a.  Echinocereus  pectinatus  var.  rigidissimus  (Engelm.) 
Engelm.  ex  Kiimpler 

Cerevs  pectinatus  (Scheidw.)  Engelm.,  var.  rigidissimus  Engelm.,  1857,  Proc.  Amer.  Acad., 

3:  279. 
EcMnocere\is  rigidissimus  Rose,  1909,  Contr.  U.  S.  Nat.  Herb.,  12 :  293. 
Echinocereiis  dasyaeanthus  Engelm.,  var.  rigidissimus  W.  T.  Marshall  in  Marshall  and 

Bock,  1941,  Cactaceae,  p.  119. 

Stems  0.7-3.5  dm.  high,  5-10  cm.  in  diameter ;  areoles  crowded,  the  spines 
completelj'  obliterating  the  surface  of  the  stem;  central  spines  none;  radial 
spines  flat,  thicker  and  stronger  than  in  the  typical  variety,  15-18,  strongly 
appressed,  6-14  mm.  long ;  flowers  between  lavender  and  magenta. 

Occurring  on  rocky  sloj^es  and  grassy  areas  in  the  Desert  Grassland  and  the 
Southwestern  Oak  Woodland  at  4,000  to  6,000  feet  elevation.  Arizona  from 
the  Baboquivari  Mountains  to  the  Santa  Catalina  Mountains,  Pima  County, 
and  southeastward  to  Santa  Cruz  and  Cochise  counties ;  northern  Sonora. 

Type  collection :  "...  from  Sonora  . . .."  According  to  Engelmann,  in  Emory,  Rept.  IT.  S. 
and  Mex.  Bound.  Surv.,  2 :  32,  1859.  "In  the  Sierras  of  Pimeria  Alta  in  Sonora,  and  farther 
west  (A.  Schott).  Flowers  in  June  and  July."  Mo. 

2b.  Echinocereus  pectinatus  var.  neomexicanus  (Coulter)  L.  Benson, 

comb.  nov. 

Echinocereus  dasyaeanthus  Engelm.,  in  Wisliz.,  1848,  Mem.  Tour.  N.  Mex.,  p.  100. 
Cereus  dasyaeanthus  Engelm.,  apud  A.  Gray,  1849,  Mem.  Amer.  Acad.,  II,  4:  50. 
Cereus  ctenoides  Engelm.,  in  Emory,  1839,  Rept.  U.  S.  and  Mex.  Bound.  Surv.,  2 :  31,  pi.  42. 
Cereus  ctenoides  Engelm.,  et  Riimpler  in  Forster,  1885,  Hand.  Cacteenk.,  ed.  2,  p.  819,  f .  109. 
Cereus  dasyaeanthus  var.  neomexicanus  Coult.,  1896,  Contr.  U.  S.  Nat.  Herb.,  3:   384. 
Not  Echinocereus  neomexicanus  Standl.,  1908,  Bull.  Torrey  Club,  35:  87. 

Stems  1-3  dm.  high,  5-7  or  10  cm.  in  diameter;  areoles  crowded,  the  spines 
usualh'  practically  obliterating  the  surface  of  the  stem ;  central  spines  mostly 
5-8,  usually  arranged  irregularly  (or  in  a  single  row  in  types  transitional  to 
the  typical  variety),  3-10  mm.  long;  radial  spines  somcAvhat  flattened,  15-20 
or  more,  not  markedly  appressed,  10-15  mm.  long;  flowers  typically  yellow 
but  sometimes  varying  to  the  magenta  series. 


Vol.  XXV]         BENSON:  REVISION  OF  ARIZONA  CACTACEAE  257 

Desert  Grassland  and  Oak  Woodland  at  4,000  to  5,000  feet  elevation.  Ari- 
zona (according  to  Professor  Thornber)  in  the  southern  part  of  the  Baboqni- 
vari  Mountains,  Pima  County,  and  (according  to  A.  A.  Nichol)  in  the  Perilla 
and  Guadaloupe  mountains,  Cochise  County,  as  well  as  (according  to  A.  A. 
Nichol)  in  jjerhaps  modified  form  in  the  Oro  Blanco  Mountains  west  of 
Nogales,  Santa  Cruz  County,  and  at  Altar  and  Pitiquito  in  adjacent  Sonora; 
Southern  New  Mexico ;  Trans-Pecos  Texas ;  Chihuahua. 

Type  collections:  (1.)  E.  dasy acanthus,  "On  the  mountains  near  El  Paso;  in  August  in 
flower  and  fruit."  Wislizenius  in  1846-7,  Mo.  (2.)  C.  ctenoides,  "From  Eagle  Pass  to  Santa 
Eosa  (Bigelow)  ;  on  the  Pecos  (Wright.)"  Texas.  Mo.  (3.)  Var.  neomexicanus,  "Type, 
Wright  366  in  Herb.  Mo.  Bot.  Gard.  Southeastern  New  Mexico."  The  necessity  for  use  of 
this  name  in  varietal  status  is  unfortunate,  for  the  little-known  epithet  neomexicanus  dis- 
places the  well-known  one,  dasy  acanthus,  and  the  epithet  neomexicanus  was  proposed  later 
in  specific  rank  for  a  plant  of  the  Echinocereus  triglochidiatus  complex. 

3.  Echinocereus  Engelmannii  (Parry)  Parry  ex  Rlimpler 

Cereus  Engelmannii  Parry  ex  Engelm.,  1852,  Amer.  Jour.  Sci.,  II,  14:  338. 

Cereus  Engelmannii  var.  variegatus  Engelm.,  1856,  Pac.  E.  E.  Eept.,  4 :  35,  pi.  5.  f .  4-7. 

Cereus  Engelmannii  var.  chrysocentrus  Engelm.,  1856,  loc.  cit.,  f .  8-10. 

Echinocereus  Engelmannii  (Parry)  Parry  ex  Eihnpler  in  Forster,  1885,  Handb.  Cacteenk., 

ed.  2,  p.  805. 
Echinocereus  Engelmannii  var.  variegatus  Engelm.,  ex  Eiimpler,  Forster,  1885,  loc.  cit., 

p.  806. 
Echinocereus  Engelmannii  var.  chrysocentrtis  Engelm.,  ex  Eiimpler,  Forster,  1885,  loc.  cit., 

p.  806. 
Echinocereus  chrysocentrus  Orcutt,  1926,  Cactography,  (3)  :  4. 

Clumps  not  dense,  stems  erect,  usually  1-25,  not  crowded,  cylindrical, 
usually  1.5-3  dm.  high,  5-7.5  cm.  in  diameter,  firm,  ribs  usually  10-13,  not 
markedly  tuberculate  at  the  areoles ;  spines  sometimes  obliterating  the  surface 
of  the  stem;  central  spines  2-6,  all  large  and  well-developed,  red,  yellow,  white, 
brown,  or  gray,  or  the  lower  one  white  and  the  others  variously  colored,  some 
of  them  deflexed,  tapering,  3.7-9  cm.  long,  stout  but  somewhat  flexible,  curv- 
ing or  twisted,  usually  flattened  at  the  bases ;  radial  spines  6-12,  similar  to 
the  central  but  smaller ;  areole  circular  or  nearly  so,  not  bearing  white  felt 
after  the  first  year ;  petals  purple  to  magenta,  the  flower  5-7.5  cm.  in  diameter, 
funnel-form;  floral  tube  about  12  mm.  long  above  the  ovary ;  fruit  red,  nearly 
spherical  to  ovoid,  about  3  cm.  long,  spiny,  shedding  the  interlocking  spines  at 
maturity,  edible. 

Hills,  plains,  alluvial  bottoms,  and  benches  in  the  Northern,  Mojavean,  and 
Sonoran  deserts  at  100  to  2,500  or  rarely  5,000  feet  elevation.  California  about 
the  Owens  River  Valley  and  in  the  Mojave  and  Colorado  deserts ;  southern 
Nevada ;  southern  Utah ;  Arizona  from  southern  Mohave  County  to  Yuma 
County,  Pinal  County,  and  western  Pima  County;  Baja  California  to  Sonora. 
Flowering  in  late  March  and  in  April. 

Type  collections:  (1.)  C.  Engelmannii,  "Mountains  about  San  Felipe,  on  the  eastern 
declivity  of  the  Cordilleras."  Colorado  Desert  in  San  Diego  County,  California,  Parry  in 


258  CALIFORNIA  ACADEMY  OF  SCIEXCES  [Pkoc.  4th  skk. 

1852,  Mo.  (2.)  Var.  varicgatus,  "On  the  Cactus  mountains  and  at  the  head  of  Williams 
Eiver;  degrees  113i/^  longitude."  Arizona,  Bigelow  in  1853-4  Mo.  (3.)  Var.  chrysocentrus, 
"It  was  found  where  C.  variegatus  [var.  varie gains']  disappears  on  the  lower  part  of 
Williams's  River,  and  was  seen  from  there  to  Mojave  Creek,  and  up  that  stream  to  the  Sierra 
Nevada."  Western  Arizona  and  California,  Bigelow  in  1853-4,  Mo.  The  species  is  l)arely 
to  be  maintained  as  distinct  from  E.  Fendleri,  and  the  separation  is  rather  arbitrary. 

3a.  Echinocereus  Engelmannii  v;n-.  Nicholii  L.  Benson,  var.  no  v. 

Plate  25,  fig.  B. 

Caulibus  10-30  erectis,  3-5  dm.  longis,  5-7  vel  8  cm.  diametro ;  aculeis  flavis ; 
petalis  colore  similibus  lavendulae. 

Stems  erect,  in  large  clumps  of  10-30,  3-5  dm.  high,  5-7  or  8  cm.  in  diame- 
ter ;  spines  all  yellow  ;  petals  lavender. 

Common  on  rocky  hillsides  in  the  Sonoran  Desert  at  1,000  to  2,000  feet 
elevation ;  not  occurring  on  the  alluvial  flats  between  the  mountains.  Arizona 
in  western  Pima  County  from  the  Organ  Pipe  Cactus  National  Monument  to 
the  Silver  Bell  Mountains ;  largely  on  the  Papago  Indian  Reservation. 

Type  collection:  Silver  Bell  Mine,  Silver  Bell  Mountains,  Pima  County,  Arizona,  L. 
Benson  10720,  March  28,  1941,  TYPE  VA  ;  ISOTYPE  B.  Mr.  A.  A.  Nichol  has  brought 
many  plants  of  this  variety  to  the  University  of  Arizona  on  the  suppostion  that  they  Avere 
a  new  entity. 

3i?.  Echinocereus  Eng-elmannii  var.  decumbens  (Clover  and  Jotter), 

L.  Benson,  comb.  nov. 
Echinocereus  decumbens  Clover  and  Jotter,  1941,  Bull.  Torrey  Club,  68:  417,  fig.  7,  1941. 

Stems  decumbent,  up  to  50  or  more,  forming  dense  masses,  elongated  and 
slender,  2-6  dm.  long,  3-5  cm.  in  diameter;  spines  yellowish  to  white  or 
pinkish,  the  lower  deflexed  and  flattened  central  spine  2.5-3.5  cm.  long,  the 
other  centrals  and  the  radials  1.5-2.3  cm.  long,  relatively  weak  and  flexible ; 
flowers  unknown. 

Talus  and  ledges  in  precipitous  canyons  of  the  Sonoran  Desert  at  about 
2,000  to  3,000  feet  elevation.  Arizona  at  Mile  161/2,  Marble  Canyon,  Colorado 
River,  Coconino  County,  and  at  Palm  Canyon,  Kofa  Mountains,  Yuma 
County  (Benson  and  Darrow  10870  VA,  B).  The  Palm  Canyon  plant  grew 
on  top  of  a  huge  boulder  where  almost  no  soil  was  present,  and  it  covered  an 
area  three  or  four  feet  in  diameter. 

Type  collection :  "Type  on  limestone  ledge  30  feet  from  the  river's  edge  and  at  the  base 
of  rocky  talus  at  Mile  161/2  Marble  Canyon,  Coconino  Co.,  Arizona  {Clover  and  Jotter  2212). 
Living  Hi)eeimen  at  the  Botanical  Gardens  of  the  University  of  Michigan,  Ann  Arbor, 
Michigan  (Gard.  16S70.  Fig.  7.)"  Specimen  now  pressed. 

4.  Echinocereus  Fendleri  (Engehn.),  Engelm.  ex  Riimpler 

Cereus  Fendleri  Engelm.,  apud  A.  Gray,  1849,  Mem.  Amer.  Acad.,  II.  4 :  51. 
Cereus  Fendleri  var.  paupcrculus  Engelm.,  A.  Gray,  1849,  loc.  cii. 

EcJiinocerevs  Fendleri  (Engelm.),  Engelm.  e.x  TJiiniiiler  in  Forstek,  1885,  Handb.  Cacteenk., 
ed.  2,  p.  801. 


Vol.  XXV]         BENSON :  BETISION  OF  ARIZONA  CACTACEAE  259 

Clumps  not  dense,  tlie  stems  1-5  or  perhaps  more,  not  crowded  against  one 
another,  ovoid,  7-15  cm.  long,  5-10  cm.  in  diameter,  flaccid,  ribs  usually  about 
9-10,  not  markedly  tuberculate  at  the  areoles ;  spines  not  obliterating  the  sur- 
face of  the  stem;  central  spine  1,  at  first  very  dark  brown  to  black,  gray  in 
age,  turned  upward  from  the  base,  tapering,  2-4.5  cm.  long,  stout,  slightly 
curving  upward  the  entire  length,  rigid,  not  flattened ;  radial  spines  9-11, 
similar  to  the  central  but  shorter  and  white  or  pale  gray ;  areole  circular,  not 
bearing  white  felt  at  maturity,  the  felt  rarely  persisting  two  years ;  petals 
magenta  to  purplish,  the  flower  about  5-7  cm.  in  diameter;  floral  tube  about 
12  mm.  long  above  the  ovary ;  fruit  ovoid,  about  3  cm.  long,  spiny,  the  spine- 
clusters  interlocking,  falling  away  as  a  sheet  at  maturity. 

Grassland  and  open  areas  in  the  Rocky  Mountain  Forests  at  5,000  to  7,000 
feet  elevation.  Northern  Arizona  in  Coconino,  Navajo,  and  Apache  counties ; 
southern  Utah ;  northern  New  Mexico. 

Type  collections:  (1.)  C.  Fendleri,  "Santa  Fe,  on  elevated  sandy  plains;  flowering  in 
June."  Fcndler,  Mo.  (2.)  Var.  pauperculus,  apparently  collected  in  the  same  vicinity  by 
Fendler,  Mo. 

4a.  Echinocereus  Fendleri  var.  rectispinus  (Peebles),  L.  Benson,  comb.  nov. 
Echinocereus  recfispmws  Peebles,  1938,  Amer.  Jour.  Bot.,  25 :  675,  f.  1,  3d. 

Stems  1-5,  usually  1-2  dm.  high,  6-9  cm.  in  diameter,  firm,  the  ribs  8-10; 
central  spine  perpendicular  to  the  stem,  straight,  1.2-2.5  cm.  long,  in  some 
areoles  accompanied  by  small  accessory  centrals,  rigid,  rather  stout;  flower 
about  6-6.5  cm.  in  diameter. 

Eocky  slopes  and  benches  in  the  Desert  Grassland  at  3,500  to  5,500  feet 
elevation.  Arizona  from  Nogales  and  the  Santa  Rita  Mountains  to  Ft.  Grant 
and  Cochise  County. 

Type  collection:  "Type:  Peebles  No.  SF  905  (fig.  1),  hills  near  Nogales,  Arizona,  eleva- 
tion 3,900  feet.  May  5,  1935,  represented  by  a  living  plant  under  cultivation  at  U.  S.  Field 
Station,  Sacaton,  Ariz.,  and  by  material  deposited  in  the  U.  S.  National  Herbarium  as 

No.  1729266." 

4b.  Echinocereus  Fendleri  var.  robustus  (Peebles),  L.  Benson,  comb.  nov. 

Echinocereus  rectispinus  Peebles  var.  robustus  Peebles,  1938,  Amer.  Jour.  Bot.,  25 :  675, 

f.  3c. 
Echinocereus  robustus  Peebles,  1940,  Jour.  Wash.  Acad.  Sci.,  30 :  219. 

Stems  5-20,  1.7^.5  dm.  high,  mostly  8-10  cm.  in  diameter,  ribs  8-13,  firm ; 
central  spine  straight,  perpendicular  to  the  stem,  2.5-7.5  cm.  long,  in  some 
areoles  accompanied  by  small  accessory  centrals,  rather  flexible,  slender ; 
flower  usually  7.5-8.5  cm.  in  diameter. 

Rocky  slopes  in  the  Sonoran  and  Chihuahuan  deserts  and  the  lower  part 
of  the  Desert  Grassland  at  2,000  to  4,000  feet  elevation.  Arizona  from  eastern 
Pinal  County  and  eastern  Pima  County  to  the  upper  Gila  River  valley  and 
the  lower  parts  of  Cochise  County.  This  is  the  only  Echinocereus  in  the  desert 
about  Tucson.  Flowering  in  April  and  May.  Closely  related  to  the  next  variety. 


260  CALIFORNIA  ACADEMY  OF  SCIENCES  [Proc.  4th  sek. 

Type  collection:  "Type:  Peebles  No.  SF  896,  Tucson  to  Sabino  Canyon,  Pima  County,  Ari- 
zona, August  27,  1935,  represented  by  a  living  plant  g^o^\^l  under  cultivation  at  Sacaton, 
Ariz.,  and  by  material  deposited  in  the  U.  S.  National  Herbarium  as  No.  1729267." 

4c.  Echinocereus  Fendleri  var.  Boyce-Thompsonii  (Orcutt),  L.  Benson, 

eoinb.  nov. 

Echinocereus  Boyce-Thompsonii  Orcutt,  1926,  Cactography,  (3) :  4. 

Stems  1-10,  1-2.5  dm.  high,  5-7.5  em.  in  diameter,  rather  firm,  ribs  12  or 
commonly  14-18  or  sometimes  more;  central  spine  2.5-4  cm.  long,  accompanied 
in  at  least  some  areoles  by  1  or  2  accessory  centrals,  perpendicular  to  the  stem 
or  more  commonly  deflexed,  straight,  flexible,  very  slender,  remarkably  light- 
colored,  either  straw-colored,  reddish,  or  white  (gray  in  the  two  preceding 
varieties)  ;  flower  usually  about  6-6.5  cm.  in  diameter.  This  variety  is  transi- 
tional in  some  of  its  forms  to  Echinocereus  Engelniannii. 

Rocky  slopes  or  grassy  uplands  in  the  upper  part  of  the  Sonoran  Desert, 
the  Desert  Grassland,  the  Southwestern  Coniferous  Woodland,  and  the  South- 
western Oak  Woodland  at  mostly  2,500  to  5,500  feet  elevation.  Central  Arizona 
from  the  vicinity  of  Seligman  southeastward  just  below  the  Mogollon  Rim  to 
the  upper  Gila  River  Valley.  Flowering  from  March  until  May. 

Type  collection :  "Grounds  of  the  Boyce-Thompson  Southwestern  Arboretum,  near  Su- 
perior, Arizona,  at  an  elevation  of  about  2,300  feet."  The  original  description  was  reprinted 
by  Peebles,  Amer.  Jour.  Bot.,  25:  677.  1938. 

4d.  Echinocereus  Fendleri  var.  Bonkerae  (Thornber  and  Bonker), 

L.  Benson,  comb.  nov. 

Echinocereus  Bonkerae  Thornber  and  Bonker,  1832,  Fantastic  Clan,  p.  71-3,  85,  pi.  oppo- 
site p.  28,  72. 

Clumps  rather  dense  for  the  group,  the  stems  usually  5-15,  1.2-2  dm.  high, 
3.5-6.5  cm.  in  diameter,  rather  firm,  ribs  11  or  13-16 ;  central  spine  6-8  mm. 
long,  accompanied  in  some  areoles  by  1  or  2  accessory  centrals,  perpendicular 
to  the  stem,  rigid,  white  or  pale  gray  tipped  with  brown ;  flower  tending  toward 
purple,  about  6-6.5  cm.  in  diameter,  the  petals  and  sepals  more  nearly  rotate 
than  in  the  other  varieties,  or  even  slightly  reflexed. 

Hills  and  slopes  in  the  Desert  Grassland  and  the  Southwestern  Oak  Wood- 
land at  3,500  to  5,000  feet  elevation;  best  developed  in  shady  situations  where 
leaf -mold  is  present.  Arizona  from  the  vicinity  of  Globe  to  the  Santa  Catalina 
Mountains  and  the  Graham  (Pinaleno)  Mountains.  Specimens  collected  by 
A.  A.  Nichol  in  1940  are  clearly  transitional  between  this  and  var.  Boyce- 
Thompsonii. 

Type  collection:  "We  are  nearing  the  beautiful  Pinal  Mountains  in  southeastern  Ari- 
zona. .  . ."  According  to  A.  A.  Nichol,  the  plant  described  by  Professor  Thornber  was  from 
near  Oracle  at  the  north  base  of  the  Santa  Catalina  Mountains.  Mr.  Nichol  planted  the 
specimens  in  the  University  of  Arizona  Cactus  Garden,  and  in  1939,  the  writer  placed  the 
remains  of  the  last  individual  in  the  University  Herbarium.  They  are  the  closest  approach 
to  a  type.  Coulter,  Contr.  U.  S.  Nat.  Herb.,  3 :  384,  1896,  described  a  plant  probably  of  this 
variety  from  Oracle  (Evans  in  1891),  placing  the  vegetative  material  near  Cereus  ctenoides 


Vol.  XXY]         BENSON :  BEVISIOK  OF  ASIZOXA  CAC TACEAE  261 

Engelm.  (Echinocereus  pectinaius  var.  neomexicanus  above).  Britton  and  Rose,  Cactaceae, 
2:  37,  1922,  published  a  photograpli  of  this  plant  (fig.  45)  as  E.  Fendleri. 

5.  Echinocereus  Ledingii  Peebles 

Echinocererts  Ledingii  Peebles,  1936,  Caet.  and  Succ.  Jour.,  8:  35. 

Clumps  not  particularly  dense ;  longer  stems  cylindrical,  1.5-4  or  5  dm. 
high,  6-9  or  10  cm.  in  diameter,  ribs  12-16,  not  markedly  tuberculate  at  the 
areoles ;  spines  not  completely  obliterating  the  surface  of  the  stem,  light  yellow 
or  straw-color;  central  spine  1,  but  often  accompanied  by  1-3  short  upper 
accessory  centrals,  deflexed  and  strongly  curved  downward  near  the  base, 
about  2  cm.  long,  stout,  rigid,  tapering;  radial  spines  about  10-12,  straight, 
about  9  mm.  long;  areole  circular  or  nearly  so,  not  bearing  white  felt;  petals 
rose-purple,  the  flower  3.5-5  cm.  in  diameter;  floral  tube  about  9  mm.  long 
above  the  ovary;  fruit  pale  green,  spherical,  a  little  less  than  1  inch  in  diam- 
eter, spiny  when  young. 

Slopes  in  the  Southwestern  Oak  Woodland  at  4,500  to  6,000  feet  elevation. 
Southeastern  Arizona  in  the  Quinlan,  Santa  Catalina,  Santa  Rita,  Huachuca, 
Graham  (Pinaleno),  and  Chiricahua  Mountains  in  Pima,  Graham,  Santa 
Cruz,  and  Cochise  counties.  Perhaps  endemic  in  Arizona.  The  localities  are 
based  upon  records  supplied  by  A.  A.  Nichol ;  specimens  have  been  obtained 
only  from  the  Graham  Mountains. 

Type  collection :  "Type  collected  at  about  4,500  feet  elevation  on  slopes  of  Mt.  Graham, 
Pinaleno  Mountains,  Graham  Co.,  Arizona,  Louis  Wankum,  July  11,  1935,  U.  S.  Nat.  Herb. 
No.  1,634,004." 

ECHINOCACTUS 

In  this  genus  only  a  few  changes  are  proposed  now,  and  other  problems  are 
deferred  pending  accumulation  of  additional  data. 

Echinocactus  acanthodes  Lemaire 

Echinocactus  acantJiodes  Lemaire,  1839,  Cact.  Gen.  Nov.  Sp.,  p.  106. 

Echinocactus  viridescens  var.  cylindraceus  Engelm.,  1852,  Amer.  Jour.  Sci.,  II :  338. 

Echinocactus  LeContei  Engelm.,  1856,  Pac.  E.  R.  Eept.,  4:  29. 

Echinocactus  cylindraceus  Engelm.,  1857,  Proc.  Amer.  Acad.,  3  :  275. 

Echinocactus  Wislisenii  Engelm.  var.  LeContei  Engelm.  in  Eothrock,  1878,  Eept.  U.  S. 

Geogr.  Surv.,  6 :  128. 
Ferocactus  LeContei  Britt.  and  EosE,  1922,  Cactaceae,  3  :  129. 
Ferocactus  acanthodes  Britt.  and  Eose,  1922,  loc.  cit. 
Ferocactus  Bostii  Britt.  and  Eose,  1922,  loc.  cit.,  p.  146. 
Echinocactus  Eertrichii  Weinberg,  1929,  Desert,  1 :  40. 
Ferocactus  acanthodes  var.  Bostii  W.  T.  Marshall  in  Marshall  and  Bock,  1941,  Cactaceae, 

p.  148. 

The  common  barrel  cactus  in  western  and  central  Arizona  does  not  seem  to 
have  any  points  of  consistent  difference  from  the  type  common  in  the  Cali- 
fornia deserts.  Previously  (Benson  et  al,  1940)  it  was  treated  as  a  species 
pending  further  investigation,  but  with  the  following  statement,  "Probably 
merely  a  form  of  Echinocactus  acanthodes." 


262  CALIFOENIA  ACADEMY  OF  SCIENCES  [Proc.  4th  ser. 

Type  collections:  (1.)  E.  acanthodes,  type  collected  in  California.  (2.)  Var.  cylindraceus, 
"Found  near  San  Felipe,  on  the  eastern  slope  of  the  California  Mountains."  Colorado 
Desert,  San  Diego  County,  Parry,  Mo.  (3.)  E.  LeContei,  "This  gigantic  .species  was  first 
noticed  by  Dr.  John  L.  LeConte,  on  the  lower  Gila,  where  also  Dr.  C.  C.  Parry  saw  it.  . .  . 
Subsequently,  Dr.  Bigelow  met  with  this  remarkable  plant,  abundantly  from  the  Cactus 
Pass,  at  the  headwaters  of  Williams'  River  down  this  stream  to  the  Colorado,  and  Avest  of 
it  till  E.  polycephalus  took  its  place."  (4.)  F.  Bostii,  "The  tj-pe  is  based  on  a  plant  collected 
in  Lower  California,  40  miles  south  of  the  International  Boundary  Line  (Eost  No.  327)." 
(5.)  E.  ncrtrichii,  "Type  specimen:  Huntington's  Botanischer  Garten  und  Museum,  San 
Marino,  California."  "Type — Weinberg  in  Henry  E.  Huntington's  Botanic  Garden,  San 
Marino,  Calif.  Locality — Tortilla  and  Gila  mountains,  Arizona." 

Echinocactus  polycephalus  Engelm.  var.  xeranthemoides  Coulter 

Echinocactus  polycephalus  Engelm.  var.  xeranthemoides  Coult.,  1896,  Contr.  U.  S.  Nat. 

Herb.,  3 :  385. 
Echinocactus  xeranthemoides  Engelm.  ex  Coult.,  1896,  loc.  cit.,  as  syn. 
Echinocactus  xeranthemoides  Engelm.  ex  Eydb.,  1917,  Fl.  Eocky  Mts.  and  adj.  Plains,  p.  579. 

Stems  rarely  more  than  12,  the  largest  ones  1-2  dm.  in  diameter,  the  clump 
usually  asymmetrial,  that  is,  the  tallest  stem  on  the  edge ;  spines  glabrous, 
yellow  or  pale  gray  tinged  with  red  or  sometimes  pink  or  light  red ;  central 
spines  spreading  at  right  angles  to  the  stem.  (In  the  tj^pical  variety:  stems 
commonly  10-30,  the  larger  ones  2-3  dm.  in  diameter,  the  clump  symmetrical, 
that  is  with  the  largest  stem  in  the  center;  spines  densely  covered  with  felt 
when  young,  red ;  central  spines  spreading  each  in  a  low"  arc.) 

Rocky,  south-facing  ledges  in  the  Southwestern  Coniferous  Woodland  at 
4,000  to  7,000  feet  elevation.  Arizona  in  the  northern  edge  of  Mohave  County 
and  along  the  Colorado  River  system  from  Lee's  Ferry  and  Canyon  Diablo 
to  the  Grand  Canyon  region ;  southern  Utah. 

Type  collection:  "Type,  Siler  of  1881  and  1883  in  Herb.  Mo.  Bot.  Card." 

MAMMILLARIA 

As  in  Echinocactus  and  Opuntia,  although  a  number  of  problems  exist,  data 
for  solution  of  some  of  them  are  inadequate.  Only  the  following  changes  are 
proposed. 

Mammillaria  vivipara  (Nutt.)  HaAv.  var.  aggregata  (Engelm.),  L.  Benson, 

comb.  nov. 

Mammillaria  aggregnia  Engelm.  in  Emory,  1848,  Notes  Mil.  Eeconn.,  p.  146. 
Cereus  aggregatus  Coult.,  1896,  Contr.  U.  S.  Nat.  Herb.,  3  :  396. 
Echinocereus  aggregatus  Eydb.,  1906,  Bull.  Torrcy  Club,  33  :  146. 
Coryphantha  aggregata  Britt.  and  Eose,  1923,  Cactaceae,  4 :  47. 

Low-growing  plant,  the  stems  at  first  solitary  but  later  forming  mounds 
2-6  dm.  high  and  up  to  6  dm.  in  diameter ;  stems  4—5  cm.  in  diameter  at  ma- 
turity, tubercles  cylindrical,  about  8-10  mm.  high,  about  6  mm.  in  diameter, 
the  surface  of  the  stem  obliterated  by  spines;  central  spines  usually  5-6,  white 
tipped  with  brown,  straight,  the  upper  ones  turned  upward,  1.5-2  cm.  long, 
the  lower  ones  spreading  at  various  angles  from  the  stem,  about  6-9  mm.  long. 


Vol.  XXV]         BEN  SON  :  BEVI S  lOX  OF  ABIZON  A  CACTACEAE  263 

m 

tapering",  slender  but  rigid,  not  flattened,  the  centrals  appearing  after  the 
radials  and  not  present  in  yomig  stems;  radial  spines  15-20,  similar  to  the 
centrals,  white,  slightly  brownish  at  the  extreme  tips,  12-18  or  25  mm.  long, 
somewhat  flattened;  petals  usually  lavender  or  pink,  oblanceolate,  rather 
abruptly  acuminate,  4-6  mm.  broad,  the  flower  about  3.5^  cm.  in  diameter ; 
fruit  green  with  a  dull  purplish  tinge,  ellipsoid,  about  2.5  em.  long,  1.8  mm. 
in  diameter. 

River  bottoms,  the  desert  floor,  or  most  frequently  on  grassy  plains  or  rocky 
slopes  in  the  upper  part  of  the  Sonoran  Desert,  the  Desert  Grassland,  and  the 
Southwestern  Oak  Woodland  and  Chaparral  at  3,000  to  5,000  or  6,000  feet 
elevation.  The  eastern  half  of  southern  Arizona;  largely  south  of  the  Mogollon 
Rim ;  New  Mexico  on  the  Gila  River  drainage ;  Sonora.  Flowering  in  May 
and  June. 

Type  collection:  "Oct.  18,  1846.  Head  waters  of  the  Gila,  6,000  feet  above  tlie  sea." 
Interpretations  of  the  type  have  alternated  between  this  plant  and  Echinocereus  triglocM- 
diatus  var.  melancanthus.  Engelmann,  loc.  cit.,  first  interpreted  the  plant  as  a  member  of 
the  Mammillaria  vivipara  complex,  and  (as  pointed  out  by  Peebles  apud  Kearney  and 
Peebles,  U.  S.  Dept.  Agric.  Misc.  Publ.,  (423)  :  605,  1942,  the  original  drawing  shows 
crossed  spirals  indicating  tubercles  on  the  stem.  The  type  was  not  preserved,  or  was  lost. 

Mammillaria  vivipara  var.  arizonica  (Engelm.)  L.  Benson,  comb,  no  v. 

Mammillaria  arizonica  Engelm.  in  Brew,  and  Wats.,  1876,  Bot.  Calif.,  ed.  1,  1:  244. 
Cactus  radiosus   (Engelm.)   Coult.  var.  arizonica  Coult.,  1894,  Contr.  U.  S.  Nat.  Herb., 

3:  121. 
Mammillaria  radiosa  Engelm.  var.  arizonica  K.  Schum.,  1898,  Gesamtb.  Kakteen,  p.  481. 
Con-ypliantlia  arizonica  Britt.  and  Eose,  1923,  Cactaceae,  4:  45. 

Stems  solitary  at  first,  later  forming  great  mounds ;  spines  predominantly 
dark  brown;  lower  central  spines  2,  one  of  these  spreading  perpendicularly 
(to  the  areole),  the  other  turned  downward;  petals  deep  pink,  linear-lance- 
olate, gradually  acuminate,  2.5-4  mm.  broad. 

Rocky  places  in  the  Southwestern  Coniferous  Woodland  and  the  Rocky 
Mountain  Forests  at  4,000  to  8,000  feet  elevation.  Southwestern  Utah ;  north- 
ern Arizona  from  Mohave  County  to  Apache  County  and  southward  to  the 
Mogollon  Rim  and  the  White  Mountains,  and  perhaps  farther  in  the  higher 
mountains. 

Type  collection :  "On  sandy  and  rocky  soil  in  Northern  Arizona,  from  the  Colorado  east- 
ward (Coues,  Palmer,  F.  BiscJioff),  and  into  Southern  Utah  (/.  E.  Johnson). . ." 

Mammillaria  vivipara  var.  deserti  (Engelm.),  L.  Benson,  comb.  nov. 

Mammillaria  chlorantha  'Enge\m.apud  Eothr.,  in  Wheeler,  1878,  U.  S.  Geogr.  Surv.  W.  of 

lOOthMerid.,  6:  127. 
Mammillaria  deserti  Engelm.  in  S.  Wats.,  1880,  Bot.  Calif.,  2  :  449. 

Cactus  radiosus  (Engelm.)  Coult.  var.  deserti  Coult.,  1894,  Contr.  U.  S.  Nat.  Herb.,  3  :  121. 
Cactus  radiosus  var.  chloranthus  Coult.,  1894,  loc.  cit. 
Cactus  radiosus  var.  Alversonii  Coult,  1894,  loc.  cit.,  p.  122. 
Mammillaria  Alversonii  Zeissold,  1895,  Monatsschr.  Kakteenk.,  5 :  70. 
Mammillaria  radiosa  Engelm.  var.  deserti  K.  Schum.,  1898,  Gesamtb.  Kakteen,  p.  481. 


264  CALIFORNIA  ACADEMY  OF  SCIENCES  [Pkoc.  rtXH  skr. 

Mammillaria  radiosa  var.  chlorantha  K.  Schum.,  1898,  loc.  cit. 

Mammillaria  radiosa  var.  Alversonii  K,  ScHUM.,  1898,  loc.  cit. 

Coryphantha  chlorantha  Britt.  and  Rose,  1923,  Cactaceae,  4:  43. 

Coryphantha  arisonica  Britt.  and  Rose,  1923,  loc.  cit.,  p.  45. 

Coryphantha  deserti  Britt.  and  Rose,  1923,  loc.  cit.,  p.  45. 

Mammillaria  arisonica  Engelm.  var.  deserti  Engelm.  ex  Davidson  and  Moxley,  1923,  Fl. 

S.  Calif.,  p.  244. 
Mammillaria  arisonica  var.  Alversonii  Engelm.  ex  Davidson  and  Moxley,  1923,  loc.  cit. 
Coryphantha  Alversonii  Orcutt,  1926,  Cactography,  (3) :  3. 

A  robust  type  with  the  stems  usually  solitary  or  sometimes  few,  these  7-20 
cm.  high,  5-7.5  cm.  in  diameter ;  central  spines  3-14,  usually  white  tipped  with 
brown,  straight,  spreading  at  all  angles,  12-18  mm.  in  length,  averaging  long, 
tapering,  stout,  rigid,  not  flattened;  radial  spines  about  20,  similar  to  the 
centrals  but  more  slender,  white ;  petals  straw-colored,  yellow,  pink,  or  purple, 
the  flowers  2.5-3.5  cm.  in  diameter. 

Rocky  slopes  in  the  Mojavean  Desert  at  1,500  to  3,500  feet  elevation.  Cali- 
fornia in  the  ranges  of  the  eastern  and  southern  parts  of  the  Mojave  Desert; 
southern  Nevada;  southwestern  Utah;  northwestern  Arizona  in  northern  and 
western  Mohave  County.  A  variable  plant  possibly  to  be  segregated  into  more 
than  one  variety  with  accumulation  of  additional  data.  The  characters  of  the 
proposed  segregates  seem  inconsistent  in  their  association. 

Type  collections:  (1.)  M.  chlorantha,  "Southern  Utah,  east  of  Saint  George,  Dr.  Parry. 
I.  E.  Johnson."  (2.)  M.  deserti,  "At  Ivanpah,  30  miles  northeast  of  San  Bernardino,  in  one 
of  the  mountain  ranges  stretching  into  the  desert,  S.  B.  Parish."  Mo.  Ivanpah  is  in  the  New 
York  Mountains,  about  140  miles  northeast  of  San  Bernardino,  California.  (3.)  Var. 
Alversonii,  "Type,  Alversou's  specimens  in  the  Herb.  Mo.  Bot.  Gard.  and  Herb.  Coulter  . . .. 

SOUTHERN  CALIFORNIA  (A.  H.  Alverson  of  1892) The  decidedly  pink  flowers  were 

sent  by  Mr.  S.  B.  Parish  from  specimens  growing  in  cultivation  in  San  Diego,  and  are  not 
from  the  original  collection  of  Mr.  Alverson." 

Mammillaria  Heyderi  Miihlenpf  ordt 

Mammillaria  Heyderi  Muhlenpfordt,  1848,  Allg.  Gartenz.,  16:  20. 

Cactus  Heyderi  Kuntze,  1891,  Rev.  Gen.  et  Sp.  PI.,  1:  260. 

N eomammillaria  Heyderi  Britt.  and  Rose,  1923,  Cactaceae,  4:  75. 

Obscure  depressed  cactus  arising  only  1-5  cm.  above  ground  level;  stem 
solitary,  shaped  like  a  turnip,  flat-topped,  half  or  two-thirds  subterranean, 
5-8  cm.  long,  7-12  cm.  in  diameter,  the  tubercles  conical,  6-10  mm.  high,  5-7 
mm.  in  diameter;  juice  of  the  plant  milky;  central  spines  2-3,  white  tipped 
with  dark  brown,  straight  and  subequal,  6-8  mm.  long,  tapering,  slender, 
rigid,  not  flattened;  radial  spines  10-15;  petals  pink;  sepals  not  fringed; 
flower  10-12  mm.  in  diameter. 

Limestone  hills  in  the  Chihuahuan  Desert  or  the  Desert  Grassland  at  3,500 
to  5,000  feet  elevation.  Along  the  Mexican  Border  in  Cochise  County  near 
Bisbee  {Peebles  SF  922,  Sac,  JJA)  and  in  the  Peloncillo  Mountains  (L.  Ben- 
son 10272,  TJA,  B).  One  of  several  Chihuahuan  Desert  plants  barely  reaching 
southeastern  Arizona  and  there  restricted  to  favorable  limestone  soils. 

Type  collection :  none  given. 


Vol.  XXV]         BENSON:  BEVISION  OF  ARIZONA  CACTACEAE  2Go 

Mammillaria  Heyderi  var.  MacDougalii  (Rose)  L.  Benson,  comb.  nov. 

Mavimillaria  MacDougalii  Eose  in  Bailey,  1916,  Stand.  Cyclop.  Hort.,  ed.  1.  4:  1982. 
Neomammillaria  MacDougalii  Britt.  and  Eose,  1923,  Caetaceae,  4:  74. 

Stems  5-7.5  or  12.5  cm.  long,  7.5-17.5  cm.  in  diameter;  central  spines 
straight  or  the  principal  one  curving  gradually  outward ;  petals  cream-colored 
or  light  yellow ;  sepals  fringed ;  flower  about  17-20  mm.  in  diameter. 

South  exposures  of  rocky  hill  crests  in  the  Desert  Grassland  and  the  South- 
western Oak  Woodland  at  4,000  to  6,000  feet  elevation;  often  in  crevices  of 
rocks  or  in  rocky  soil.  Arizona  from  the  Baboquivari  Mountains  eastward 
to  the  Santa  Catalina  Mountains  and  to  southern  Cochise  County ;  probably  in 
adjacent  Sonora.  Often  associated  with  Echinocereus  pectinatus  var.  rigi- 
dissimus,  the  Arizona  rainbow  cactus.  Flowering  late  in  March  and  in  April. 

Tj^pe  collection :  "Common  in  the  mountains  about  Tucson,  Ariz.,  where  it 
was  collected  by  D.  T.  MacDougal,  for  whom  it  is  named." 

The  dimorphic  fruits  of  some  of  the  typical  Mammillaria  species,  including 
If.  Heyderi  and  the  variety  MacDougalii  have  been  well-known  for  some  time, 
and  Peebles,  Cact.  and  Succ.  Jour.,  13 :  143-5,  1941,  has  reported  and  illus- 
trated similar  types  (clavate-red  and  spherical-green)  in  Mammillaria  micro- 
carpa.  They  occur  also  in  Mammillaria  fasciculata  (15  miles  west  of  Silver 
Bell,  Pima  County,  Arizona,  L.  Benson  10606,  VA,  B). 

The  plants  described  previously  as  Mammillaria  microcarpa  Engelm.,  M. 
fascicidata  Engelm.,  31.  Wilcoxii  Tourney,  M.  viridiflora  (Britt.  and  Rose) 
Boed.,  and  M.  Mainae  K.  Brandegee  are  in  need  of  further  study,  but  at 
present  data  are  inadequate. 

LITERATURE  CITED 

Benson,  Lyman 

1939.  Notes  on  taxonomie  techniques.  Torreya,  39 :  73-75. 

1941.  Taxonomie  studies.  American  Journal  of  Botany,  28:  358-364. 

1942.  The  relationship  of  Ranunculus  to  the  North  American  Floras.  American  Journal  of 
Botany,  29 :  491-500. 

Benson,  Lyman,  J.  J.  Thornber,  A.  A.  Nichol,  and  Lucretia  Breazale  Hamilton 

1940.  The  cacti  of  Arizona.  University  of  Arizona  Biological  Science  Bulletin  No.  5.  134 
pp.  52  pi.  Tucson,  Arizona. 

Peebles,  Eobert  H. 

1941.  Variation  in  the  fruit  of  Mammillaria  microcarpa.  Cactus  and  Succulent  Journal, 
13:  143-145. 

1942.  Preservation  of  cactus  material.  Cactus  and  Succulent  Journal,  14:  3-8. 

Shreve,  Forrest 

1936.  The  plant  life  of  the  Sonoran  Desert.  Scientific  Monthly,  42  :  195-213. 
1942.  The  Desert  Vegetation  of  North  America.  Botanical  Eeview,  8:  195-246. 

Shantz,  Homer  Leroy  and  Eafael  Zon 

1924.  Natural  vegetation.  Atlas  of  American  Agriculture,  U.  S.  Dept.  of  Agriculture, 
Washington,  D.  C. 


EXPLANATION  OF  PLATE 

Plate  25 

Opunlia  and  Echinocereus :  A,  Opuntia,  1,  Opitnlia  covipressa  var.  macro- 
rhiza,  collected  at  Pearce,  Arizona,  by  A.  A.  Nichol  in  1940,  2,  Opuntia  com- 
pressa  var.  macrorhiza,  from  sand  hills  near  Guion,  Arkansas,  Demaree  23490, 
3,  Opuntia  compressa,  from  the  Atlantic  seaboard,  received  from  cultivation  in 
Massachusetts;  B.  Echinocereus  Engelmannii  var.  Xicholii,  growing  in  the 
Ajo  Mountains,  Arizona.  (Photograph  A  by  Eobert  A.  Darrow;  B  by  Walter 
S.  Phillips.) 


[  2<H'  ]  ^ 


PROC.  CALIF.  ACAD.  SCI..  4TH  SERIES.  VOL.  XXV.   NO.    10 


[BENSON]    PLATE    25 


[267] 


PROCEEDINGS 

OF  THE  -£ 

CALIFORNIA  ACADEMY  OF  SCIENCES   l-^ 

Fourth  Series 
Vol.  XXV,  No.  1 1,  pp.  269-290,  4  text  figs.  November  10,  1944 


ALICE  EASTWOOD 
SEMI-CENTENNIAL  PUBLICATIONS 

No.  11 
ENDEMISM  IN  CREPIS 

BY 

ERNEST  B.  BABCOCK 

Professor  of  Genetics 
University  of  California  ' 

I.  INTRODUCTION 

ONE  RESULT  of  the  exteiisive  investig-ations  on  the  genus  Crepis  has  been  the 
formulation  of  a  fairly  definite  hypothesis  concerning  the  origin,  devel- 
opment and  migrations  of  this  essentially  monophj^letic  group  of  plants.  This 
hypothetical  history  of  the  one  hundred  and  ninety-six  species  of  Crepis  has 
been  found  to  be  consistent  with  a  great  mass  of  other  evidence  concerning  the 
origin  and  migrations  of  floras  in  the  northern  hemisphere.  A  review  of  this 
evidence. together  with  a  detailed  discussion  of  the  data  on  Crepis  will  be 
presented  in  a  monograph  which  is  in  press. 

The  criteria  of  phylogenetic  relations  in  Crepis  represent  five  different  dis- 
ciplines. The  primary  basis  for  classifying  all  the  species  into  twenty-seven 
sections  and  arranging  the  sections  in  an  approximate  phyletie  series  is  com- 
parative morphology.  Second  in  importance  have  been  the  data  on  chromo- 
some numbers  in  one  hundred  and  thirteen  of  the  species  (cf.  Babcock  and 
Jenkins,  1943).  Thirdly,  the  cytogenetic  analysis  of  certain  interspecific 
hybrids  has  confirmed  the  conclusions  reached  concerning  the  phyletie  rela- 
tions of  the  parent  species  as  determined  on  the  basis  of  comparative  morphol- 
ogy and  chromosome  numbers,  and  has  revealed  the  evolutionary  process  by 
which  reduction  in  chromosome  number  in  the  series  6-5-4-3  has  been  accom- 
plished (cf.  Babcock,  1942,  pp.  176-178;  Tobgy,  1943).  Fourthly,  purely 
genetic  data  on  many  interspecific  hybrids  have  substantiated  the  grouping  of 

[  269  ] 


Fig.  1.  Crepis  geracioides  Hausskn.  a,  plant,  XV2;  i,  flower-head,  x  2 ;  c,  a  floret  lacking 
tlic  ovary,  x  4  ca.;  C,  trielionie  from  the  corona-tube,  x  100  ca.;  d,  anther-tube  slit  and  opened 
out,  x8  ca.;  e,  detail  of  anther-appendages,  x  32  ca.;  f,  g,  two  achenes  and  a  pappus-seta 
from  each,  x  8  ca.;  h,  W,  two  inner  involucral  bracts  showing  outer  and  inner  faces  respec- 
tively, X  4  ca.;  i,  haploid  set  of  chromosomes,  x  1500. 


Fig.  2.  Crepis  senecioides  Delile.  a,  b,  c  and  1-,  plants  and  parts,  x  i^ ;  d,  m,  florets  lacking 
the  ovary,  x4  ca.;  e,  n,  anther-tubes  slit  and  opened  out,  x  8  ca.;  f,  o,  details  of  anther- 
appendages,  X  32  ca.;  g,  h,  p,  q,  aehenes  and  single  pappus-setae,  X  8ca.;  r,  haploid  set  of 
chromosomes,  x  1500. 


272  CALIFOBXIA  ACADEMY  OF  SCIENCES  [Proc.  4th  ser. 

the  species  into  sections  on  a  morphological  basis ;  and  the  genetic  analysis 
of  hybrids  has  shown  the  importance  of  grene  mutation  as  a  cause  of  speeiation 
in  Crepis.  Fifthly,  a  study  of  geographic  distribution  in  the  genus  as  a  whole, 
taken  section  by  section,  when  considered  in  the  light  of  well-establislied  paleo- 
botanical  evidence,  has  led  to  the  formulation  of  a  phylogenetic  history  of 
the  genus. 

In  order  to  give  the  reader  a  concrete  notion  of  the  morphological  contrasts 
between  the  most  primitive  and  most  advanced  types  of  Crepis  species,  one  of 
each  is  illustrated  (figs.  1  and  2).  The  differences  in  size  and  habit  of  the 
plant,  in  size  of  the  heads,  florets,  fruits,  etc.,  are  so  obvious,  it  seems  hardly 
necessary  to  point  them  out.  The  primitive  species,  C.  geracioides,  is  a  montane 
perennial  which  is  endemic  in  the  southern  Balkan  Peninsula.  The  advanced 
species,  C.  senecioides,  is  a  precocious,  desert  annual  which  has  been  collected 
at  a  number  of  stations  scattered  along  the  Egyptian-Tunisian  littoral.  Be- 
cause of  its  wide  distribution  it  is  not  considered  to  be  an  endemic  species  in 
the  restricted  sense  in  which  that  term  is  used  in  the  present  paper.  But,  since 
it  is  the  most  advanced  species  in  the  genus  on  morphological  grounds,  it  has 
been  used  to  illustrate  the  extreme  reduction  in  size  which  has  accompanied 
development  of  the  annual  habit,  extreme  precocity,  and  remarkable  longevity 
of  the  tiny  seeds  (achenes) ,  all  of  which  adapt  it  so  well  to  desert  conditions. 
The  fact  that  these  two  species,  which  represent  the  earliest  and  latest  stages 
in  a  very  long  developmental  history,  now  exist  on  opposite  sides  of  the  Medi- 
terranean Sea,  which  is  only  some  600  kilometers  (400  miles)  wide  in  this 
region,  is  sufficient  to  pique  one's  curiosity  as  to  whence  and  how  they  came 
to  be  there. 

Stated  as  briefly  as  possible,  the  answer  is  as  follows.  Crepis  originated  in 
Central  Asia  early  in  the  Tertiary  period.  Some,  at  least,  of  the  most  primitive 
species  existed  at  low  elevations,  under  mesophytic  conditions.  But  migration 
into  higher  altitudes,  accompanied  by  speeiation,  must  have  occurred  during 
the  early  development  of  the  genus  and  before  the  extensive  migrations  took 
place  to  the  northeast,  southeast,  northwest  and  southwest  from  Central  Asia. 
The  migrations  to  the  southwest  affected  a  much  larger  number  of  Crepis 
species  than  the  other  three  migrations.  The  general  trend  of  the  southwesterly 
migration  passed  through  Turkestan  and  Iran  and  then  followed  one  of  three 
main  routes:  (1)  through  northern  Asia  Minor  into  the  Balkan  Peninsula 
and  southern  Europe;  (2)  through  southern  Asia  Minor,  Iraq,  Syria  and 
Palestine  into  Egypt;  and  (3)  through  southern  Arabia  into  Abj^ssinia  and 
tropical  Africa.  These  migrations  were  accompanied  or  followed  by  increas- 
ing dessication  in  southwestern  Asia  and  the  Mediterranean  region  and  this, 
together  with  the  many  topographical  changes  Avhich  occurred,  resulted  in 
increased  speeiation  throughout  that  general  region.  Thus  C.  geracioides 
almost  certainly  migrated  from  Central  Asia  around  the  south  end  of  the 
Caspian  Sea,  through  northern  Asia  Minor,  into  the  Balkan  Peninsula  where 
it  still  exists  in  tliree  localities.  But  C.  senecioides  almost  certainly  developed 


Vol.  XXV]  BABCOCK:  ENDEMISM  IN  CEEPIS  273 

where  it  now  exists  from  some  less  advanced  ancestor  which  was  unable  to 
tolerate  the  increasing  aridity  of  the  climate.  That  ancestor,  in  turn,  probably 
developed  from  a  still  more  primitive  type  somewhere  in  the  Iran-Iraq  region. 
A  presentation  of  the  general  background  of  evidence  for  these  rather  dog- 
matic statements  must  await  the  publication  of  the  monograph. 

One  of  the  most  interesting  aspects  of  the  geographic  distribution  of  Crepis 
is  the  large  amount  of  endemism  present  and  the  distribution  of  the  endemic 
areas.  It  is  the  purpose  of  the  present  paper  to  examine  this  evidence  in  order 
to  determine  to  what  extent  it  agrees  with  the  phyletic  history  briefly  outlined 
above,  which  was  worked  out  for  the  genus  as  a  whole.  At  the  same  time  it  may 
be  possible  to  derive  some  conclusions  concerning  endemism  in  Crepis  which 
will  have  general  significance. 

The  present  study  is  confined  to  the  Old  World  species  inasmuch  as  the 
data  on  endemism  in  the  New  World  species  have  already  been  reported  (Bab- 
cock  and  Stebbins,  1938).  With  reference  to  the  history  of  the  genus  as  a 
whole,  the  conclusion  was  reached  that  all  of  the  American  species  either 
migrated  from  Asia  across  the  Beringian  land-bridge  before  the  Pleistocene 
glaciation  could  prevent  such  migrations,  or  that  they  were  derived  from 
species  of  Asiatic  origin. 

The  term  endemic,  in  its  usual  biological  connotation,  is  applied  to  a  plant 
or  animal  which  is  restricted  in  its  distribution  to  a  single  country,  region  or 
area.  This  definition  becomes  much  more  definite  as  soon  as  the  size  of  the 
region  or  area  is  stated.  During  the  years  spent  in  accumulating  data  of  vari- 
ous kinds  concerning  Crepis  species,  the  writer  came  to  realize  that  a  great 
many  of  them  are  endemic  in  the  general  sense  defined  above ;  and  that  a  con- 
siderable number  of  them  are  also  of  very  restricted  distribution.  Some  of 
these  restricted  species  are  insular,  being  confined  to  one  or  other  of  such 
islands  as  Crete,  Sicily,  Majorca,  Madeira  or  Fuerteventura  and  Lanzarote 
of  the  Canary  Archipelago.  Others  are  restricted  to  a  single  limited  district 
on  a  continent,  as  in  the  case  of  the  22-chromosome  forms  of  C.  monticola  and 
C.  pleurocarpa,  two  of  the  oldest  endemic  American  species  of  Crepis  (cf. 
Babcock  and  Stebbins,  1938) .  Still  others  are  similarly  restricted  by  ecological 
factors,  but  they  occur  in  two  or  more  small  districts  and  these  may  be  either 
near  together  or  widely  separated  within  a  given  endemic  area.  An  example  of 
the  latter  is  C.  geracioides;  but  the  three  localities  at  which  C.  geracioides  is 
known  to  occur  are  separated  by  a  maximum  distance  of  approximately  150 
kilometers  (90  miles) .  Hence  this  species  can  hardly  be  described  as  a  narrow 
endemic  except  in  a  purely  relative  sense,  as  when  compared  with  such  a  wide- 
spread species  as  C.  senecioides. 

In  order  to  obtain  more  definite  information  as  to  the  amount  and  possible 
significance  of  endemism  in  Crepis,  it  was  decided  to  set  arbitrarily  a  lower 
limit  of  fifty  kilometers  (30  miles)  and  an  upper  limit  of  150  kilometers  (90 
miles) ;  and  to  classify  all  the  species  whose  maximum  straight-line  distance 
between  known  localities  falls  within  the  lower  limit  as  "narrow  A"  ;  all  those 


274  CALIFOBNIA  ACADEMY  OF  SCIENCES  [Proc.  4th  ser. 

whose  maximum  distance  between  known  localities  exceeds  the  upper  limit  as 
"wider" ;  while  those  falling  between  the  two  limits  were  classed  as  "narrow 
B."  It  was  found  that  there  were  so  few  species  in  the  "narrow  B"  group  that, 
for  the  sake  of  simplicity,  the  "narrow^  B"  and  "wider"  groups  were  lumped 
together,  thus  recognizing  only  two  categories,  "narrow"  and  "wider,"  with 
the  dividing  line  at  50  kilometers. 

On  this  basis  it  was  found  that  there  are  forty-eight  species  of  Crepis  for 
which  the  available  data  indicate  a  "narrow"  distribution.  Twenty-eight  of 
these  are  known  from  only  one  locality'  while  the  other  twenty  are  known  from 
two  or  more  localities.  Of  the  twenty-eight  that  are  known  from  a  single  lo- 
cality, twentj'-f our  are  known  to  the  writer  from  just  one  collection ;  but  the 
other  four  are  known  to  have  been  collected  several  times  at  the  one  locality. 
Three  of  these  four  localities  are  mountain  peaks  in  Greece ;  the  other  is  the 
eastern  promontory  of  Madeira  Island.  The  twenty-four  localities  at  which 
only  a  single  collection  has  been  made  are  mostly  in  regions  w^hich  have  been 
less  thoroughly  explored  botanically,  such  as  Asia  Minor,  the  Caucasus, 
Turkestan  and  tropical  Africa.  Many  of  these  localities  are  in  mountainous 
countrj'^  and  some  are  mountain  peaks.  Since  many  of  the  "wider"  endemics 
are  known  to  be  very  local  in  their  distribution,  and  half  of  the  "narrow"  ones 
are  known  from  two  or  more  collections,  it  is  fairly  probable  that  the  majority 
of  the  species  known  from  only  one  collection  will  turn  out  CA'cntually  to  be 
actually  "narrow"  as  here  defined.  At  any  rate  the  purposes  of  the  present 
paper  can  be  served  better  by  recognizing  them  as  "narrow"  than  by  ignor- 
ing them. 

Classification  of  the  remainder  of  the  Crepis  species  as  endemic  or  not  en- 
demic was  accomplished  by  defining  a  series  of  endemic  areas  and  then  by 
accepting  as  endemic  only  those  known  to  be  restricted  to  a  single  area.  Many 
of  these  areas  are  widely  recognized  as  rich  in  endemic  species  of  plants.  By 
this  method  ninetj^-one  species  were  selected  as  "wider"  endemics.  This  does 
not  mean  that  these  species  are  continuously  distributed  over  a  district  more 
than  150  km.  (90  miles)  in  diameter.  On  the  contrary  these  "wider"  endemics 
are  all  more  or  less  local  in  their  distribution ;  although  a  few  are  fairly 
abundant  in  certain  districts.  Thus  in  the  case  of  C.  geracioides  it  happens 
that  the  three  known  localities  are  about  150  kilometers  distant  from  one  an- 
other ;  whereas  C.  terglouensis,  another  very  primitive  endemic,  is  known  from 
at  least  twenty  high  alpine  stations  ranging  from  south-central  Switzerland 
to  central  Tirol  and  Upper  Austria.  In  marked  contrast  with  these  are  such 
insular  species  as  C.  cretica  and  C.  canariensis  which  are  abundant  on  the 
islands  where  they  exist.  A  few  species  have  been  omitted  which  occur  mostly 
in  one  endemic  area  but  are  also  represented  in  another  and  hence,  on  a  less 
restricted  criterion,  might  have  been  considered  as  endemic.  Interesting  ex- 
amples are  C.  pygmaea,  a  very  primitive  12-chromosome  species  of  Spain,  the 
Pyrenees,  the  southwestern  Alps  and  northern  Italy;  C.  smyrnaea,  another 
12-chromosome  primitive  species  known  from  only  three  localities  in  southern 


YoL.XXV]  BABCOCK:  EXDEMISM  IN  CEEPIS  275 

Greece  and  western  Asia  Minor;  C.  pontana,  the  most  primitive  10-chromo- 
some  species  of  Crepis,  which  occurs  from  the  eastern  Alps  to  the  western  part 
of  the  Balkan  Peninsula ;  and  C.  albida,  the  next  most  primitive  10-chromo- 
some  species  which  is  polymorphic  and  is  distributed  from  the  Grand  Atlas 
Mts.,  in  Morocco  through  Spain,  the  mountains  of  southern  France  and  the 
Maritime  Alps.  It  becomes  difficult  to  know  where  to  draw  the  line,  if  one 
begins  to  go  beyond  the  limits  of  a  single  endemic  area.  But,  on  this  restricted 
basis,  there  are  still  one  hundred  and  thirty-three  endemic  species  which  is 
72  per  cent  of  the  one  hundred  and  eighty-five  Old  World  species. 

II.  THE  DISTRIBUTION  OF  THE  ENDEMIC 
REGIONS  AND  AREAS  FOR  CREPIS 

The  endemic  regions  and  areas  for  Crepis  are  defined  in  Table  1  and  shown 
on  the  accompanying  map  (fig.  3).  On  this  map  the  large  size  of  the  tropical 
African  area  (no.  19)  will  be  one  of  the  first  things  to  catch  the  attention.  It 
should  be  emphasized  that,  in  the  latter  area,  all  of  the  eighteen  endemic 
species  found  there  are  extremely  local  in  distribution.  All  but  three  occur 
on  only  one  or  two  mountains  or  in  a  single  mountain  range.  These  mountains 
and  ranges  are  shown  as  solid  circles  or  narrow  ellipses  on  the  map.  The  other 
three  species  are  knoAvn  from  only  single  stations  which  are  at  low  elevation 
and  these  are  shown  as  solid  squares  on  the  map.  Following  is  a  key  to  the 
symbols  used  to  designate  these  locations  in  area  No.  19  and  three  others  in 
area  No.  10. 

The  long  winged-shaped  ellipse  in  area  No.  10  is  the  Pamir-Ferghana-Altai 
system  of  mountains. 

KEY  TO  MAP  SYMBOLS 

Regions  are  indicated  by  Roman  numerals.  Areas  are 
indicated  by  Arabic  numerals. 

Mountains  shown  hy  solid  circles  and  ellipses: 

AT.  Ala-tau  Mts.,  Central  Asia. 

CH.  Mt.  Chirinda,  southeastern  Africa. 

DA.  Darvas  Mts.,  eastern  Kussian  Turkestan. 

CU.  Congo-Uganda  plateau  north  of  Lake  Kiwu,  Africa. 

HI.  Himalaya  Mts.,  Kashmir  region,  high  alpine. 

KE.  Mt.  Kenya,  eastern  tropical  Africa. 

KI.  Mt.  Kilimanjaro,  eastern  tropical  Africa. 

KU.  Mountains  south  of  Lake  Ki'\vu,  tropical  Africa. 

MC.  Mt.  Cameroon,  western  tropical  Africa. 

ME.  Mt.  Meru,  adjacent  to  Kilimanjaro. 

NY.  Mountains  of  Nyasaland  and  adjacent  Congo. 

EM.  Mt.  Kungwe  and  Mt.  Mbeya,  tropical  Africa. 

UK.  Urundi  Province,  Kisozi,  tropical  Africa. 

UL.  Uluguru  Mts.,  Tanganyika  Prov.,  Africa. 

UT.  Ulu-tau  Mts.,  northern  Eussian  Turkestan. 


176 


CALIFOBXIA  ACADEMY  OF  SCIEXCES  [Proo.  4th  ser. 


Single  lowland  stations  in  Africa: 

EC.     Elizabethville,  southeastern  Belgian  Congo. 
FU.     Fort  Portal,  Uganda. 
CA.     Cului,  Angola. 

TABLE  1 
Endemic  Regions  and  Areas  for  Old  World  Crepis 


Hegions 
I.  Southwestern  Europe;  Alps;  north- 
western Morocco 

II.  Balkan  Peninsula 
III.  Asia  Minor ;  Syria-Palestine ;  Cri- 
mea 

IV.  Caucasus ;  Iran 

V.  Turkestan-Pamir-Altai  region 

VI.  Northwestern  India 

VII.  Southeastern  Asia 

VIII.  Eastern  Siberia 
IX.  Northern  Japan 
X.  Abyssinia ;  Eritrea ;  southwestern 
Arabia 

XI.  Africa,  tropical 
XII.  Lybia- Morocco -Cadiz   (littoral)  ; 
southern    and    western    maritime 
Italy  and  southern  France;   Ma- 
deira and  Canary  Islands 


Areas 

1.  S.  Spain  and  N.W.  Morocco 

2.  N.E.  Spain  and  "W.  France 

3.  European  Alps 

4.  Balkan  Peninsula 

5.  Asia  Minor 

6.  Syria  and  Palestine 

7.  Crimea 

8.  Caucasus  region 

9.  Iran 

10.  Turkestan  (E.  and  N.  Eussian)  and  the 
Pamir-Altai  mountains 

11.  N.AV.  Himalaya  Mountains,  high  alpine 

12.  N.W.  India-Baluchistan,  low  montane 

13.  S.E.  Asia  (=  S.E.  Tibet,  S.W.  China,  N. 
Burma,  Laos,  Annam) 

14.  E.  Siberia  (Okhotsk  district) 

15.  Hokkaido-Sakhalin 

16.  S.W.  Arabia  in  Yemen  Province 

17.  Eritrea 

18.  N.  Abyssinia,  mountains 

19.  Africa,  tropical  (isolated  localities) 

20.  Lybia  (littoral) 

21.  Tunisia  (middle  and  southern) 

22.  E.  Algeria  (littoral) 

23.  S.  Italy  and  Toscana,  Sicily,  Sardinia, 
Corsica ;  S.  France 

24.  W.  Algeria,  N.  Morocco,  Cadiz  (mostly 
littoral) 

25.  Madeira  and  Canary  Islands 


Tlie  distribution  of  the  twelve  endemic  regions  defines  the  east-west  distri- 
bution of  the  genus  as  a  whole  almost  completel}',  the  gaps  between  the  en- 
demic regions  being  occupied  more  or  less  by  the  various  widespread  spet^ies. 
In  this  connection  it  should  be  repeated  that  the  total  evidence  on  geographic 
distribution  in  Crepis,  considered  in  relation  to  the  evidence  on  phylogeny, 
definitely  indicates  that  the  center  of  origin  and  early  development  of  Crepis 
was  in  Central  Asia,  in  the  Altai-Tian  Shan  mountain  region  in  the  north- 
eastern part  of  region  V  (fig.  3) ;  and  that  the  present  wide  distribution  of 
the  genus  was  accomplished  through  migration  in  four  general  directions 
from  the  center.  One  of  these  trends  was  towards  the  northwest,  into  what  is 
now  Russia.  Only  five  species  exist  today  in  northern  Europe  as  a  result  of  that 


Vol.  XX Y] 


BABCOCK :  ENDEMISM  IN  CBEPIS 


277 


trend  of  mig-ration  and  none  of  these  is  endemic.  Another  migration  trend 
was  to  the  east  and  northeast,  resulting  in  two  endemics  in  Japan,  one  in  the 
Okhotsk  district  of  Siberia,  (region  VIII,  fig.  3),  and  all  of  the  American 
species.  Still  another  trend  was  to  the  southeast,  resulting  in  the  nine  endemic 
species  found  in  region  VII.  Most  important  of  all  the  migration  trends  from 
Central  Asia,  in  terms  of  the  number  of  species  involved,  was  the  southwest- 


Fig.  3.  Map  of  the  Old  World  showing  the  endemic  regions  (Eoman  numerals)  and  areas 
(Arabic  numerals)  for  Crepis,  (In  regions  V  and  XI  solid  circles  indicate  single  mountains 
and  ellipses,  mountain  ranges  or  systems;  solid  squares  indicate  single  lowland  stations. 
See  key  to  symbols  in  text.) 

erly  trend  across  Turkestan  and  Iran.  From  that  region  the  three  branches 
already  mentioned  led  to  northwest  and  tropical  Africa;  to  the  Caucasus, 
Asia  Minor  and  southern  Europe;  and  to  the  eastern  Mediterranean  and 
North  African  littoral.  The  point  to  be  emphasized  here  is  that  the  distribu- 
tion of  the  endemic  regions  for  Crepis  is  entirely  consistent  with  the  generic 
history  briefly  outlined  above. 

III.  PHYLOGENY  AND  ENDEMISM  IN  CREPIS 

Based  on  the  evidence  from  comparative  morphology  and  cytogenetics  it 
has  been  possible  to  classify  all  the  species  of  Crepis  on  a  relative  phylogenetic 
basis  with  some  degree  of  assurance.  For  purposes  of  the  present  discussion 
they  have  been  grouped  into  three  classes,  the  more  primitive,  the  interme- 
diate, and  the  more  advanced  species.  In  Table  2  are  shown  the  relations  of 
the  endemic  species  in  these  phylogenetic  groups  to  the  size  of  area  occupied 


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VoL.XXV]  BABCOCK:  ENDEMISM  IN  CBEPIS  279 

by  each  species  (narrow  or  wider),  to  its  altitude  class  (alpine,  montane  or 
low),  its  moisture  classification  (as  occupying  a  moist,  sub  humid  or  arid  en- 
vironment), its  life-duration  class  (perennial  or  annual-biennial),  and  its 
variability  class  (monomorpliic  or  polymorphic). 

Considering  first  the  totals  in  the  right  hand  column,  it  will  be  noted  that 
twenty-nine  of  the  endemic  species  are  primitive  and  twentj^'-nine  are  ad- 
vanced, whereas  seventy-five,  or  more  than  one-half  of  them,  are  intermediate. 
Evidently  Crepis  may  be  described  as  a  conservative  genus  and  this  becomes 
more  patent  when  it  is  realized  that  of  the  twenty-nine  advanced  endemics 
only  some  half-dozen  desert  or  montane  annuals  can  be  considered  as  very 
highly  specialized  species.  There  is  apparently  no  correlation  between  the 
phj^logenetic  grouping  and  extent  of  distribution  (narrow  vs.  wider),  since 
about  one-third  of  all  the  endemics  are  narrow  and  two  thirds  are  wider  and 
these  same  proportions  hold  roughly  for  each  of  the  three  phylogenetic  classes. 
In  altitudinal  distribution  there  is  positive  correlation  between  primitive  type 
and  alpine  altitude  and  very  strong  positive  correlation  between  advanced 
type  and  low  altitude.  In  the  moisture  classes  we  find  that,  of  the  one  hundred 
and  four  endemics  in  the  primitive  and  intermediate  groups  taken  together, 
just  one-fifth  occur  under  arid  conditions,  M'hereas  four-fifths  of  the  advanced 
endemics  occur  under  arid  conditions.  Hence  there  is  strong  positive  correla- 
tion between  advanced  type  and  arid  environment.  In  the  life-duration  classes 
the  differences  are  most  striking — all  the  primitive  and  intermediate  endemics 
are  perennial  whereas  nearly  all  of  the  advanced  endemics  are  annual  or  (in  a 
few  cases)  biennial.  In  regard  to  variability  there  is  a  stronger  tendency 
towards  polymorphism  in  the  advanced  than  in  the  intermediate  or  primitive 
endemics.  All  of  the  primitive  and  intermediate  endemics  are  perennial  species 
and  more  than  four-fifths  of  them  are  monomorphic  and  occur  at  montane  or 
alpine  elevations  and  under  suhhumid  or  moist  conditions.  Most  of  the  ad- 
vanced endemics  are  annual  species  and  two-fifths  of  them  are  polymorphic; 
none  occur  at  alpine  elevations  and  about  three-fourths  of  them  are  found  at 
low  elevations  and  under  arid  conditions. 

Having  noted  the  lack  of  correlation  between  extent  of  distribution  (as 
expressed  by  the  arbitrary  classes  "narrow"  and  "wider"  (and  the  three 
phylogenetic  groups,  it  may  be  of  interest  to  examine  the  relations  between 
extent  of  distribution  and  the  other  four  classifications  of  endemic  Crepis 
species.  In  Table  3,  in  order  to  reveal  the  relations  involved  more  clearly,  the 
actual  numbers,  given  in  the  upper  two  rows,  are  expressed  in  the  lower  two 
rows  as  percentages  of  the  respective  totals,  given  in  the  right-hand  column. 
The  totals  given  in  the  middle  row  are  merely  for  the  purpose  of  checking 
against  the  other  tables.  Comparing  these  percentages,  it  is  clear  that  there  is 
little  or  no  positive  correlation  between  narrow  and  wider  distribution  and 
phylogenetic  grouping.  But  it  is  equally  clear  that  there  is  positive  correlation 
of  wider  distribution  with  low  altitude  and  arid  climate.  As  for  duration  of 
life  and  variability,  about  four-fifths  of  both  the  narrow  and  the  wider  en- 


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VoL.XXV]  BABCOCE:  EXDEMISM  IN  CEEPIS  281 

demies  are  perennial ;  wliereas  all  of  the  narrow  endemics  are  monomorphic, 
but  one-third  of  the  wider  endemics  are  polymorphic. 

Considering-  these  facts  in  the  light  of  the  conclusions  derived  from  Table  2, 
it  may  be  inferred  that  the  advanced,  annual  endemic  species  which  are  poly- 
morphic are  mostly  of  wider  distribution  and  occur  at  low  altitude,  under  arid 
conditions.  This  inference  has  been  verified  by  reexamination  of  the  original 
data.  There  are  twenty -five  annual  endemic  species  (Table  2)  of  which  twenty- 
three  occur  in  an  arid  climate  and  all  but  one  of  these  at  low  altitude.  Only 
eighteen  are  known  to  have  a  wider  distribution ;  but,  of  the  other  seven,  six 
are  known  as  yet  only  from  the  type  locality  and  further  collections  may  show 
that  some  of  these  also  have  a  wider  distribution. 

The  contrast  between  this  group  of  advanced  endemics  and  the  strictly  al- 
pine endemics  is  most  striking.  Of  the  thirty-one  alpine  endemics,  all  are 
perennial ;  twenty-nine  are  monomorphic  and  only  two,  polymorphic ;  while 
all  but  one  occur  under  subhumid  or  moist  conditions.  Six  of  them  are  primi- 
tive, aiid  twenty -five  intermediate — there  are  no  advanced  alpine  species.  This 
means,  of  course,  that  the  alpine  endemics  are  all  relatively  old  species  and 
adapted  to  mesophytic  conditions;  whereas  the  lowland  endemics  of  advanced 
type  are  relatively  young  species  and  adapted  to  xerophytic  conditions. 

But  the  contrast  between  the  two  gToups  does  not  hold  for  size  of  distribu- 
tional area.  Of  the  thirty-one  alpine  endemics  twenty-one  are  wider  and  only 
ten  narrow  in  distribution.  Furthermore,  there  are  ten  other  alpine  species  of 
Crepis  which  are  so  extensive  in  geographic  distribution  that  they  are  not 
considered  to  be  endemic.  For  example,  C.  conyzaefolia  is  a  primitive  alpine- 
subalpine  species  distributed  from  the  Pyrenees  to  the  Balkan  Peninsula  and 
in  northern  Asia  Minor,  Transcaucasia,  northern  Persia  and  the  Altai  region. 
Such  evidence,  together  with  the  prevalence  of  wide  distribution  among  the 
alpine  endemics,  certainly  indicates  that  the  alpine  endemics  were  formerly 
more  widely  distributed  and  have  become  restricted  to  their  present  locations 
presumably  through  radical  changes  in  the  environment. 

The  evidence  on  chromosome  numbers  is  in  good  agreement  with  the  forego- 
ing generalizations.  In  Table  4  fifty -three  diploid  endemic  species  are  classified 
opposite  their  numbers  6,  5,  4,  or  3,  and  in  relation  to  phylogenetic  grouping- 
and  the  other  classifications  used  in  the  preceding  tables.  Phylogenetically, 
the  6-paired  endemics  are  all  primitive  or  intermediate,  none  are  advanced ; 
whereas  tlie  5-paired  endemics  are  about  evenly  divided  among  the  three 
groups  and  the  4-paired  endemics  are  mostly  intermediate  or  advanced.  The 
one  3-paired  endemic,  C.  fuUginosa,  is  advanced,  widely  distributed,  montane, 
xerophytic,  annual,  and  polymorphic.  In  distribution  all  of  the  6-paired  en- 
demies  are  wider  as  well  as  most  of  the  5's  and  4's ;  but  the  small  number  of 
narrow  endemics  that  have  been  examined  eytologically  may  be  partly  due  to 
difficulties  in  obtaining  them  in  living  condition.  The  data  on  altitudinal  dis- 
tribution indicate  no  correlations  except  in  the  4-paired  species  which  are 
mostly  of  low  elevation.  As  for  moisture  relations,  most  of  the  fifty-three 


282  CALIFORNIA  ACADEMY  OF  SCIENCES  [Pkoc.  4th  ser. 

species  are  about  equally  divided  between  subliumid  and  arid  and  this  holds 
for  each  of  the  chromosome  number  classes.  Under  duration  of  life  we  find 
that  all  of  the  6-paired  endemics  are  perennial,  whereas  about  one-third  of 
both  5's  and  4's  are  annual.  Obviously  reduction  in  length  of  life  has  gone 
along  with  reduction  in  the  number  of  chromosomes.  As  for  variability,  there 
is  little  difference  between  the  chromosome  number  classes  in  the  proportion 
of  monomorphic  and  polymorphic  species.  The  most  significant  facts  derived 
from  this  analysis  are  :  (1)  The  6-paired  species  are  mostly  primitive  and  all 
perennial,  hut  they  are  about  equally  distributed  among  the  altitude  and  mois- 
ture classes.  (2)  The  4-paired  species  are  mostly  intermediate  or  advanced 
and  of  low  elevation,  but  they  are  about  equally  divided  between  mesophytic 
and  xerophytic  environments.  (3)  These  facts  seem  to  indicate  that  adapta- 
tion from  mesophytic  to  xerophytic  cofiditions  has  been  going  on  in  this  genus 
over  a  very  long  period  of  time. 

IV.  THE  DISTRIBUTION  AND  PHYLOGENY  OF 
CREPIS  ENDEMICS 

In  Table  5  each  of  the  endemic  Crepis  species  is  tabulated  according  to  th< 
endemic  region  in  which  it  occurs  and  its  phyletic,  altitude  and  life-duratioi 
class ;  also  the  fifty-three  endemics  that  have  had  their  chromosomes  countec 
are  tabulated  according  to  chromosome  number  as  well  as  the  region  where 
they  occur. 

Considering  first  the  primitive  endemics,  it  will  be  noted  that  two  grades 
are  recognized,  A  and  B.  The  thirteen  species  in  the  A  group  are  definitely 
more  primitive  than  those  of  the  B  group  on  morphological  grounds.  Eleven| 
of  these  most  primitive  endemics  are  either  alpine  or  montane,  and  restricte( 
to  southwestern  Europe  or  the  Balkan  Peninsula  or  to  a  few  high  mountains| 
in  tropical  Africa.  The  other  two  are  alpine  species,  one  in  the  western  Hima- 
laya Mountains  and  the  other  in  the  northern  islands  of  Japan.  Thus  the  most 
primitive  Crepis  endemics  are  now  mostly  distributed  at  great  distances  froi 
the  assumed  center  of  origin  for  the  genus.  Since  most  of  the  advanced  en^ 
demies  are  closer  to  Central  Asia  than  southwestern  Europe,  the  distributionj 
of  the  most  primitive  and  most  advanced  endemics  in  Crepis  conforms  ii 
general  with  Matthew's  principle  of  radial  distribution  of  older  types  from 
common  center  accompanied  by  the  development  of  more  advanced  types 
nearer  the  center  (IMattliew,  1915).  This  conformity  with  Matthew's  principle 
is  also  apparent  from  the  distribution  of  all  the  species  (including  the  en-| 
demies)  in  several  of  the  more  primitive  sections  of  Crepis.  These  facts  provide 
a  sound  basis  for  the  hypothetical  history  of  Crepis  which  starts  with  the 
origin  and  early  development  of  the  genus  in  Central  Asia. 

Considering  next  the  totals  for  the  twelve  regions  (Table  5.  left-hand  coin 
umn),  it  is  clear  that  the  greatest  concentrations  of  endemics  exist  in  thai 
Balkan  Peninsula,  in  Asia  Minor  and  adjacent  areas,  in  tropical  Africa,  and 
in  the  Mediterranean  littoral.  But  these  four  regions  differ  as  to  the  phylo- 


VoL.XXV]  BABCOCK:  ENDEMISM  IN  CBEPIS  283 

genetic  status  of  their  endemics.  In  the  Balkan  Peninsula  close  to  one-half  of 
the  endemics  are  intermediate,  whereas,  one-fourth  are  primitive  (A+B)  and 
one-fourth,  advanced.  These  twenty-six  species  are  similarly  divided  between 
montane,  alpine  and  low  elevations ;  the  seven  advanced  species  are  all  annuals. 
Evidently  the  highly  diversified  conditions  in  that  region  have  favored  the 
persistence  of  both  primitive  and  intermediate  endemics  as  well  as  the  develop- 
ment of  advanced  endemics.  In  Asia  Minor  and  the  Mediterranean  littoral, 
however,  the  endemics  are  all  either  intermediate  or  advanced.  But  the  fact 
that  there  are  no  primitive  endemics  in  Asia  Minor  is  probably  due  to  the  gen- 
erally more  arid  climate  of  that  region  as  compared  with  the  Balkan  Peninsula 
as  a  whole.  In  tropical  Africa,  on  the  other  hand,  we  find  just  the  opposite 
situation.  The  eighteen  endemics  are  all  either  intermediate  or  primitive ;  and 
this  is  also  true  of  four  other  tropical  African  species  which  are  so  widely  dis- 
tributed that  they  are  not  included  here  as  endemics.  This  absence  of  advanced 
species  in  tropical  Africa  probably  indicates  that  the  more  uniform  and  favor- 
able conditions  of  that  region  have  favored  the  persistence  of  more  primitive 
types  without  inducing  the  development  of  very  advanced  types. 

Compared  with  the  four  regions  that  are  richest  in  endemics,  the  following 
regions  have  only  one-third  to  one-half  as  many  in  each  region  :  southwestern 
Europe,  the  Caucasus-Iran  region,  Abyssinia  and  adjacent  small  areas,  and 
southeastern  Asia.  One  of  these,  the  Abyssinian  region,  differs  from  the  other 
four  in  having  all  the  phyletic  groups  represented.  The  t^n  endemics  of  the 
Abyssinian  region  are  all  montane  except  the  two  primitive  species  which  are 
alpine ;  and  yet  four  of  these  montane  species  are  advanced.  It  will  be  recalled, 
however,  that  certain  other  advanced  species  are  montane.  This  is  true  of  the 
one  3-paired  endemic,  C.  ftiliginosa,  which  occurs  in  southern  Greece,  and  of 
its  4-paired  close  relative,  C.  cretica.  The  three  other  regions  contain  only 
perennial  endemics,  none  of  w^hich  is  advanced,  some  being  intermediate  and 
some  primitive.  Southwestern  Europe  and  tropical  Africa  are  important 
regions  for  the  most  primitive  endemics  since  nine  of  the  twenty-six  endemics 
found  in  those  two  regions  are  classed  as  primitive-A.  Thus,  at  the  western 
and  southwestern  extremes  of  distribution  for  Crepis,  we  find  the  greatest 
concentration  of  the  most  primitive  endemics  in  the  genus. 

In  order  to  give  another  picture  of  the  morphological  contrasts  between 
these  most  primitive  endemics  and  the  most  advanced  endemics  in  the  genus, 
C.  tergloucnsis,  an  alpine  species  found  only  in  the  European  Alps,  is  shown 
in  fig.  4,  a-f,  and  C.  ftiliginosa,  a  lower  montane  species  of  southern  Greece, 
in  fig.  4,  a'-f '.  The  plants  and  their  parts  are  all  drawn  at  comparable  scales. 
It  will  be  noted  that,  in  their  haploid  sets  of  chromosomes,  the  two  species  rep- 
resent the  extremes  of  variation  found  in  Crepis  in  both  number  and  size  of 
the  chromosomes. 

With  further  reference  to  the  chromosomes  of  fifty-three  of  the  endemic 
species  (Table  5) ,  it  is  clear  that  the  6-paired  endemics  are  restricted  to  regions 
I  to  IV  except  for  one  species,  C.  hashmirica,  of  northwestern  India.  In  fact 


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Vol.  XXV] 


BABCOCK:  EXDEMISM  IX  CEEPIS 


285 


rffT{\ 


0)7Ur 


Fig.  4.  a-/,  Crepis  ierglouensis  (Hacq.)  A.  Kern,  a,  plant,  x  %  ;  l»,  floret  lacking  the 
ovary,  x3;  c,  anther-tube  slit  and  opened  out,  x6;  cZ,  detail  of  anther-appendages,  x24; 
e,  achene,  x  6;  /,  haploid  set  of  chromosomes,  x  925.  a'-f,  Cre-pis  fuliginosa  S.  et  S.  all  as 
above. 


two-thirds  of  them  occur  in  the  mountains  of  southwestern  Europe  and  the 
Balkan  Peninsula.  But  the  5-paired  endemics  are  widely  distributed,  and  the 
4-paired  species  still  more  widely,  with  a  strong  representation  in  the  Medi- 
terranean littoral.  The  evidence  from  chromosome  numbers  is  consistent  with 
the  morphological  evidence  in  showing  that  the  oldest  endemics  are  mostly  in 
the  mountains  of  southern  Europe  whereas  the  j^oungest  endemics  are  mostlj^ 
in  the  Mediterranean  littoral  and  closely  adjacent  areas.  This  evidence  from 


286  CiLIFOHXIA  ACADEMY  OF  SCIEXCES  [Proc.  4th  ser. 

chromosome  numbers  supports  the  general  hypothesis,  based  on  morphology 
and  geographic  distribution  in  the  genus  as  a  whole,  concerning  the  origin 
and  distribution  of  the  genus.  This  hypothesis  includes  the  assumption  that 
two  of  the  important  causes  of  increased  speciation  in  the  genus  were  the 
climatic  and  topographic  changes  which  occurred  in  southwestern  Asia  and 
the  Mediterranean  region  during  late  Tertiary  and  early  Quarternary  time. 
In  Crepis  the  endemic  areas  most  affected  by  tliese  changing  conditions  are 
lto9,  21,22and24(cf.fig.  3). 

V.  THE  GENERAL  SIGNIFICANCE  OF  ENDEMISM  IN 

CREPIS 

In  addition  to  the  support  found  in  this  study  of  endemism  in  Crepis  for 
the  hypothetical  history  of  the  genus  which  had  been  previously  formulated, 
it  appears  that  some  of  the  evidence  on  Crepis  endemics  has  a  bearing  on  im- 
portant problems  of  plant  distribution.  One  such  problem  concerns  the  so- 
called  age  and  area  hypothesis  (Willis,  1940)  which  assumes  that  the  age  of 
a  species  or  group  is  proportional  to  the  area  it  occupies. 

The  "age  and  area"  concept.  From  the  foregoing  review  it  is  clear  that 
among  the  endemic  species  of  Crepis  there  are  two  extremely  diverse  groups  : 
(1)  ihe  primitive,  perennial,  alpine  endemics  which  are  relatively  old,  mono- 
morphic,  and  adapted  to  mesophytic  conditions ;  and  (2)  the  advanced,  annual 
lowland  endemics  M^hich  are  relatively  young,  more  highly  variable,  and 
adapted  to  xerophytic  conditions. 

Now  the  present  distribution  of  these  two  groups  does  not  conform  to  the 
"age  and  area"  concept.  The  old,  alpine  endemics,  however,  are  believed  by 
most  students  of  floristics  to  have  been  more  widespread  before  the  Pleistocene 
epoch  and  to  have  moved  into  the  narrow  niches  that  they  now  occupy  from 
glacial  refugia  (Braun-Blanquet,  1923,  pp.  260-261).  The  fact  that  many  of 
them  are  of  relatively  wide  distribution,  although  they  exist  only  in  a  special 
type  of  environment,  supports  this  idea.  On  the  other  hand,  the  young,  low- 
land, annual  endemics  are  just  as  widely  distributed  as  the  alpine  species  are 
at  present;  and  there  are  a  number  of  lowland,  annual  species,  not  endemics, 
for  example,  C.  joetida  and  C.  pulchra,  which  are  about  as  widely  distributed 
as  the  forerunners  of  the  alpine  endemics  ever  could  have  been.  Hence  the 
"age  and  area"  concept  breaks  down  completelj^  with  respect  to  these  tM'o 
groups  of  endemic  species.  As  has  been  pointed  out  by  Stebbins  (1942)  the 
concepts  of  "senescence"  and  "competition"  are  no  more  satisfactory  than 
"age  and  area"  in  attempting  to  account  for  such  diverse  groups  of  species. 
But  the  concept  that  the  monomorphic,  alpine  species  are  genetically  homo- 
geneous whereas  the  variable  lowland  species  are  genetically  heterogeneous, 
i.e.,  composed  of  several  or  many  biotypes,  provides  an  acceptable  explanation 
of  their  differences  in  variability  and  distribution. 

The  nature  of  the  "narrow"  Crepis  endemics.  This  genetic  approach  to  the 
problem  of  the  differences  between  the  widely  distributed  alpine  and  low- 


YoL.XXY]  BABCOCK:  ENDEMISM  IX  CEEPIS  287 

laud  endemics  also  provides  a  satisfactory  explanation  of  the  nature  of  the 
"narrow"  Crepis  endemics.  Of  the  forty-eig'ht  Old  World  species  which  were 
classified  as  "narrow"  many  are  known  only  from  their  type  locality  or  else 
the  information  about  their  distribution  is  vague ;  but  there  are  several  con- 
cerning which  it  is  possible  to  make  fairly  definite  statements.  These  are  listed 
below  in  their  phylogenetic  groups,  together  with  pertinent  data. 

Narrow  Endemics — All  Presumably  Monomorphic 

Primitive 

1.  C.  Mlimandscliarica  O.  Hoffm.  Section  8.  Known  only  from  Mt.  Kilimanjaro  at  four 
stations,  2700-3500  meters,  in  forest. 

2.  C.  cameroonica  Babe.  Section  8.  Known  with  certainty  only  from  Mt.  Cameroon  at  three 
stations,  2000-4000  meters,  in  meadows. 

Intermediate 

3.  C.  taygetica  Babe.  Section  10.  Known  from  five  collections,  all  made  at  a  place  called 
"Porta"  at  about  2300  meters  altitude  on  Mt.  St.  Elias  in  the  Tavgetus  Mts.,  of  southern 
Greece.  Exposed  rocks. 

4.  C.  crocifolia  Boiss.  et  Heldr.  Section  10.  Knol^^l  from  three  or  four  collections  made  at  a 
place  called  Megala  Zonaria,  at  about  2400  meters  altitude,  on  the  same  mountain  as 
the  preceding.  Exposed  rocks. 

5.  C.  athoa  Boiss.  Section  10.  Seven  collections,  all  on  Mt.  Athos,  Hagion  Oros  Peninsula, 
northern  Greece.  Exposed  rocks. 

Advanced 

6.  C.  eritreensis  Babe.  Section  20.  Three  or  four  localities  in  southern  Eritrea  from  800 
to  1400  meters  elevation.  One  collection  was  from  "a  comparatively  moist,  rocky  for- 
mation." 

7.  C.  tyhalciensis  Vierh.  Section  20.  Known  from  one  specimen,  collected  in  southern  Crete. 
Since  Crete  has  been  botanized  by  numerous  collectors,  it  is  very  probable  that  this 
species  would  be  represented  by  more  collections  if  it  were  widely  distributed  on  the 
island.  Certainly  insular. 

8.  C.  divaricata  (Lowe)  F.  Schultz.  Section  25.  Known  with  certainty  from  four  collections 
made  on  the  isolated  eastern  promontory  of  Madeira  Island  where  it  has  been  nearly 
exterminated  by  grazing.  (Eeported  in  1837-1850  from  the  tiny  Desertas  Islands  near 
Madeira.)  Certainly  insular. 

9.  C.  Noronhaea  Babe.  Section  25.  Six  collections,  all  from  Porto  Santo,  a  small  island  in 
the  Madeira  Archipelago.  Certainly  insular. 

10.  C.  Forskalii  Babe.  Section  27.  Two  localities  in  Yemen  Province,  southwestern  Arabia, 
between  1300  and  2500  meters  elevation,  in  the  coffee  belt. 

It  is  noteworthy  that  three  of  these  species,  C.  divaricata,  C.  Noronhaea  and 
C.  tyhakiensis,  are  insular  and  of  very  restricted  distribution.  However,  they 
are  relatively  advanced  species.  Therefore  their  morphological  homogeneity 
(assuming  that  C.  tyhakiensis  is  actually  monomorphic)  must  be  due  to  genetic 
homogeneity  rather  than  to  senescence.  Five  others  in  the  above  list,  C.  kili- 
mandscJiarica,  C.  cameroonica,  C.  tay'getica,  C.  crocifolia  and  C.  athoa,  are  each 
found  on  only  a  single  mountain.  They  are  almost  certainly  "depleted"  species 
(Stebbins,  1942)   and  they  probably  became  isolated  through  widespread 


288  CALIFORNIA  ACADEMY  OF  SCIENCES  [Proc. -Ith  ser. 

changes  in  topography  and  climate  during  late  Tertiary  and  Pleistocene  times. 
In  these  also  the  population  must  be  relatively  small  and  homogeneous.  The 
other  two  species  may  also  be  "depleted"  species,  although  it  must  be  admitted 
that  the  available  information  on  their  distribution  and  variability  is  scanty. 

SUMMARY 

1.  The  twelve  endemic  regions  for  Crepis,  comprising  twenty-five  areas, 
define  the  east -west  distribution  of  the  genus  almost  completely. 

2.  Seventy -two  per  cent  of  the  Old  World  Crepis  species  are  endemic  in  the , 
sense  that  thej^  are  restricted  to  a  single  subcontinental  geographic  area. 
Among  these  endemic  species  are  two  very  diverse  groups,  namely  the  most 
primitive,  perennial  species,  half  of  which  are  alpine  relics,  and  the  advanced,! 
annual  species,  most  of  which  occur  at  low  altitudes  under  arid  conditions. 
The  remaining  endemics  comprise  an  intermediate  series  connecting  these  tw( 
extremes.  But  comparatively  few  of  the  alpine  perennials  and  of  the  lowlanc 
annuals  have  narrow  distributions,  i.e.,  their  distributional  areas,  as  at  present 
known,  are  mostly  over  150  kilometers  in  extent.  From  this  evidence  it  is  in-| 
f erred  that  the  alpine  relics  were  formerly  of  wider  distribution  and  that  thej 
moved  into  their  present  niches  from  glacial  refugia ;  whereas  the  lowland, 
annual  endemics  have  become  adapted  to  a  more  arid  climate  but  under  con- 
ditions favoring  genetic  heterogeneity.  From  the  evidence  on  morphology  and 
distribution  as  well  as  on  chromosome  numbers  it  is  inferred  that  the  process 
of  adaptation  from  mesophytic  to  xerophytic  conditions  went  on  in  the  genus 
as  a  whole  over  a  long  period  of  time. 

3.  The  distribution  of  the  most  primitive  and  most  advanced  endemic  species 
of  Crepis  supports  the  conclusion,  derived  from  the  evidence  on  phylogeny 
and  distribution  of  the  genus  as  a  whole,  that  the  center  of  origin  was  in  Cen- 
tral Asia. 

4.  The  "age  and  area"  concept  of  Willis  breaks  down  completelj^  in  the  light 
of  the  evidence  on  the  two  diverse  groups  of  endemic  species  mentioned  above. 

5.  The  concept  of  "senescence"  fails  to  explain  the  existence  of  narrow  en- 
demics in  Crepis  since  this  group  includes  some  of  the  most  primitive  and 
most  advanced  species  in  the  genus.  Since  they  are  either  insular  or  terrestrial, 
"depleted"  species,  the  population  is  more  or  less  restricted  in  size  and  rela- 
tively homogeneous. 


VoL.XXV]  BABCOCE:  ENDEMISM  IN  CBEPIS  289 

-    REFERENCES 
Babcock,  E.  B. 

1942.  Systematies,  cytogenetics  and  evolution  in  Crepis.  Bot.  Eev.,  8:139-190. 

Babcock,  E.  B.,  and  G.  L.  Stebbins,  Jr. 

1938.  The  American  Species  of  Crepis.  Carnegie  Inst.  Wash.,  Pub.  No.  504:1-199. 

Babcock,  E.  B.,  and  J.  A.  Jenkins. 

1943.  Chromosomes  and  phylogeny  in  Crepis  III.  The  relations  of  one  hundred  and  thir- 
teen species.  Univ.  Calif.  Publ.  Bot.,  18:241-292. 

Braun-Blanquet,  J. 

1923.  tyber  die  Genesis  der  Alpenflora.  Verh.  Naturf.  Ges.  Basel,  35:243-261. 

Matthew,  W.  D. 

1915.  Climate  and  Evolution.  Ann.  N.  Y.  Acad.  Sci.,  24:171-318. 

Stebbins,  G.  L.,  Jr. 

1942.  The  genetic  approach  to  problems  of  rare  and  endemic  species.  Madrono,  6:241-258. 

TOBGY,  H.  A. 

1943.  A  cytological  study  of  Crepis  fuliginosa,  C.  neglecta,  and  their  Fj  hybrid,  and  its 
bearing  on  the  mechanism  of  phylogenetic  reduction  in  chromosome  number.  Jour. 
Genetics,  45 : 6  7-1 11. 

Willis,  J.  C. 

1940.  The  Course  of  Evolution.  207  pp.  Cambridge  University  Press. 


PROCEEDINGS 
OF  THE 

CALIFORNIA  ACADEMY  OF  SCIENCES 

Fourth  Series 


;««r^  t 


Vol.  XXV,  No.  12,  pp.  291-306,  pis.  26-27 


November  10,  1944 


ALICE  EASTWOOD 
SEMI-CENTENNIAL  PUBLICATIONS 

No.  12 

NICOTIANA  ARENTSII— A  NEW,  NATURALLY 
OCCURRING  AMPHIDIPLOID,  SPECIES 

BY 

T.  H.  GOODSPEED 

Department  of  Botany 
University  of  California* 


IN  1917,  Winge  pointed  out  that  the  arithmetic  progression  observed  in  the 
chromosome  numbers  of  some  genera  might  be  the  product  of  interspecific 
hybridization  followed  by  chromosome  doubling.  Eight  years  later,  Clausen 
and  Goodspeed  showed  that  such  doubling  had  occurred  in  a  fertile  F^  hybrid 
between  Nicotiana  glutinosa  and  N.  Tdbacum.  This  initial  verification  of 
Winge's  hypothesis  served  to  direct  interest  toward  the  incidence,  causes  and 
artificial  induction  of  amphidiploidy  (cf.  Goodspeed  and  Bradley,  1942).  It 
also  led  to  an  evaluation  of  the  applicability  of  Winge's  hypothesis  in  pictur- 
ing evolutionary  relationships  in  the  genus  Nicotiana. 

Fundamentally,  Nicotiana  is  a  New  World  genus  and,  with  the  exception  of 
5  species  showing  9  or  10  pairs  of  chromosomes,  all  its  North  and  South  Ameri- 
can species  possess  either  12  or  24  pairs.  Since  there  is  no  evidence  that  any  of 
the  24-paired  species  is  autopolyploid,  it  has  been  assumed  that  they  were 
amphidiploid  in  origin  (Goodspeed,  1934).  Proof  of  such  origin  would  con- 
tribute materially  to  the  systematic  botany  of  the  genus  and  to  an  interpre- 
tation of  its  species  distribution  in  the  two  Americas.  For  two  of  the  24-paired 
species,  such  proof  is  at  hand.  Thus,  iV.  Tahacuni  (n  =  24)  has  been  shown  to  be 

*  Contribution  No.  112  from  the  University  of  California  Botanical  Garden. 

Acknowledgment  is  made  of  grants  received  from  the  Committee  for  Inter-American 
Artistic  and  Cultural  Eelations,  the  American  Philosophical  Society,  the  American  Academy 
of  Arts  and  Sciences,  the  Coolidge  Foundation,  and  to  a  number  of  private  donors,  in  sup- 
port of  the  studies  reported  upon.  The  author  is  indebted  to  Dr.  H-M.  Wheeler  and  M.  C. 
Thompson  for  assistance  in  the  preparation  of  this  article. 

[291] 


292  CALIFOBNIA  ACADEMY  OF  SCIENCES  [Proc.  4th  ser. 

au  amphidiploid  product  of  hybridization  between  progenitors  of  the  modern 
species  N.  sylvesiris  (n  =  12)  and  N.  tomentosa  (n  =  12)  or  its  relatives  (Good- 
speed  and  Clausen,  1928) .  These  two  species  are  peculiar  to  the  Andes  of  Peru, 
Bolivia,  or  Argentina,  and  one  of  the  products  of  the  three  University  of  Cali- 
fornia Botanical  Garden  Expeditions  to  the  Andes  has  been  the  demonstration 
that  today  the  distribution  of  the  putative  parental  species  overlaps.  Simi- 
larly, progenitors  of  two  other  present  day  Peruvian-Bolivian-Argentine 
species,  N.  paniculata  (n=12)  and  N.  undnlata  (n=12) ,  were  undoubtedly  the 
parents  of  the  amphidiploid  species  N.  rustica,  the  other  well-known  24-paired 
species  of  commerce  (Goodspeed,  1934,  1936). 

Despite  cytogenetic  evidence  in  favor  of  amphidiploid  origin,  it  has  not  been 
possible  to  reproduce  artificially  a  duplicate  of  modern  N.  T  aha  cum,  or  of 
N.  rustica,  nor  is  there  evidence  of  tlie  existence  of  at  least  the  former  species 
in  a  wild  state.  In  other  words,  although  there  is  good  reason  to  believe  that 
the  24-paired  American  species  of  Nicotiana  are  of  amphidiploid  origin  and 
while  amphidiploidy  has  been  induced  in  at  least  27  experimentally  produced 
Fj  intersiT^cific  hybrids  (Bradley  and  Goodspeed,  1943),  no  naturally  oc- 
curring American  species  has  hitherto  been  found  for  which  such  origin  could 
be  shown.^  It  now  appears  that  the  recently  discovered  N.  Arentsii  is  such  a 
species. 

In  November  1938  one  of  the  members  of  the  Second  University  of  Cali- 
fornia Botanical  Garden  Expedition  to  the  Andes,  Dr.  Cesar  Vargas  C,  Pro- 
fessor of  Botany  and  Director  of  the  Botanical  Museum  in  the  University  of 
Cuzco,  Peru,  found  N.  Arentsii  in  southeastern  Peru,  north  of  Lake  Titicaca, 
near  the  Bolivian  border,  in  and  about  the  village  of  Limbani.  Limbani  is 
situated  in  the  Dept.  Puno,  Prov.  Sandia,  at  an  altitude  of  about  3500  m., 
in  a  narrow  box  canyon  on  the  eastern  watershed  of  the  Cordillera.  The  region 
is  remote,  climatically  inhospitable,  and  little  known  botanically.  Nearby, 
at  somewhat  lower  altitudes,  is  the  beginning  of  the  Amazonian  rain  forest. 
The  following-  extract  from  Dr.  Vargas'  notes  is  included  here,  in  part  because 
of  its  general  botanical  interest.  Below  Limbani,  "the  river  of  the  same  name 
runs  swiftly  and  precipitately,  a  turbulent  and  foam-flecked  stream  which 
springs  from  the  thawing  snows  of  the  nearby  Cordillera.  The  village  mostly 
consists  of  hovels  and  cabins  nearly  all  made  of  wliitish  granitic  rock  with  roofs 
of  straw.  The  only  street  is  the  tortuous  highway,  lined  on  both  sides  by  the 
principal  habitations  while  the  rest  are  scattered  here  and  there  on  contiguous 
slopes.  On  the  walls  of  the  canyon  are  a  series  of  shelves  or  terraces,  now  in 
ruins.  Nobody  knows  how  many  hundreds  of  years  ago  they  were  constructed 
through  the  laboriousness  and  perseverance  of  a  people  who  disappeared  long 

^  Kostoff  (1939)  has  commented  upon  the  origin  of  a  32-paired  race  of  the  Australian 
species  N.  suavcolms  (n  =  16)  or  its  close  relatives  whieh  was  first  grown  in  the  University 
of  California  Botanical  Garden  (Wheeler,  1935).  For  tliis  race  Kostoff  has  suggested  the 
name  N.  Eastii  (but  does  not  formally  describe  it  as  a  new  species)  and  considers  it  an 
amphidiploid  product  of  hybridization  between  N.  suaveolens  and  N.  maritima.  We  have 
grown  more  than  one  32-paired  race  from  Australia  and  their  morphological  distinctions 
appear  to  complicate  the  problem  somewhat. 


VOL.XXV]         GOODSPEED  :  NICOTIANA  ARENTSII  SP.  NOV.  293 

ago  and  for  wliom  the  soil  and  the  plant  constituted  the  most  cherished  gifts 
of  nature.  . .  .  However,  what  is  most  cheering  ...  is  the  abundance  of  vegeta- 
tion .  .  .  herbaceous  and  shrub  forms  as  far  as  my  eyes  reach ;  even  the  rocky 
cliffs  offer  a  varied  flora.  Along  the  road  there  are  found  several  species  of 
the  families  Melastomaceae,  Ericaceae,  Canipanulaceae,  etc.  .  .  .  beautiful  and 
graceful  clusters  of  Bomarea  hang  among  Baccharis,  Cantua,  and  other 
bushes ;  under  the  rocks  and  on  the  demolished  terraces  grow  two  species  of 
Begonia,  Solanum,  Centropogon,  Monnina  and,  in  the  dank  turf,  species  of 
Viola  and  Calceolaria.  The  air  was  humid,  the  soil  moist."  The  Quechua  name 
for  the  new  Nicotiana  species  was  "Cjamasairi." 

Four  years  later,  Mr.  Roy  D.  Metcalf,  a  member  of  the  Third  University  of 
California  Botanical  Garden  Expedition  to  the  Andes,  during  an  exploration 
of  the  Inambari  and  Tambopata  drainage  basins,  collected  N.  Arentsii  again. 
He  found  it  restricted  to  the  immediate  vicinity  of  the  village  of  Limbani  and 
commonly  around  habitations,  in  some  cases  growing  on  walls  and  fences. 
According  to  his  report,  it  prefers  semi-shade,  moist  but  not  wet  surroundings 
and  gravelly  soil  among  rocks.  The  preference  of  N.  Arentsii  for  disturbed 
ground  and  the  partial  protection  provided  by  habitations  is  characteristic 
of  many  South  American  species  of  Nicotiana.  In  Limbani  the  Indians  did 
not  make  any  use  of  the  plant.  Although  Mr.  Metcalf 's  itinerary  northward 
from  Chucuito  (Dept.  and  Prov.  Puno)  at  the  northwest  end  of  Lake  Titicaca, 
took  him  as  far  as  the  Santo  Domingo  mine  (immediately  north  of  the  Inam- 
bari river)  in  the  province  of  Sandia  and  as  far  east  as  the  towns  of  Sandia 
and  Cuyocuyo,  he  did  not  again  encounter  this  species.  He  states  that  N.  un- 
dulata  was  also  called  Cjamasairi  by  the  natives. 

From  the  original  seed  of  N.  Arentsii  collected  by  Dr.  Vargas,  100  plants 
have  been  grown  in  the  University  of  California  Botanical  Garden  at  Berke- 
ley, California.  Of  these,  25  were  grown  in  1940,  75  in  1941.  In  the  former 
year,  all  plants  were  set  out  in  open  ground;  in  the  latter  year,  35  were  planted 
out,  38  held  outdoors  in  large  pots,  and  two  allowed  to  flower  in  pots  in  the 
greenhouse.  Potted  plants  from  seed  sown  on  January  10,  1941,  began  to 
bloom  August  28,  1941.  Those  plants  transplanted  to  open  ground  from  seed 
sowai  on  February  16,  1940,  and  January  10,  1941,  began  to  bloom  in  the 
middle  of  the  next  October.  To  test  their  reported  perennial  capacities  the 
field  culture  was  not  discarded  at  the  close  of  the  1940  growing  season.  Al- 
though the  winter  (of  1940)  was  moderate  in  temperature,  the  plants  died 
after  setting  seed.  All  plants  held  in  large  pots  failed  to  develop  lower 
branches.  Their  inflorescences  were  narrower  than  those  of  field  plants. 
Finally,  plants  blooming  in  the  greenhouse  lacked  the  rose-red  pigmentation 
in  calyx  and  corolla,  characteristic  of  field  grown  plants. 

In  all  these  garden  cultures  the  branching  was  variable  and  frequently 
asymetric.  There  was  also  a  strong  tendency  toward  fasciation  of  elements  of 
the  inflorescence.  Generally,  the  union  did  not  extend  beyond  the  first  leaf  of 
the  axillary  shoot.  The  branch  thus  affected  would  depart  from  the  main  axis 


294  CALIFORNIA  ACADEMY  OF  SCIENCES  [Pkoc.  4th  ser.      i 

at  a  decidedly  more  acute  angle  than  that  of  a  normal  shoot.  Faseiation  did 
not  appear  in  a  population  of  13  plants  grown,  under  equivalent  conditions, 
in  1943  from  selfed  seed  of  a  1942  plant.  Despite  certain  definite  but  inter- 
grading  variations  in  leaf  shape  and  inflorescence  type,  the  appearance  of  the 
original  populations  was  as  uniform  as  might  be  expected  from  seed  of  wild 
plants. 

On  the  basis  of  herbarium  and  garden  studies,  N.  Arcntsir  was  found 
clearly  to  combine  prominent  morphological  features  of  two  other  species  of 
Nicotiana — i.e.,  N.  wigandioides  Koch  and  Fint"  and  N.  undulata  R.  and  P." 
The  former  is  a  subarborescent  species,  less  massive  than  N.  tomentosa  but 
attaining  a  height  of  several  meters  and  capable  of  living  a  number  of  years 
in  the  Berkeley  climate.  Especiall}^  on  younger  stem  parts,  long  straight 
silvery  hairs  stand  out  prominently ;  there  being,  however,  a  glabrous  region 
below  the  insertion  of  the  leaf.  The  length,  and  to  some  degree  the  direction, 
of  these  soft,  glistening  hairs  is  modified  in  N.  Arentsii.  The  red-purple  colora- 
tion of  N.  Arentsii  manifested  in  stem,  leaf,  calyx,  and  a  pinkish  blush  to  the 
corolla  is  derived  from  N.  wigandioides.  Plants  of  both  species  blossoming  in 
the  greenhouse  instead  of  outdoors,  lack  it.  Slow  maturing,  despite  early 
sowing,  is  characteristic  of  both,  N.  undulata  on  the  other  hand,  contributes 
to  N.  Arentsii  relativelj^  low  stature  and  annual  to  limited  perennial  habit, 
greater  leafiness  in  the  inflorescence  region,  and  a  large  adaxial  calyx  lobe.  In 
certain  plants  it  modifies  the  calyx  coloration  to  a  blue-black. 

Generally  speaking,  N.  wigandioides  is  a  tropical  or  subtropical  species  as 
yet  known  only  from  central  (western)  Bolivia.  It  may  be  associated  with  the 
eastern  headwaters  of  the  Beni  river  for  it  has  been  collected  a  short  distance 
beyond  the  edge  of  this  drainage  system  (Cardenas  2800,  Inca  Corral,  Dept. 
Cochabamba,  Prov.  Chapare,  1700  m. ;  Steinhach  5754,  Cerro  de  Ineachaca, 
Dept.  Cochabamba,  Prov.  Sacaba,*  2500  m.)  and  well  within  it  in  valleys  in 
Dept.  La  Paz  (Rushy  75a,  Mulford  Biol.  Explor.  Amazon  Basin,  Caiiamina, 
Prov.  Inquisivi,  1100  m. ;  Troll  2812,  Lambate,  Prov.  Sud  Yungas,  3400  m.; 
Buchtien  3905,  Unduavi,  Prov.  Nor  Yungas,  3200  m. ;  Bushy  2434,  near 
Yungas,  1200  m.).  It  may  also  occur  to  the  north  and  it  is  unfortunate  that 
we  have  been  unable  to  explore  the  tributaries  of  the  Rio  Beni  in  Bolivian 
territory  adjacent  to  the  Peruvian  area  where  N.  Arentsii  was  found. 

Speaking  of  the  vegetation  at  his  last  cited  locality,  H.  H.  Rusby  (1888,  p. 
181)  writes:  ''Descending  to  7000  feet,  we  enter  the  great  Andean  forests 
which  become  heavier  and  heavier,  though  scarcely  denser,  as  we  descend.  The 
trunks  and  greater  branches  are  scarcely  to  be  seen  for  the  epiphytes  upon 
them,  chief  of  which  are  orchids,  bromeliads,  ferns,  mosses,  and  aroids.  At 
5500  feet,  we  strike  the  coca  and  cinchona  belt,  and  at  4000  feet  we  find  the 
heat  becoming  oppressive  and  the  air  sultry.  From  3500  to  5500  feet  is  prob- 

-  A  detailed  description  of  this  new  species  appears  at  the  end  of  this  report  where  photo- 
graphs of  it  and  of  N.  undulata  and  N.  wigandioides  ■will  be  found. 

^  Undoubtedly  a  mistake  and  should  read  "the  canton  of  Saeaba,  in  Prov.  Chapare." 


Vol.  XXV]         GOODSPEED  :  NICOTIANA  ABENTSII  SP.  NOV.  295 

ably  the  region  of  greatest  rain-fall."  Beyond  general  pictures  such  as  this 
there  is  little  description  of  habitat  for  N.  wigandioides  except  the  remarks 
of  Dr.  Martin  Cardenas  on  his  collection  of  this  species  from  Inca  Corral.  He 
refers  to  the  region  as  "Ceja  de  Monte ;  the  'Ceja'  is  a  kind  of  beginning  of  the 
Yungas."  The  term  Yungas  is  used  here  as  a  collective  designation  for  Bolivian 
tropical  valley  terrain.  He  states  that  N.  ivigandioides  grows  "on  wet  soil." 

N.  undidata  is  a  widely  distributed  species  which,  by  contrast,  thrives  in 
high,  cold,  bleak  situations  and  generally  fails  to  compete  successfully  as  the 
climate  becomes  less  rigorous  and  the  region  can  support  a  richer  vegetation. 
Apparently,  it  does  not  descend  the  eastern  flank  of  the  Andes  but  every- 
where finds  a  congenial  habitat  in  the  Puna  zone  to  the  west.  Its  range  is 
continuous  from  northern  Argentina  (Univ.  Calif.  Bot.  Card.  Exped.  Andes 
6323,  coll.  West,  20  km.  w.  of  Humahuaca,  Prov.  Jujuy,  3600  m.)  to  central 
Peru  (Univ.  Calif.  Second  Bot.  Card.  Exped.  Andes  10985,  coll.  Stork, 
Muilo.  10  km.  w.  of  Tarma,  Dept.  Junin,  3300  m.)  and  it  has  even  been 
found  in  northern  Peru  (Univ.  Calif.  Third  Bot.  Card.  Exped.  Andes 
30781,  coll.  Metcalf,  road  from  Cajamarca,  3200  m.).  Near  Oroya,  Peru,  it 
forms  almost  pure  colonies  where  only  it  and  Bidens  can  tolerate  the  fumes 
from  nearby  smelters  (Univ.  Calif.  Second  Bot.  Card.  Exped.  Andes  10986, 
coll.  Stork) ;  between  Candarave  and  Puno  it  occurs  in  the  hard-packed  road- 
bed (Univ  Calif.  Third  Bot.  Card.  Exped.  Andes  30387,  coH.  Metcalf)  ;  in 
Chucuito,  Huancane,  Putina,  Asillo  and  other  windswept  altiplano  towns  of 
the  Dept.  Puno  (ibid.  30677,  30704,  30718,  30424)  it  is  common  in  the  streets, 
on  refuse  dumps,  and  in  the  protection  of  houses  and  fences.  Metcalf  reports : 
"While  crossing  the  altiplano,  the  one  plant  that  one  expects  to  find  is  this 
Nicotiana.  "When  there  is  nothing  else,  there  is  always  this  . .  .  while  never 
growing  in  great  abundance  in  any  one  site,  it  none  the  less  covers  sparsely 
huge  areas.  Koughly,  N.  undulata  occurs  from  10,000  feet  to  over  14,000  feet." 

Although  N.  wigandioides  has  not  been  found  near  the  limited  known  range 
of  distribution  of  N.  Arentsii,  the  other  species,  N.  undulata,  which  appears 
to  have  entered  into  its  amphidiploid  origin,  occurs  within  30  km.  of  it.  In  the 
La  Paz  region  of  Bolivia,  N.  undulata  and  N.  wigandioides  have  been  col- 
lected within  this  distance  and  in  Dept.  Cochabamba  they  are  found  within 
10  km.  of  each  other.  It  is,  therefore,  not  too  much  to  say  that  in  past  time 
these  two  species  have  been  in  contact,  if  they  are  not  so  at  present.  In  the 
dissected  eastern  edge  of  the  Peruvian-Bolivian  Andes,  alps  and  tropics  are 
remarkably  juxtaposed  and  wherever  a  cleared  pathway  opens,  whether  it  be 
by  diastrophic  movement,  streams,  or  man-made  paths,  Nicotiana  species  will 
seek  to  colonize  the  disturbed  ground  to  the  limit  of  their  climatic  tolerance. 

Above,  it  has  been  pointed  out  that  N.  Arentsii  is  morphologically  a  mosaic 
of  the  distinctive  characters  peculiar  to  two  other  species  of  Nicotiana.  Atten- 
tion has  also  been  called  to  the  fact  that,  at  one  time,  the  range  of  distribution 
of  these  two  species  undoubtedly  overlapped  and  that  today  one  of  them  is 
known  to  be  practically  in  contact  with  N.  Arentsii.  These  two  lines  of  evi- 


296  CALIFOSXIA  ACADEMY  OF  SCIENCES  [Proc.  4th  ser. 

dence  suggest  that  this  new  species  originated  as  a  hybrid  between  progenitors 
of  the  two  other  species,  chromosome  doubling  then  occurring  in  the  hj^brid. 
The  following  cytogenetic  observations  largely  confirm  this  suggestion. 

It  is,  first,  to  be  noted  that  the  chromosome  number  of  N.  Are7itsii  is  24 
pairs  while  both  N.  undulata  and  N.  wigandioides  possess  12  pairs.  In  an 
earlier  report  (Goodspeed  1934),  it  was  shown  that  the  somatic  karyotype  of 
N.  undulata  consists  of  chromosomes  with  median  or  submedian  centromere, 
one  of  the  submedian  pairs  satellited.  In  the  case  of  N.  wigandioides  there  are 
8  pairs  with  median  or  submedian  centromere  and  4  with  subterminal,  one  of 
the  subterminal  satellited.  As  regards  size,  there  is  little  difference  between 
the  two  complements;  less,  indeed,  than  between  the  largest  and  smallest 
chromosomes  within  a  complement.  Both  karyotypes  belong  to  the  larger,  but 
not  the  largest  chromosome  size  groups  in  the  genus.  Detailed  study  of  its 
chromosome  morphology  shows  that  the  complement  of  N.  Arentsii  represents 
a  summation  of  the  two  species  types,  with  mostly  median  or  submedian  cen- 
tromere but  a  few  with  subterminal.  The  scant  size  distinctions  between  the 
N.  undulata  and  N.  wigandioides  complements  do  not,  however,  make  it  feasi- 
ble to  homologize  full  complements  but  it  may  be  said  that  the  morphological 
equivalents  of  the  satellited  pair  characteristic  of  each  of  the  two  species  has 
been  identified  in  N.  Arentsii. 

Meiotic  configurations  (pollen  mother  cells)  of  each  of  the  three  species 
were  studied  to  determine  whether  any  characteristic  N.  undulata  or  N. 
wigandioides  features  reappear  in  N.  Arentsii.  In  each  of  the  12-paired 
species,  two  IM  bivalent  types  w^ere  selected  as  sufficiently  distinctive  to  be 
identified  :  in  iV.  undulata  (1)  'a  completely  terminalized  one-ehiasma  bivalent 
with  oblong  partners  and  (2)  a  completely  terminalized  thick  ring  bivalent; 
in  N.  wigandioides  (1)  an  incompletely  terminalized  one-chiasma  bivalent  in 
which  two  droplet  partners  are  separated  by  a  central  (generally  small)  bead 
and  (2)  a  not  quite  terminalized,  two-chiasma  "diamond"  bivalent.  Each  of 
these  types  reappeared  in  N.  Arentsii. 

In  N.  undidata  and  N.  wigandioides,  meiotic  chromosome  behavior  is  normal 
but  in  N.  Arentsii  pairing  was  sometimes  found  to  be  irregular,  even  in  plants 
which  produced  only  good  pollen  grains.  It  is  not  possible  to  say  that  this 
condition  prevailed  in  every  plant,  as  only  a  few  were  examined  in  first 
metaphase,  but  the  fact  that  it  was  true  of  every  plant  examined  is  significant. 
It  was  noted  that  from  2  to  6  chromosomes  (often  in  more  than  50  per  cent  of 
the  PMC)  either  failed  to  pair  or  formed  trivalents.  In  20  second  metaphase 
counts  of  one  plant,  this  irregularity  reduced  to  85  per  cent  the  PMC  showing 
24-24  distribution  of  chromosomes,  the  remaining  15  per  cent  showing  23-25. 

Beyond  a  determination  of  chromosome  number,  the  cytology  of  N.  Arentsii 
had  not  been  studied  at  the  time  hybrids  of  this  new  species  with  N.  undulata 
and  N.  wigandioides  were  made  and  grown.  A  few  plants  each  of  both  F^ 
hybrids  were  secured  and  inasmuch  as  the  preponderance  of  meiotic  material 
available  proved  to  be  from  monosomic  hybrids  made  with  the  same  N.  Arentsii 


Vol.  XXV]         GOODSPEED  :  NICOTIAN  A  ABENTSII  SP.  NOV.  297 

parent,  it  is  clear  that  the  particular  N.  Are7itsn  plant  involved  in  the  crosses 
was  itself  monosomic.  Apart  from  somewhat  complicating  the  determination 
of  extent  of  pairing,  this  situation  did  not  alter  the  essential  evidence,  briefly 
summarized  as  follows : 


Fi  Hybrid 

PMC 

counted 

Plant 
number 

Chromosome 

Constitution 

Pairs  per  . 
Range 

PMC 
Average 

N.  undulata 

14 

F-341  P54: 

36 

11-12 

11.71 

X                      >■ 
N.  Arentsii 

25 

F-385  P51 

35 

11-13 

11.96 

N.  Arentsii 

25 

F-386  P55 

36 

11-13 

11.92 

X                            > 

25 

r-386  P58 

35 

11-13 

12.08 

N.  wigandioides 

25 

F-386  P59 

35 

11-13 

11.96 

In  the  case  of  F^  N.  Arentsii  x  N.  wigandioides,  somewhat  more  than  50  per 
cent  of  the  counts  represent  12  bivalents  only.  The  remainder  involve  combina- 
tions ranging  from  8n  +  Sm  to  13n  with  the  largest  class  lOn  +  liv  and  the  next 
llii  +  liii-  Almost  the  same  types  of  association  occurred  in  the  case  of  F^  N. 
undulata  x  N.  Arentsii  except  that  the  proportion  of  12ii  was  approximately 
40  per  cent.  In  view  of  the  extent  of  multivalent  formation  in  the  two  hybrids 
just  mentioned,  the  occurrence  of  some  pairing  in  the  hybrid  which  represents 
the  third  side  of  the  triangle,  F^  N.  undulata  x  N.  wigandioides,  was  not  un- 
expected. Thus,  a  range  of  from  2  to  9  pairs  was  observed  in  100  pollen  mother 
cells  while  the  average  number  of  pairs  was  approximately  5. 

For  our  present  purpose,  it  is  sufficient  to  call  attention  to  the  fact  that  the 
cytogenetic  evidence  shows  that  the  gametic  sets  of  both  N.  undulata  and  N. 
wigandioides  are  represented  by  a  full  set  of  homologous  chromosomes  in  the 
gametic  set  of  N.  Arentsii  and  that  the  amount  of  pairing  in  F^  N.  undulata  x 
N.~  wigandioides  is  low.  In  other  words,  the  cytogenetic  findings  are  consistent 
Avith  the'  evidence  from  external  morphology  and  from  distribution  which 
indicate  that  N.  Arentsii  represents  an  amphidiploid  product  of  hybridization 
between  those  other  two  species  of  Nicotiana. 

Nicotiana  Arentsii  Goodspeed,  sp.  nov. 

Nicotiana  herbacea;  caule  crasso,  conspicue  pubescente;  foliis  petiolatis, 
late  ellipticis,  rotundo-ovatis  vel  subcordatis;  floribus  in  paniculas  foliosas 
dispositis ;  calyce  magno,  lobis  inaequalibus ;  corolla  dilute  straminea  v.  prope 
alba,  obconico-clavata,  limbo  plus  minusve  zygomorpho,  lobis  obtusis;  stami- 
nibus  inclusis,  abrupte  curvatis,  ad  partem  tubulatam  corollae  inferne  affixis. 

Type. — University  of  California  Second  Botanical  Garden  Expedition  to  the  Andes, 
1938-39,  9666  (U.  C.  Herbarium  665156).  Coll.  C.  Vargas  C,  Nov.  21,  1938,  Limbani,  Dept. 
Puno,  Prov.  Sandia,  Peru.  Alt.  3400  m.  Close  to  the  road  and  houses.  "Cjamasairi."  Other 
authentic  material:  University  of  California  Third  Botanical  Garden  Expedition  to  the 
Andes,  1942^3,  30529.  Coll.  E.  D.  Metcalf,  May  16,  1942,  in  town  of  Limbani,  Dept.  Puno, 
Prov.  Sandia,  Peru.  Alongside  huts,  north  end  of  town.  Alt.  3350  ni. 

Slowly  maturing  annual  or  limited  perennial  1-2  m.  high.  Stem  commonly 
reddish  purplish,  rather  densely  clothed  with  distinct  but  not  long,  weak, 


298  CALIFOBNIA  ACADEMY  OF  SCIENCES  [Proc.  4th  ser. 

silvery-white  hairs,  ultimately  oldest  parts  glabrate,  thinly  corky,  woody 
within ;  main  axis  stiffly  erect,  2-2  cm.  thick,  chief  branches  few,  irregular  in 
length  and  distribution,  usually  some  long  and  rapidly  ascending.  Leaves 
pubescent ;  blade  to  25  or  30  cm.  long  and  about  20  cm.  wide,  rotund-ovate  or 
subcordate  or  the  uppermost  often  lance-ovate,  base  variable,  apex  acute  or 
acuminate,  veins  evident,  sometimes  indenting  upper  surface  a  little,  margin 
slightly  undulate;  petiole  %-^/'io  as  long  as  blade;  frequentlj^  red  coloration 
in  petiole,  midrib,  veins,  margin  or  even  hairs.  Inflorescence  paniculate,  35-50 
cm.  long,  narrow  or  broad,  lower  limits  indistinct  by  very  gradual  reduction 
of  leaves  to  sessile,  broadly  lanceolate  bracts ;  central  axis  erect,  thick,  pubes- 
cent but  seldom  viscid ;  laterals  numerous,  frequently  rather  crowded,  variable 
in  leng-th,  generally  rigid  and  spreading,  paniculately  or  occasionally  cymosely 
branched,  ±:  viscid;  ultimate  pedicels  about  2  mm.  long,  later  6  mm.  Flowers 
abundant,  odorless  or  nearly  so.  Calyx  green,  green  suffused  with  red-purple, 
or  green  blended  with  blue-black,  pubescent,  10-14  mm.  long,  broadly  cylindric 
or  subcampanuiate ;  membranous  area  below  each  sinus  narrow,  short ;  lobes 
shorter  than  tube,  broadly  triangular-ovate,  plane,  unequal,  largest  never 
extending  to  corolla  mouth.  Corolla  distinctly  pubescent  exteriorly  except  in 
the  partially  glabrous  limb;  tubular  part  15-20  mm.  long,  7-9  mm.  wide, 
broadly  ob conic  from  short,  narrower  base,  apically  somewhat  swollen  im- 
mediately below  the  slightly  contracted  mouth,  pale  greenish  yellow  with  pink 
or  purplish  tingeing  above;  entire  limb  (including  rimmed,  hexagonal  mouth) 
to  17  mm.  across,  in  bud  spirally  plicate  and  often  bright  pink,  in  flower 
spreading,  pale  straw  color  frequently  conspicuously  suffused  with  pink 
outside,  rarely  also  some  pink  inside ;  lobes  broad,  obtuse,  3  upper  ones 
smaller,  commonly  a  little  deflexed.  Stamens  inserted  about  5  mm.  from  very 
base  of  corolla  (at  or  near  where  the  obconieal  expansion  begins)  ;  filaments 
thick,  pale  greenish,  4  equal  or  subequal,  wooly  and  erect  for  several  mm. 
above  insertion,  then  glabrous  and  curving  outward  abruptly  to  rest  against 
corolla,  extending  to  about  1  mm.  below  corolla  mouth,  1  sigmoid,  not  wooly 
above  insertion,  commonly  appearing  somewhat  shorter ;  anthers  yellow-green. 
Pollen  pale  straw  color.  Hypogynous  disk  thick,  red  or  red-orange.  Ovary 
rotund-ovoid,  greenish  or  purplish;  style  thick,  faintly  greenish;  stigma 
green,  discoid-pulvinate,  median  groove  shallow.  Capsule  included,  7-9  mm. 
long,  elliptic-ovoid,  brown,  on  dehiscence  upper  half  split  into  2  divergent, 
notched  valves.  Seeds  broadly  elliptic-ovoid,  0.7-0.8  mm.  long,  rather  dark 
brown,  surface  shallowly  reticulate,  meshes  moderately  large,  deeply  lobed 
by  bounding  ridges,  embryo  straight.  Somatic  chromosome  number  48. 

Distinguishable  from  N.  undulata  R.  &  P.  by  later  maturity ;  longer,  more 
distinct,  glistening  hairs;  larger,  rotund-ovate  leaf;  larger  flower;  longest 
calyx  lobe  plane,  not  boat-shaped  in  flower  or  fruit ;  corolla  limb  sometimes^ 
pinkish,  never  greenish ;  from  .¥.  ivigandioides  Koch  &  Fint.  by  shorter  life 
span ;  smaller  leaf ;  markedly  unequal  calyx  segments ;  smaller  corolla  limb. 

Named  in  honor  of  Mr.  George  Arents,  to  whom  botany  is  indebted  for  a| 
comprehensive  compilation  of  the  history  and  culture  of  tobacco. 


Vol.  XXV]         GOODSPEED  :  NICOTIANA  ARENTSII  SP.  NOV.  299 

SUMMARY 

A  new  species  of  Nicotiana,  N.  Arentsii  (n  =  24) ,  from  southeastern  Peru  is 

described,  with  comments  upon  its  morphology,  distribution  and  cytology. 

Similar  comments  are  made  in  the  case  of  two  other  Nicotiana  species,  natives 

of  Peru  and  Bolivia — N.  undulata  (n  =  12)  and  N.  tvigandioides  (n  =  12).  The 

morphological  and  distributional  evidence  points  to  an  amphidiploid  origin 

for  N.  Arentsii  in  which  the  other  two  species  are  involved.  This  hypothesis  is 

confirmed  by  the  cytology  of  the  three  species  and  of  the  F^  hybrids  made 

between  them. 

(Since  the  above  ■was  written,  it  has  been  possible  to  reexamine  the  herbarium  material 
of  two  Bolivian  collections  of  H.  H.  Rusby  which,  before  Nicotiana  Arentsii  was  recognized, 
were  identified  as  N.  wigandioides.  It  is  now  clear  that  they  were  taken  from  plants  of  N. 
Arentsii.  Both  are  Rusby's  No.  822  and  both  are  from  Mapiri  (Dept.  La  Paz,  Prov.  Lare- 
caja)  but  one  is  labelled  "5000  ft.,  April,  1886"  and  the  other  "2500  ft..  May,  1886."  Mapiri 
is  approximately  200  km.  southeast  of  the  Limbani  area  in  which  the  collections  of  N. 
Arentsii  described  above  were  obtained.) 


LITERATURE  CITED 

Bradley,  Muriel  V.  and  Goodspeed,  T.  H. 

1943.  Colchicine-induced  alio-  and  autopolyploidy  in.  Nicotiana.  Proc.  Nat.  Acad.  Sci.  29 : 
295-301. 

Clausen,  R.  E.,  and  Goodspeed,  T.  H. 

1925.  Interspecific  hybridization  in  Nicotiana.  II.  A  tetraploid  glutinosa-tabacum  hybrid, 
an  experimental  verification  of  Winge's  hypothesis.  Genetics  10:278-284. 

Goodspeed,  T.  H. 

1934.  Nicotiana  phylesis  in  the  light  of  chromosome  number,  morphology,  and  behavior, 
Univ.  Calif.  Publ.  Bot.  17:369-398. 

1936.  Phylesis  in  Nicotiana.  Abstract,  Proc.  VI  Internat.  Bot.  Congr.  1:164-165. 

Goodspeed,  T.  H.,  and  Bradley,  Muriel  V. 
1942.  Amphidiploidy.  Bot.  Rev.  8:271-316. 

Goodspeed,  T.  H.,  and  Clausen,  R.  E. 

1928.  Interspecific  hybridization  in  Nicotiana.  VIII.  The  sylvestris-tomentosa-tabactim 
hybrid  triangle  and  its  bearing  on  the  origin  of  tabacum.  Univ.  Calif.  Publ.  Bot. 
11:245-256. 

KOSTOFF,  DONTCHO 

1939.  The  origin  of  the  tetraploid  Nicotiana  from  Bathurst.  Curr.  Sci.  8:110-111. 

RusBY,  H.  H. 

1888.  An  enumeration  of  the  plants  collected  by  Dr.  H.  H.  Rusby  in  South  America.  1885- 
1886.  I.  Bull.  Torr.  Bot.  Club  15:177-184. 

Wheeler,  H-M. 

1935.  Studies  in  Nicotiana.  II.  A  taxonomie  survey  of  the  Australasian  species.  Univ. 
Calif.  Publ.  Bot.  18:45-68. 


i 


EXPLANATION  OF  PLATES 


PLATE   26 

Left:  Nicotiana  Arentsii,  grown  from  seed  of  Univ.  Calif.  Second  Bot.  Gard. 
Exped.  Andes  9666,  Limbani,  Dept.  Puno,  Peru,  coll.  C.  Vargas  C. 

Center:  N.  vndvlata,  grown  from  seed  of  Univ.  Calif.  Bot.  Gard.  Exped. 
Andes  7173,  Pisacc,  Dept.  Cuzco,  Peru,  coll.  James  West. 

Right :  N.  wigandioides,  grown  from  seed  collected  by  Jose  Steinbach,  near 
Cochabaniba,  Bolivia. 


i 


{  3U2  J 


ID 
CM 

UJ 
h 
< 

Q. 

Q 
III 
Lli 

a. 
w 
D 
0 
0 

o 


PLATE   27 

Upper  row :  Nicotiana  ujidulata,  from  seed  collected  by  A.  Weberbauer,  at 
Huancavelica,  Peru.  The  species  is  polymorphic  in  corolla  shape  and  size.  The 
race  figured  is  relatively  large  flowered  with  the  tubular  part  of  its  corolla 
broad  and  little  dilated  below  the  mouth. 

Middle  row :  N.  Arentsii,  source  as  in  plate  26. 

Lower  row:  N.  wiyandioides,  source  as  in  plate  26. 

(Black  paper  has  been  inserted  in  the  corolla  tubes  of  the  sectioned  flowers 
and  one  or  two  stamens  removed  to  show  the  pistil.  In  N.  undulata  and  N. 
Arentsii  those  removed  included  a  short  stamen ;  in  N.  wigandioides  the  two 
removed  are  no  shorter  than  those  remaining.) 


[304] 


PROC.  CALIF.  ACAD.  SCI..  4TH  SERIES.  VOL.  XXV.   NO.    12  [GOODSPEED]    PLATE    27 


[  305] 


PROCEEDINGS 

OF  THE  ^ 

CALIFORNIA  ACADEMY  OF  SCIENCESX 

Fourth  Series 
Vol.  XXV,  No.  13,  pp.  307-322,  figs.  1-13  November  10.  1944 


ALICE  EASTWOOD 
SEMI-CENTENNIAL  PUBLICATIONS 

No.  13 

THE  CYTOGENETICS  OF  HYBRIDS  IN  BROMUS  II. 
BROMUS  CARINATUS  AND  BROMUS  ARIZONICUS 

BY 

G.  L.  STEBBINS  JR.,  H.  A.  TOBGY,  and  JACK  R.  HARLAN 

University  of  California 

INTRODUCTION 

THE  FACT  that  Bromus  carinatus  H.  and  A.  and  its  relatives  constitute  a 
complex  of  polyploid  entities,  mostly  if  not  entirely  of  allopolyploid  or 
hybrid  origin,  has  been  brought  out  in  the  first  paper  of  this  series  (Stebbins 
and  Tobgy,  1944).  In  an  earlier  survey  of  chromosome  numbers  (Steb- 
bins and  Love,  1941)  the  octoploid  number  of  chromosmes  (2n=56)  was 
counted  in  all  representatives  of  this  complex  collected  in  California.  Counts 
made  by  us  of  collections  from  Oregon,  Washington,  Idaho  and  British  Colum- 
bia have  revealed  the  same  number,  so  that  we  can  state  with  some  assurance 
that  the  Pacific  Coast  members  of  this  complex  are  predominantly  if  not  en- 
tirely octoploid,  with  the  exception  of  a  type  originally  collected  by  one  of 
us  (Harlan)  in  Central  Arizona.  The  28  collections  of  the  B.  carinaius  com- 
plex made  in  this  state  were  found  to  be  divided  into  two  series  on  the  basis 
of  morphological  and  ecological  characteristics.  The  first  series,  found  mostly 
in  desert  washes,  roadsides,  and  waste  places  at  lower  altitudes  consists  of 
strictly  annual  plants  with  relatively  long-  awns ;  while  the  second,  found  pre- 
dominantly at  altitudes  of  5000  feet  (1500  meters)  or  higher,  consists  of 
perennials,  with  broad  g'lumes ;  usually  many-flowered  spikelets,  and  shorter 
awns.  The  first  series  agree  with  the  original  description  of  B.  carinatus  var. 
arizonicus  Shear  (1900),  and  would  be  identified  as  B.  carinatus  according  to 

I  307  1 


308  CALIFORXIA  ACADEMY  OF  SCIENCES         [Proc.  4th  ser. 

the  key  of  Swallen  (1942)  in  the  Arizona  flora  of  Kearney  and  Peebles.  The 
second,  since  all  but  one  of  the  specimens  collected  have  essentially  glabrous 
foliage,  "svould  be  considered  B.  polyanthus  Shear,  according  to  the  treatment 
of  Swallen  (Joe.  cit.).  Chromosome  counts  on  five  different  collections  of  this 
second  series  revealed  56  chromosomes  in  them,  as  in  Pacific  Coast  members 
of  the  complex.  A  detailed  description  of  one  collection  of  it  (no.  116),  with 
an  account  of  its  crossing  relationships  with  other  octoploid  species,  is  pre- 
sented elsewhere  (Stebbins  and  Tobgy,  1944). 

The  annual,  long-awned  series,  on  the  other  hand,  was  found  to  possess 
uniformly  the  somatic  number  2n=84,  or  42  bivalents  at  meiosis,  in  material 
from  21  different  collections  (see  map,  fig.  4).  Furthermore,  the  same  chromo- 
some number  was  found  in  a  similar  annual  form  from  Tehachapi,  Kern 
County,  California.  When  these  duodecaploids  were  grown  in  the  same  field 
at  Berkeley  along  with  a  large  series  of  octoploids  from  California  as  well  as 
Arizona,  it  was  found  that  certain  morphological  characteristics  made  them 
readily  distinguishable  from  the  perennial  or  subperennial,  long-awned  octo- 
ploid types  from  lower  altitudes  in  California,  as  well  as  from  the  short-awned 
octoploids  found  at  higher  altitudes  in  both  states.  For  this  reason,  the  specific 
distinctness  of  the  84-chromosome  type  was  suspected,  and  the  testing  of  this 
hypothesis  by  means  of  hybridization  seemed  desirable.  The  results  of  this 
hybridization  and  their  implications  in  regard  to  the  status  and  possible  origin 
of  the  84-chromosome  type  are  here  presented. 

MATERIAL  AND  METHODS 

Seeds  from  the  Arizona  collections  were  all  planted  in  the  field  in  the  sum- 
mer of  1941.  Although  each  progeny  from  a  single  collection  was  with  a  few 
exceptions  remarkably  uniform,  there  was  a  great  deal  of  variation  between- 
collections  (Harlan,  in  press).  Some  of  the  seedlings  behaved  like  desert 
ephemerals,  producing  only  one  or  two  tillers,  which  flowered  in  a  few  weeks 
when  only  10  or  15  cm  high,  and  produced  only  2  or  3  spikelets,  dying  when 
the  seed  matured.  Others  behaved  like  winter  annuals,  and  in  the  spring  pro- 
duced very  tall  plants  with  several  tillers  and  large  panicles.  It  was  from  one 
of  the  latter  type  that  the  parent  stock  for  hj^bridization  with  octoploid  B. 
carinatus  was  selected.  The  particular  collection,  no.  125,  came  from  south  of 
Lake  Meade,  near  Boulder  Dam,  Mojave  County.  The  octoploid  strain  used 
for  hybridization  was  a  typical  B.  carinatus  strain  from  Berkeley,  represented 
in  the  University  of  California  Herbarium  by  the  collection  Stebbins  no.  2677, 
and  described  elsewhere  as  strain  no.  5  (Harlan,  in  press ;  Stebbins  and  Tobgy, 
1944).  The  hybridization  was  made  in  the  field  in  May  1942,  by  Mr.  Henry 
Moser,  according  to  the  method  described  earlier  (Stebbins  and  Tobgy,  loc. 
cit.).  The  cytological  techniques  used  are  described  in  the  same  paper. 


Vol.  XXY]       STEBBINS-TOBGY-HAELAX  :  HYBRIDS  IN  BROMUS  II         300 

MORPHOLOGY  AND  SYSTEMATIC  DESCRIPTION  OF 
THE  SPECIES  AND  THE  HYBRID 

The  morphological  and  cytological  data  presented  below  leaves  no  doubt 
that  the  84-chroinosome  type  is  a  species  sharply  distinct  from  any  now  recog- 
nized in  the  B.  carinatus  complex,  even  though  its  diagnostic  characteristics 
are  somewhat  meager.  Therefore,  a  taxonomic  description  of  it  will  be  given 
at  the  outset,  so  that  it  may  thereafter  be  referred  to  by  its  correct  name. 
Although  the  type  specimen  of  Shear's  B.  carinatus  var.  arizonicus  has  not 
been  seen,  and  the  original  description  is  somewhat  scanty,  nevertheless  there 
is  little  doubt  that  it  belongs  to  the  84-chromosome  species  described  here.  All 
of  the  collections  made  by  Harlan  from  situations  similar  to  Tucson,  its  type 
locality,  are  of  this  species,  and  it  is  the  only  member  of  the  complex  known 
from  lower  altitudes  in  Arizona.  One  collection,  no.  105,  in  which  84  chromo- 
somes were  counted,  came  from  the  Santa  Cruz  Valley  only  35  miles  from  the 
tj^pe  locality.  It  may  be  described  as  follows  (the  taxonomic  section  is  the 
work  of  the  senior  author) . 

Bromus  arizonicus  (Shear)  Stebbins  n.  comb. 

Planta  annua;  culmi  humiles  vel  alti;  paniculae  parvae  spiculis  paucis, 
vel  ampla  spiculis  numerosis;  glumae  elongatae,  secunda  infimam  lemmam 
aequans;  lemmata  ad  marginem  hirsuta,  dorso  scabra  vel  glabra,  lemmata 
superiores  conspicue  bidentata. 

Plants  annual ;  culms  15-140  cm  high,  with  3-5  nodes.  Leaves  somewhat 
glaucous,  glabrous  or  pubescent.  Panicle  sometimes  much  reduced,  but  in  well 
developed  plants  ample,  the  branches  spreading,  in  groups  of  5-6  at  the  lower 
nodes,  the  longest  bearing  5-8  spikelets ;  branches  often  finely  hirsute,  the 
hairs  more  slender  than  in  B.  carinatus.  Spikelets  mostly  5-7  flowered ;  glumes 
subequal  or  distinctly  unequal,  elongate,  the  upper  glume  ahotit  equalling  in 
length  the  lowest  lemma.  Lemmas  of  medium  length,  mostly  11-14  mm  long, 
sparsely  or  densely  hirsute  near  the  edges,  short-scahrous  on  the  hack,  iiden- 
tate,  the  teeth  rather  prominent,  those  of  the  upper  lemmas  often  icith  the 
lateral  nerves  extending  into  them. 

Bromus  carinatus  var.  arizonicus  Shear,  U.  S.  Div.  x\grost,  Bull,  no,  23 : 
62.1900.  Type  in  the  U.  S.  National  Herbarium,  from  near  Tucson,  Arizona, 
C.  G.  Pringle  in  1884.  B.  carinatus  calif  amicus  Shear,  U.  S.  Dept.  Agr.  Div. 
Agrost.  Bull.  23  :60,  at  least  in  part. 

Central  and  southern  Arizona  to  southern  and  central  California,  in  desert 
areas  and  open,  disturbed  ground  of  valleys ;  an  introduced  weed  in  the  north- 
western portion  of  its  present  range.  The  following  specimens,  in  the  herbaria 
of  the  University  of  California  (UC)  and  the  California  Academy  of  Sciences 
(CAS),  are  typical:  CALIFORNIA:  Mark  West  Creek,  jBoZancZer  3985  (UC)  ; 
Davis,  Yolo  County,  Stelhins  3356  (UC)  ;  Alcalde,  Fresno  County,  Eastwood 
13492  (CAS)  ;  Tulare,  Davy  3055  (CAS);  Kettleman  City,  Kings  County, 


310 


CALIFORNIA  ACADEMY  OF  SCIEXCES  [Proc.  4th  ser. 


Hoover  442  (UC)  ;  Bakersfield,  Kern  County,  Davy  1896  (UC)  ;  San  Emigdio 
Caiion,  Kern  County,  Davy  1985  (UC)  ;  Tehachapi,  Kern  County,  Harlan 
128  (UC) ;  Paso  Robles,  San  Luis  Obispo  County,  Barher  in  1900  (UC)  ;  Santa 
Maria,  Santa  Barbara  County,  Eastwood  344  (CAS)  ;  Pasadena,  Los  Angeles 
County,  Grant  117,  117a  (UC,  CAS)  ;  Claremont,  Los  Angeles  County,  Davy 
in  1902  (UC)  ;  San  Bernardino,  Parish  in  1889  (UC)  ;  Colorado  Desert,  Riv- 
erside County,  Hall  5980  (CAS)  ;  Fall  Brook,  San  Diego  County,  Jones  3108 


Fig.  1.  Bronins  carbial us,  sti'ain  no.  5. 
Fig.  2.  Bromus  arizonicus  x  carmatus,  no.  306.5. 
Fig.  3.  Bromus  arizonicus,  no.  311.1  (progeny  of  Harlan  no.  125).  Figs,  la,  2a,  3o,  spike- 
lets,  X  1.  Figs,  lb,  2b,  3b,  lower  glumes;  Ic,  2c,  Sc,  upper  glumes;  Id,  2d,  3d,  lemmas  and 
paleae,  all  x  2.  Figs.  Ic,  2e,  3e,  apices  of  lemmas,  x  4.  Figs.  If,  2f,  of,  portions  of  surface  of 
lemmas,  with  midrib  at  right  and  margin  at  left,  showing  pubescence,  x  8. 


(CAS)  ;  San  Diego,  Orciitt  1178  (UC)  ;  between  Del  Mar  and  Lower  Cali- 
fornia, Lemmon  in  1888  (CAS)  ;  Santa  Catalina  Id.,  Branclegce  in  1890 
(UC) ;  Santa  Cruz  Id.,  Brandegee  in  1888  (UC)  ;  Santa  Rosa  Id.,  Hoffmann 
in  1929  (CAS)  ;  San  Clemente  Id.,  Murlarger  56  (UC)  ;  San  Nicolas  Id., 
J.  T.  Howell  8226  (CAS).  BAJA  CALIFORNIA :  San  Quintin,  Epling  and 
Stewart  in  1936  (CAS)  ;  24.3  miles  east  of  Rosario,  Wiggins  5291  (UC,  CAS). 
ARIZONA:  Wickenburg,  Maricopa  County,  W.  W.  Jones  in  1921  (UC)  ; 
Tempe,  Maricopa  Count)'-,  Gillespie  5519  (UC)  ;  Fort  Lowell,  Thornher  535 
(UC)  ;  25  miles  southeast  of  Lake  Meade,  Mohave  County,  Harlan  125  (UC)  ; 
east  of  King-man,  Mohave  County,  Harlan  123  (UC)  ;  Pinal  Mts.,  southwest 
of  Globe,  Gila  County,  Harlan  112  (UC)  ;  Salt  River  Canyon,  Gila  County, 
Harlan  115  (UC)  ;  Sacaton,  Pinal  County,  Harlan  103  (UC)  ;  east  of  Flor- 


Vol.  XXV]       STEBBINS-TOBGY-HAELAN :  HYBRIDS  IN  BEOMUS  II         311 

ence,  Pinal  County,  Harlan  107  (UC)  ;  Red  Rock,  Pinal  Comity,  Harlan 
105  (UC). 

As  stated  preYioiisl.y,  the  chromosome  number  2n=84  has  been  determined 
in  all  of  the  Harlan  collections  cited. 

Without  having  seen  the  type  specimen,  the  writer  cannot  determine 
whether  B.  arizonicus  is  the  same  as  B.  carinatus  calif ornicus  Shear,  or  B. 


^7^^. 


Fig.  4.  Map  showing  distribution  of  Harlan  collections  of  Bromus  arizonicus  (2n  =84), 
solid  circles,  and  B.  ijolyanthus  vel  aff.  (2?i  =  56),  hollow  squares,  in  Arizona.  The  5000  foot 
contour  level  is  drawn. 

calif  ornicus  Nutt.  Nuttall's  specific  name  is  a  nomen  nudum  (cf.  Shear  1900), 
and  his  specimen,  according  to  Shear,  "is  a  mere  scrap,"  of  which  the  locality 
of  collection  is  indefinite.  Nevertheless,  the  specimens  cited  by  Shear  under 
B.  carinatus  calif  ornicus,  since  all  come  from  San  Diego  or  Lower  California, 
are  very  likely  B.  arizonicus. 

The  characteristics  which  distingiiish  B.  arizonicus  from  all  other  members 
of  the  subgenus  Ceratochloa  in  the  Western  United  States  are  the  strictly 
annual  habit  (the  plants  always  die  when  the  seeds  have  matured),  the  elon- 
gate glumes,  the  hirsute  pubescence  near  the  margin  of  the  lemmas,  and  the 
rather  prominent  lobes  of  at  least  the  upper  lemmas  (fig.  2) .  All  of  the  typical, 
octoploid  members  of  the  B.  carinatus  complex  as  grown  in  Berkeley  pro- 
duced some  new  tillers  after  the  seed  had  ripened,  except  when  the  plants 


312 


CALIFORNIA  ACADEMY  OF  SCIEXCES  [Proc.  4th  ser. 


were  obviously  diseased.  In  all  of  them  the  lowest  lemma  is  distinctly  longer 
than  the  upper  glume ;  its  pubescence,  whether  long  or  short  is  more  or  less 
evenly  distributed  over  the  surface ;  and  the  lobes  are  very  short,  with  the 
latei-al  nerves  ending  far  below  their  base  (fig.  1). 

The  relative  distributions  of  B.  arizonicus  and  the  octoploids  referred  to 
B.  folyanthiis  are  shown  on  a  map  (fig.  4)  giving  the  localities  from  which 
chromosome  counts  were  obtained.  Only  one  collection  from  California  has 


TABLE  1 

Comparisons  of  the  Diagnostic  Characteristics  of  B.  carinatus, 
B.  arizonicus,  and  Their  Fi  Hybrid 


Number  of  nodes  of  culms 

Pubescence  of  top  node 

Pubescence  of  upper  culm  leaf 

Maximum  number  of  branches  at  node 

of  panicle 

Maximum  number  of  florets  per  spikelet 

Length  second  glume 

Length  second  lemma 

Ratio,  glume/lemma 

Pubescence  of  second  lemma 

Lobes  of  upper  lemmas 


B.  carinatus 
(no.  5) 


5-6 
+  below 

+  lower 


5-8 

8-9 
14.3 
16.4 

0.87 
+  + 
throughout 
small 


Fi  hybrid 
(no.  125  X  no.  5) 


5 

+  + 
throughout 
+  +  upper 
+  lower 

4-5 

7-9 

16.0 

15.2 

1.05 

+  +  margins 

+  back 
intermediate 


B.  arizonicun 
(no.  125) 


4-5 

+ 

throughout 

+  upper 

0  lower 

4-6 

6-7 

12.4 

12.3 

1.01 

+  +  margins 

0-+  back 

prominent 


been  counted,  that  from  Tehachapi  (no.  128),  but  the  external  morphology 
of  the  other  California  specimens  cited  leaves  no  doubt  as  to  their  specific 
identity.  At  Davis,  California,  the  northern  limit  of  its  known  range,  B. 
arizo7iicus  is  obviously  a  recent  introduction  along  the  railroad  tracks,  and  it 
is  very  likely  introduced  at  the  San  Joaquin  Valley  localities  also. 

The  particular  strain  of  B.  arizonicus  used  for  hybridization  was  one  of  the 
most  vigorous  and  latest  in  flowering  of  all.  As  recorded  elsewhere  (Harlan, 
1942;  Stebbins  and  Tobgy,  1944),  the  no.  5  strain  of  B.  carinatus  is  also  a 
very  vigorous  one.  It  is  about  intermediate  or  somewhat  late  in  flowering 
compared  to  other  Berkeley  strains,  but  earlier  than  most  of  the  montane 
types.  It  is  distinguished  from  most  other  B.  carinatus  strains  from  Berkeley 
by  its  rather  large  spikelets  and  lemmas;  the  average  length  of  the  lemmas 
and  awns  in  B.  carinatus  as  a  whole  is  not  very  different  from  that  in  B. 
arizonicus.  The  chief  distinguishing  characteristics  of  the  two  parental  strain* 
are  given  in  Table  1. 

No  hybrid  seeds  were  obtained  out  of  42  florets  emasculated  when  B.  ari' 
zonicus  was  used  as  the  pollen  parent  on  B.  carinatus.  In  the  reciprocal  cross, 


Vol.  XXV]       STEBBINS-TOBGT-HABLAN :  HYBRIDS  IN  BBOMUS  II        313 

B.  arizo7iicus  5  x  -S.  carinatus  ^,  37  seeds  were  obtained  out  of  40  florets 
emasculated.  Of  these,  10  were  sown  in  September,  1942.  Four  were  lost 
through  an  accident,  and  six  were  planted  in  the  field  in  November.  Of  these 
4  turned  out  to  be  hybrids.  Morphologically,  they  were  intermediate  between 
their  parents,  except  that  in  number  of  florets  per  spikelet  they  were  nearer 
their  B.  carinatus  parent,  and  in  relative  size  of  glumes  and  lemmas,  like  B. 
arizonicus  (Table  1).  They  proved  to  be  completely  pollen  and  seed  sterile,  a 
condition  which  would  be  expected  from  the  cytological  situation  described 
below.  Three  of  them  died  during  May,  1943,  even  before  their  annual  B. 
arizonicus  parent.  A  fourth  showed  some  tendency  to  grow  out  after  being  ciit 
back,  as  does  B.  carinatus,  but  died  during  the  fall  of  1943.    . 

CYTOLOGY  OF  THE  PARENTS  AND  HYBRID 

1.  B.  carinatus:  The  meiotic  chromosome  behavior  of  strain  no.  5  of  B.  cari- 
natus used  in  this  study  has  been  reported  earlier  (Stebbins  and  Tobgy, 
1944).  The  main  features  may  be  outlined  here.  At  diakinesis  and  I-meta- 


♦ftt*J^»V 


:  >.«•  • 


^^     ^^ 


,-'' 


Figs.  5-7,  chromosome  pairing  at  first  metaphase  in:  5,  B.  carinatus;  6,  B.  arizonicus ; 
7,  B.  arizonicus  x  carinatus.  All  reproduced  X  1300. 


314 


CALIFORNIA  ACABEMY  OF  SCIENCES  [Pkoc.  4th 


SER. 


8 


•To  5 


%  % 


•  • 


U^  \ 


10 


m ' 


<    •  I 


w    > 


'rv 


1  1 


^ 


^.<| 


f; 


2u 


13 


Figs.  8-13.  Photomicrographs  showing  chromosome  behavior,  all  reproduced  X  800. 
8,  B.  carinatus,  I  metapliase;  9,  B.  arisoiiicus,  diakinesis;  10  and  11,  B.  arisonicus  x  cari- 
natuti,  I  mctaphase:  12  and  13,  B.  arisonicus  x  carinatiis,  I  anapliase. 


Vol.  XXV]       STEBBINS-TOBGY-HARLAN :  HYBEIBS  IN  BEOMUS  II         315 

phase  there  were  always  28  bivalents ;  no  univalents  or  multivalents  were  ever 
seen.  Seven  bivalents  could  be  distinguished  by  their  conspicuously  large  size, 
while  the  remaining  21  bivalents  were  uniformly  smaller  and  may  therefore 
be  termed  medium-sized  (figs.  5  and  8).  At  I-metaphase  most  of  the  bivalents 
had  two  chiasmata,  one  in  each  arm,  and  occasionally  some  of  the  large  biva- 
lents had  three  chiasmata.  The  majority  of  the  bivalents,  large  and  medium- 
sized,  were  therefore  pairing  in  both  arms,  although  rarely  one  or  two  of  them 
were  pairing  in  one  arm  only.  In  9  per  cent  of  the  I-anaphase  cells  and  in  1  per 

TABLE  2 
Chromosome  Pairing  at  Meiosis  in  Bromus  arizonicus  X  carinaius 


Combination 

No.  of  cells* 

Per  cent 

niiii 

mjj 

m, 

ii 

7 

14 

14 

7 

12(2) 

33.5 

6 

15 

15 

7 

6 

16.2 

5 

16 

16 

7 

6 

16.2 

4 

17 

17 

7 

9(2) 

24  3 

3 

18 

18 

7 

2(1) 

5.4 

2 

19 

19 

7 

2(1) 

5.4 

Total     . .  . 

37 

100.0 

'  The  numbers  in  parentheses  refer  to  6  cells  in  which  two  of  the  chromosomes  regularly  forming  a  mji  ap- 
peared instead  as  2mi. 


cent  of  the  Il-anaphase  cells  one  inversion  bridge  and  a  fragment  were  seen. 
Almost  99  per  cent  of  the  tetrads  had  4  normal  microspores ;  in  the  remaining 
1  per  cent,  one  or  two  micronuclei  were  also  present.  There  was  81.0  per  cent 
stained  pollen. 

2.  B.  arizonicus:  At  diakinesis  and  I-metaphase  of  B.  arizonicus  there  were 
always  42  medium-sized  bivalents  (figs.  6  and  9)  ;  univalents  and  multivalents 
were  significantly  missing.  There  was  no  differentiation  into  large  and  small 
bivalents  and  none  of  the  bivalents  could  be  classified  in  the  large  size  category 
characteristic  of  seven  bivalents  in  B.  carinatus.  In  almost  half  of  the  I-meta- 
phase cells  all  the  bivalents  were  paired  in  both  arms  and  in  the  other  half 
one,  two,  or  three  bivalents  were  paired  in  one  arm  only.  There  was  less  in- 
version hyljridity  exhibited  in  the  B.  arizonicus  plant  examined  than  in  that 
of  B.  carinatus.  Only  1  per  cent  of  the  I-anaphase  cells  and  0.4  per  cent  of  the 
Il-anaphase  cells  showed  one  chromatid  bridge  and  one  fragment.  About  2.5 
per  cent  of  the  tetrads  had  one  or  two  micronuclei  in  addition  to  the  4  normal 
microspores,  but  the  remaining  97.5  per  cent  had  4  large  microspores  only. 
There  was  90  per  cent  stained  pollen. 

3.  B.  arizonicus  x  B.  carinatus.  The  F^  hybrid  plants  (2n  =  70)  between  B. 
arizonicus   (2n  =  84)   and  B.  carinatus  (2n  =  56)  received  42  medium-sized 


31G 


CALIFORNIA  ACADEMY  OF  SCIENCES  [Proc.  4th  ser. 


chromosomes  from  B.  arizonicus  and  21  medium  +  7  large  chromosomes  from 
B.  carinatus.  Thirty-seven  cells  at  I-metaphase  were  completely  analyzed  in 
the  hybrid  (Table  2). 

Pairing  of  the  parental  chromosomes  followed  a  comparatively  simple 
scheme  which  may  be  described  as  follows : 

1.  The  7  large  chromosomes  from  B.  carinatus  were  always  seen  as  7  unpaired  univalents 
(li  in  the  table). 

2.  At  least  2  sets  of  7  medium  chromosomes  from  B.  arizonicus  also  appeared  as  14  un- 
paired univalents  (m,  in  the  table). 

3.  At  least  14  medium  bivalents  were  almost  always  seen  (mn  in  the  table)  ;  these  being 
formed  from  the  consistent  pairing  of  2  sets  of  7  medium  chromosomes  from  B.  arizonicus 
with  2  corresponding  sets  from  B.  carinatus. 

4.  A  maximum  number  of  7  trivalents  was  formed  (mm  in  the  table) ;  these  resulted 
from  partial  or  complete  homology  between  one  set  from  B.  arizonicus,  another  from  the 
same  species,  and  a  third  from  B.  carinatus.  When  the  maximum  number  of  7  trivalents 
was  formed,  there  were  14  medium  bivalents,  14  medium  univalents  and  7  large  univalents 
in  the  cell.  With  decreasing  numbers  of  trivalents,  the  number  of  bivalents  and  univalents 
was  correspondingly  increased.  A  typical  I-metaphase  cell  is  shoA^Ti  in  figure  7,  and  two 
other  cells  in  figures  10  and  11.  The  percentage  of  cells  with  these  verying  numbers  of 
trivalents  is  given  in  the  last  column  of  Table  2.  A  slight  reduction  in  the  normal  pairing 
in  a  few  cells  was  found  (cf.  footnote.  Table  2). 

TABLE  3 
Sporad  Analysis  in  B.  arizonicus  X  carinatus 


Number  of  cells 

Number  of  micronuclei 

Totals 

Per  cent 

in  sporad 

7-11 

12-16 

17-21 

4  cells 

5  cells     

29 

15 
8 
2 

51 
23 
13 

4 

27 
6 
4 

1 

107 
44 
25 

7 

58.5 
24.0 

6  cells 

13.7 

7  cells 

3.8 

Totals 

Percentages 

54 
29.5 

91 
49.7 

38 
20.8 

183 

100.0 

There  was  good  evidence  of  extensive  inversion  hybriditj'  in  the  F^.  This 
was  shown  by  bridge-fragment  configurations  in  48.5  per  cent  of  the  cells  at 
I-anaphase  and  by  16.5  per  cent  of  the  cells  at  Il-anaphase.  Usually  only  one 
bridge  and  occasionally  two  were  seen  at  I-anaphase,  and  only  once  was  a  cell 
with  three  bridges  encountered  (fig.  13).  All  the  bridges  at  Il-anaphase  were 
genuine  second  division  bridges  and  not  relics  from  the  first  division.  There- 
fore the  extent  of  inversion  hybridity  in  the  hybrid  may  be  represented  as 
one  bridge  in  65  per  cent  of  the  pollen  mother  ceUs.  It  could  not  be  always 
determined  whether  the  inversions  were  confined  to  the  bivalents  or  were 
found  in  the  trivalents  also,  but  evidence  was  obtained  of  bridges  in  both 
types  of  configurations.  Therefore  it  is  likely  that  the  two  groups  of  parental 


Vol.  XXY]       STEBBINS-IOBGY-EABLAN  :  HYBBIDS  IN  BEOMUS  II        317 

chromosomes  whicli  regularly  pair  differ  by  inversions  as  well  as  the  two 
partly  homologous  sets  £rom  B.  arizonicus  which  enter  into  the  trivalents. 
Since  heterozygosity  for  inversions  was  found  to  a  small  degree  in  both 
parents,  it  is  reasonable  to  suppose  that  most  of  the  pairing  sets  differ  by  at 
least  a  few  inversions. 

The  majority  of  the  univalent  chromosomes  divided  equationally  and  the 
two  daughter  univalents  separated  to  the  two  poles  at  I-anaphase,  but  occa- 
sionally some  of  the  univalents,  though  already  divided,  would  fail  to  undergo 
the  anaphasic  separation  and  would  pass  entirely  to  one  pole  or  the  other 
(figs.  12  and  13).  There  was  excessive  chromosome  lagging  at  the  second 
division,  which,  together  with  that  at  the  first  division,  may  best  be  repre- 
sented by  the  very  complex  condition  of  the  sporads  (Table  3) . 

"With  such  conditions  prevailing  in  the  sporads  the  repeated  observation  of 
complete  sterility  of  both  pollen  and  ovules  is  the  expected  condition. 

DISCUSSION 

Specific  distinctness  of  B.  arizonicus.  The  experimental  evidence  presented 
here  shows  that,  because  of  the  extreme  abnormality  of  meiosis  in  their  Fi 
hybrid,  and  the  complete  inviability  of  both  pollen  and  ovules,  B.  arizonicus 
and  B.  carinatus  are  completely  incapable  of  interbreeding,  that  is  of  ex- 
changing genes.  Therefore,  they  would  be  recognized  as  distinct  species  under 
any  modern  definition  of  this  entity  (cf.  Dobzhanskj^  1941,  p.  373).  Further- 
more, the  cytological  evidence  indicates  that  B.  arizonicus  would  be  equally 
intersterile  with  any  other  octoploid  member  of  the  B.  carinatus  complex 
(cf.  Stebbins  and  Tobgy,  1944).  The  two  species  clearly  are  not  made  up  of 
the  same  chromosomal  material.  Out  of  the  haploid  complement  of  42  in  B. 
arizonicus,  21  chromosomes  are  nearly  or  quite  homologous  to  21  of  the  haploid 
set  of  28  in  B.  carinatus,  7  are  partly  homologous  with  7  of  these  21  carinatus 
chromosomes  and  14  are  entirely  different.  The  resemblance  between  B.  ari- 
zonicus and  B.  carinatus,  therefore,  lies  principally  in  their  common  posses- 
sion of  21  pairs  of  medium-sized  (m)  chromosomes. 

Cytological  constitution  and  possiMe  origin  of  B.  arizonicus.  In  another 
paper  (Stebbins  and  Tobgy,  1944)  it  was  shown  that  the  21  medium-sized 
chromosomes  (m)  iiiB.  carinatus  are  almost  completely  homologous  with  the 
21  chromosomes  which  form  the  entire  haploid  complement  of  the  South 
American  species,  B.  catharticus  Valil  (B.  unioloides  HBK).  In  that  paper, 
these  sets  were  designated  AABBCC,  so  that  the  genome  formula  of  the  various 
octoploid  members  of  the  B.  carinatus  complex  was  written  as  AABBCCLL, 
LL  representing  the  sets  of  7  large  chromosomes.  The  present  evidence  shows 
that  B.  arizonicus  possesses  the  AABBCC  set,  perhaps  somewhat  modified,  but 
lacks  the  7  pairs  of  large,  or  the  LL  chromosomes.  Of  its  other  three  sets,  one 
may  be  designated  as  Co  C2,  because  of  its  similarity  to  one  set  of  the  catharti- 
cus group,  this  partially  homologous  set  being  arbitrarily  designated  as  the 
CC  one.  The  remaining  two  sets  of  B.  arizonicus  may  be  designated  DDEE. 


318  CALJFOKXIA  ACADEMY  OF  SCIENCES  [Proc.  4th  ser. 

Thus  the  g-enome  formula  of  B.  arizonicus  may  be  written  AABBCjCiCaCo 
DDEE.  It  should  be  clearly  understood  that  this- formula  is  used  only  to 
express  pairing  relationships ;  the  actual  genie  content  of  the  genomes  prob- 
ably varies  considerably  from  species  to  species,  and  they  undoubtedly  differ 
in  the  various  species  by  minor  structural  variations. 

This  cytological  constitution  indicates  that  B.  arizonicus  is  an  allopoly- 
ploid derived  bj^  chromosome  doubling  from  a  hybrid  between  B.  catharticus 
or  a  close  relative  of  it  and  some  other  species  with  21  pairs  of  medium-sized 
chromosomes.  The  identity  of  this  other  parent  is  not  yet  established,  but 
based  on  the  morphological  differences  between  B.  arizonicus  and  B.  cai'inatus, 
its  main  features  can  be  postulated.  They  are  as  follows : 

1.  It  should  be  an  annual,  preferably  found  in  desert  or  at  least  semiarid  situations. 

2.  Its  panicle  branches  should  be  slender,  but  should  be  glabrous  or  finely  short-pubescent, 
rather  than  coarsely  scabrous  as  in  B.  catharticus. 

3.  Its  glumes  should  be  elongate,  definitely  exceeding  the  lower  lemmas.  The  outer  glume 
should  have  chiefly  3  nerves. 

4.  Its  lemmas  should  be  rounded  on  the  back  or  weakly  keeled,  should  be  long-hirsute 
near  the  margins,  conspicuously  bidentate  at  the  apex,  and  long-awned. 

5.  It  should  form  a  maximum  of  7  bivalents  in  hybrids  with  B.  catharticus  or  B.  carinatus. 

The  morphological  characteristics  listed  here  are  found  in  only  one  species 
of  Bromus  known  to  us,  B.  Trinii  Desv.  This  species  also  has  the  required  21 
pairs  of  medium-sized  chromosomes,  as  determined  by  Knowles  (1944)  as 
well  as  by  the  senior  author.  Its  pairing  relationships  with  B.  catharticus  and 
B.  carinatus  have  not  yet  been  tested.  The  hypothesis  may,  however,  be  ad- 
vanced that  B.  arizonicus  is  an  allopolyploid  derived  from  B.  catharticus  and 
B.  Trinii  or  close  relatives  of  these  species.  It  is  hoped  that  this  can  be  tested 
in  the  near  future  by  appropriate  hybridization  experiments.  The  hypothesis 
involves  some  difficulties  from  the  historical  point  of  view,  since  both  of  the 
putative  parents  are  generally  believed  to  be  native  only  to  South  America, 
and  to  be  recent  introductions  into  the  area  now  occupied  by  B.  arizonicus. 
This  problem,  however,  had  best  be  considered  after  the  evidence  in  favor  of 
the  proposed  hypothesis  has  become  more  definite. 

There  is  one  cytological  discrepancy  found  in  the  pairing  behavior  of  the 
chromosomes  of  B.  arizonicus  in  the  pure  species  as  compared  with  their  be- 
havior in  its  hybrid  with  B.  carinatus.  The  trivalents  found  in  the  hybrid 
indicates  definitely  a  partial  homology  between  two  of  the  sets  in  B.  arizonicus. 
That  they  could  not  possibly  consist  of  two  sets  from  B.  carinatus  and  one 
from  B.  arizonicus  is  evident  from  the  fact  that  in  another  hybrid,  B.  carina- 
ius  X  B.  mollis,  the  chromosomes  nearly  all  remain  unpaired,  showing  that 
there  is  little  or  no  homology  between  any  of  the  four  sets  found  in  B.  carinatus 
(Knowles  1944) .  The  difficulty  is  that  these  two  partially  homologous  sets  of  B. 
arizonicus  should  occasionally  form  quadrivalents  in  the  pure  species.  How- 
ever, in  several  different  strains,  examined  by  each  of  the  three  authors,  the 
presence  of  42  bivalents  was  observed  with  strikingly  consistent  regularity, 


Vol.  XXV]       STEBBINS-TOBGY-HARLAN :  HYBRIDS  IN  BROMUS  II         319 

and  the  partial  homology  between  two  of  the  sets  was  not  even  suspected  until 
the  cytology  of  the  hybrid  was  studied.  Two  explanations  for  this  phenomenon 
are  suggested.  The  first  is  that  advanced  by  Miintzing  and  Prakken  (1940) 
for  a  similar  behavior  in  Phleum  pratense,  which  was  also  found  in  the  same 
species  by  Myers  (1941) .  They  assume  that  in  Phleum  and  other  genera  there 
is  a  "special  genotypically  controlled  tendency  to  bivalent  formation"  (p.  494) . 
The  other  explanation  is  that  of  differential  affinity,  or  preferential  pairing, 
advanced  by  Darlington  (1927,  p.  198)  to  explain  the  lack  of  multivalents  in 
the  allopolyploid  Primula  kewensis,  and  by  Skirm  (1942),  to  explain  the 
difference  in  pairing  found  in  two  different  tetraploicls  of  Tradescantia,  both 
obtained  from  the  same  diploid  hybrid.  From  the  data  now  at  hand,  there  is 
no  way  of  concluding  which  is  the  best  explanation  for  the  situation  in  Bromus 
arizonicus. 

In  conclusion,  the  following  points  have  arisen  from  this  study  which  are 
of  particular  interest  to  students  of  the  taxonomy  of  the  Gramineae.  In  the 
first  place,  whatever  the  second  parent  of  B.  arizonicus  is,  it  most  certainly 
belongs  neither  to  the  section  Ceratochloa  nor  to  the  section  Bromopsis,  as 
postulated  for  the  second  parent  of  the  octoploid  B.  carinatus  and  its  relatives 
(Stebbins  and  Tobgy,  loc.  cit.) .  The  morphological  characteristics  listed  above 
would  immediately  remove  the  species  from  the  former  section,  while  in  the 
latter  the  chromosomes  are  large,  like  the  LL  set  of  B.  carhiatus  (Stebbins 
and  Love,  1941).  Both  B.  carinatus  and  B.  arizonicus,  therefore,  are  allopoly- 
ploids from  intersectional  hybrids,  but  the  two,  sections  thus  connected  by 
allopolyploidy  are  different  in  each  case.  Therefore  the  cytogenetic  evidence 
entirely  supports  the  recognition  of  Bromus  as  a  single  genus,  rather  than  the 
separation  of  Ceratochloa  as  generically  distinct,  as  has  been  done  by  a  num- 
ber of  agrostologists.  It  speaks  even  more  emphatically  against  the  splitting 
of  Bromus  into  several  different  genera,  as  was  proposed  by  Nevski  (1934). 

In  the  second  place,  the  particular  type  of  relationship  which  exists  between 
B.  carinatus  and  B.  arizonicus  will  probably  be  found  to  be  rather  frequent 
when  more  genera  of  this  family  are  studied  cytogenetically,  since  polyploidy 
is  so  prevalent  in  them.  They  are  a  typical  example  of  species  which  are  mor- 
phologically very  similar,  and  cytogenetically  partly  very  similar  and  partly 
very  different.  Their  similarities  are  due  to  their  common  possession  of  the 
B.  catharticus  set.  Their  morphological  differences  represent,  in  a  much  di- 
luted form,  the  differences  between  two  different  sections  of  the  genus.  Both 
of  these  sections  diverge  from  typical  Ceratochloa  in  their  much  less  strongly 
keeled  lemmas  and  much  longer  awns ;  hence  both  B.  arizonicus  and  B.  carin- 
natus  diverge  from  B.  catharticus  in  the  same  respects.  B.  carinatus  and 
B.  arizonicus  may  therefore  be  spoken  of  as  allosyngenic  amphidiploids  (or 
allopolyploids),  since  they  differ  in  respect  to  one  of  their  original  parents, 
and  have  the  other  in  common.  The  likelihood  is  that  in  all  genera  possessing 
large  polyploid  complexes,  allosyngenic  amphidiploids  form  one  of  the  chief 
sources  of  taxonomic  difficulty. 


320  CALIFORNIA  ACAVEMY  OF  SCIEXCES  [Pkoc.  4th  ser. 

SUMMARY 

1.  A  new  species,  B.  arizonicus,  is  described,  based  on  B.  carinatus  variety 
arizonicus  Shear.  This  species  has  the  somatic  chromosome  number  2n  =  84, 
as  compared  with  2n  =  56  for  B.  carinatus. 

2.  Tlie  hybrid,  B.  arizonicus  x  B.  carinatus,  can  be  obtained  only  when  B. 
arizonicus  is  the  female  parent.  It  is  morphologically  intermediate  between 
its  parents  and  is  completely  sterile. 

3.  At  meiosis  this  hybrid  regularly  shows  a  maximum  of  7  trivalents,  with 
the  rest  of  the  chromosomes  existing  as  bivalents  and  univalents.  The  7  large 
chromosomes  obtained  from  B.  carinatus  have  no  homologues  in  B.  arizonicus, 
and  therefore  are  always  unpaired,  as  are  also  14  medium-sized  chromosomes 
from  B.  arizonicus. 

4.  The  presence  of  bridge-fragment  configurations  in  either  the  first  or  the 
second  division  of  meiosis  in  65  per  cent  of  the  pollen  mother  cells  indicates 
that  the  chromosomes  of  two  or  more  of  the  pairing  sets  differ  hy  a  number  of 
inverted  segments. 

5.  The  cytogenetic  evidence  amply  demonstrates  the  specific  distinctness  of 
B.  carinatus  and  B.  arizonicus.  Their  genomic  formulae  may  be  written  as 
follows :  for  B.  carinatus  AKB^GS^J-iJj^,  for  B.  arizonicus  AABBC1C1C2C2 
DDEE.  The  A,  B,  and  Ci  sets  constitute  the  chromosomal  complements  of  B. 
catharticus.  It  is  possible  that  B.  arizonicus  possesses  this  complement  in  a 
slightly  modified  form. 

6.  The  hypothesis  is  advanced  that  B.  arizonicus  is  an  allopolyploid  derived 
from  doubling  of  the  chromosome  number  in  a  hybrid  between  B.  catharticus 
or  a  close  relative  of  that  species  and  some  other  species  with  21  pairs  of 
medium-sized  chromosomes,  not  belonging  to  the  subgenus  Ceratochloa. 

7.  The  cytogenetic  evidence  here  presented  supports  the  retention  of 
Bromus,  sens,  lat.,  as  a  single  genus. 

8.  The  relationship  of  B.  carinatus  and  B.  arizonicus  is  described  as  that  of 
allosyngenic  amphidiploids.  This  type  of  relationship  is  believed  to  be  a  com- 
mon source  of  taxonomie  difficulty  in  genera  which  have  highly  developed 
polyploid  complexes. 


Vol.  XXV]       STEBBINS-TOBGT-HARLAN :  HYBRIDS  IN  BEOMUS  II         321 

LITERATURE  CITED 

Darlington,  C.  D. 

1937.  Eecent  Advances  in  Cytology,  2nd  ed.  671  pp.  Philadelphia,  Blakiston's. 

Harlan,  J.  R. 

1942.  A  survey  of  the  genetic  variability  in  the  Bromus  carinatus  complex.  Thesis  (Ph.D.), 
Univ.  of  California. 

Knowles,  p.  F. 

1944.  Interspecific  hybridizations  of  Bromus.  Genetics  29:  128-140. 

MiJNTZiNG,  A.,  and  R.  Prakken 

1940.  The  mode  of  chromosome  pairing  in  Phleiim  twins  with  63  chromosomes  and  its 
cytogenetic  consequences.  Hereditas  26:463-501. 

Myers,  W.  M. 

1941.  Meiotie  behavior  of  Phleum  pratense,  Pldeum  siCbulaium,  and  their  Fi  hybrid.  Journ. 
Agr.  Res.  63:649-659. 

Nevski,  S.  a. 

1934.  Sched.  Herb.  Flor.  Asiae  Mediae,  in  Acta  Univ.  Asiae  Mediae,  Tashkent,  Ser.  VIII 
b,  Bot.,  Fase.  17:14-20. 

Shear,  C.  H. 

1900.  Studies  on  North  American  grasses.  A  revision  of  the  North  American  species  of 
Bromus  occurring  north  of  Mexico.  U.  S.  Bept.  of  Agric.  Div.  Agrost.  Bull.  no.  23. 
66  pp. 

Skirm,  G.  W.' 

1942.  Bivalent  pairing  in  an  induced  tetraploid  of  Tradescantia.  Genetics  27 :  635-640. 

Stebbins,  G.  L.  Jr.,  and  E.  M.  Love 

1941.  A  Cytological  study  of  California  forage  grasses.  Am.  Jour.  Bot.  28  :  371-382. 

Stebbins,  G.  L.,  Jr.,  and  H.  A.  Tobgy 

1944.  The  cytogenetics  of  hybrids  in  Bromus.  I.  Hybrids  within  the  section  Ceratochloa. 
Am.  Jour.  Bot.  31:1-11. 

Swallen,  J.  R. 

1942.  Gramineae,  in  Kearney  and  Peebles,  Flowering  Plants  and  Ferns  of  Arizona.  U.  S. 
Dept.  Agric.  Misc.  Publ.  no.  423.  1069  pp. 


PROCEEDINGS 

OF  THE 


Uj 


CALIFORNIA  ACADEMY  OF  SCIENCES  V. 

FoLirtli  Series 

Vol.  XXV,  No.  14,  pp.  323-356,  pis.  28-3G,  3  charls,  3  hislogianis 

November  iG,  1944 


ALICE  EASTWOOD 
SEMI-CENTENNIAL  PUBLICATIONS 


No.  14 

THE  IMPORTANCE  OF  FIELD  HYBRIDS  IN 
DETERMINING  SPECIES  IN  THE 

GENUS  CEANOTHUS 

BY 

HOWARD  E.  McMINN 

Professor  of  Botany,  Mills  College 

«VERY  FIELD  BOTANIST,  liaviiig  tlie  faculty  of  careful  observation,  is  repeat- 
edly impressed  v^ith  the  number  of  apparent  hybrids  which  occur  in 

I  certain  genera  of  seed-bearing  plants.  The  genus  Ceanothus  is  one  of  these 
genera.  According  to  Katherine  Brandegee,  Dr.  C.  C.  Parry  was  the  first  to 
call  attention  to  the  prevalen'ce  of  natural  hybrids  in  this  genus.  Mrs.  Brande- 
gee in  her  "Studies  of  Ceanothus"  (1894)  discussed  the  various  hybrids  which 

^she  had  observed  in  the  field  and  herbaria.  She  was  of  the  opinion  that  most 
of  the  hybrids  between  members  of  the  same  section  were  fertile  but  she  gave 
no  supporting  evidence.  She  stated  that,  "The  only  infertile  hybrid,  within 
its  section,  known  to  me  is  No.  69,  C.  incanus  x  papillosus.  In  this  the  ovaries 
are  more  or  less  abortive  and  no  fruit  was  formed."  If  we  accept  the  sterility 
test  for  distinct  biological  species,  the  two  parents,  C.  incanus  and  C.  pajyil- 

Hosus,  in  this  cross  belong  to  two  different  species  complexes  (cenospecies  of 
the  experimentalist).  John  Thomas  Howell  (1940a)  "presents  the  case  of  hy- 
bridization between  certain  species  of  the  section  Cerastes.  His  conclusion 
is  worthy  of  note :  "My  general  conclusion  is  that  anyone,  interested  in  the 
taxonomy  and  the  possible  evolution  of  entities  in  the  subgenus  Cerastes  of 
Ceanothus,  cannot  disregard  the  probable  effect  of  hybridization,  and  that 
some  of  our  puzzling  entities  can  be  more  definitel}^  limited  as  taxonomic 
units  with  the  proper  study  and  interpretation  of  intermediates  of  suspected 

[  323  ] 


324 


CALIFORNIA  ACADEMY  OF  SCIENCES        [Proc.  4th  ser. 


hybrid  orijiin."  ^lany  botanists  have  been  reluctant  to  accept  the  importance 
of  hybridization  in  the  evolution  of  the  numerous  forms  of  Ceanothus  and 
some  ignore  it  entirely.  However,  anyone  who  has  done  mue'h  serious  field 
study  of  this  genus  will  come  to  agree  with  Howell's  general  conclusion. 

It  is  the  opinion  of  the  writer  tluit  the  number  of  true  biological  species 
(ecospecies  of  the  experimentalist)  existing  at  a  given  time  in  any  genus  of 
plants  cannot  be  known  until  extensive  experimental  studies  for  testing  the 
fertility  of  hybrids  can  be  carried  out.  Until  more  logical  methods  for  the 
determination  of  true  biological  or  distinct  species  are  set  forth,  botanists 
will  do  well  to  test  out  the  numerous  variants  w'ithin  a  genus  by  putting 
them  through  the  biological  sterility  sieve  advocated  by  Clausen,  Keck,  and 
Hiesey  (1939)  shown  in  table  1. 

Table  1.  Tlir  conci))!  of  sprrirs. 


Degree  (if 
separation 

Hylirids  fertile, 

second  generation 

vigorous 

Hybrids  partially 

sterile, 

second  generation 

weak 

Hybrids  sterile 
or  none 

\       Internal 

\ 
External 

In  different 
environments 

Distinct  subspecies 
ECOTYPES 

Distinct  species 

ElCOSPECIES 

Distinct 

species  cn))iplc.res, 
CENOSPECIES 

In  the  same 
environment 

Local  variations 
of  one  species 

BIOTYPES 

Species  overlapping 

in  common  territory 

(with  hybrid  swarms) 

Experimental  methods  require  much  time  and  labor,  and  it  is  evident  to 
botanists  who  study  woody  plants  that  the  adoption  of  the  experimental 
method  will  be  slow,  and  until  such  adoption  other  methods  now  employed 
by  workers  will  continue  to  be  used.  However,  by  being  aware  of  the  impor- 
tance of  experimental  methods,  field  workers  will  become  more  alert  to  the 
existence  of  natural  hybrids  and  to  geographic  and  ecological  variations. 
By  recognizing  field  hybrids  and  applying  to  them  the  same  principles  used 
by  Clausen,  Keck,  and  Hiesey  in  determining  experimentally  cenospecies, 
ecospecies,  ecotypes,  and  biotypes,  it  has  been  possible  to  gain  much  helpful 
information  in  determining  species  complexes  (cenospecies),  distinct  or  bio- 
logical species  (ecospecies),  subspecies  (ecotypes),  and  local  variations  (bio- 
types) in  the  genus  Ceanothus. 

In  this  genus  there  are  two  rathei-  well-defined  sections  or  sub-genera, 
Euceanothus  with  leaves  alternate  and  fruit  not  horned,  and  Cerastes  with 
typically  opposite  leaves  and  horned  fruit  ( Plate  28) .  In  a  recent  publication, 
by  Van  Rensselaer  and  the  writer  (1942)  the  Euceanothus  section  includes 
thirty-three  binomials  (so-called  species),  seventeen  trinomials  (varieties), 
many  minor  variations,  four  named  natural  hybrids,  and  many  other  hybrids. 


Vol.  XXV]        McMIKN:   FIELD  HTBEIDS  IN    CEAXOTHUS 


325 


The  Cerastes  section  includes  twenty-three  binomials,  nine  trinomials,  many 
minor  variations,  four  named  natural  hybrids,  and  other  hybrids.  Three 
named  natural  hybrids  between  members  of  the  two  sections  are  also  included. 
The  writer  believes  that  the  number  of  existing  distinct  species  (ecospecies) 
is  much  fewer  than  represented  by  the  number  of  binomials  given  in  manuals, 
and  that  more  subspecies  (ecotypes)  and  local  variations  (biotypes)  will  be 
recognized  when  sufficient  knowledge  of  the  behavior  of  the  hybrids  is  known. 
Some  of  this  behavior  can  be  learned  by  careful  observation  in  the  field. 

INTEKSECTIONAL  HYBRIDS 

If  observed  hybrid  plants  fail  to  set  fruit  year  after  year,  one  may  assume 
with  some  degree  of  certainty  that  they  are  sterile,  and  that  their  parents 
belong  to  distinct  species  complexes  (cenospecies).  This  situation  is  partially 

CHART  1 

Wfttural  HTbridB  betireen  Snaciea 
of  the  Sections  Ceraatea  and  Sueeajiothus 


45,  C.  rlgidufl 

43.  C,  purpureus 

t?.     C.  prostratus 

7ar.  1 


61.  X  C.  Vei t  ch  ianus 

T  C.  serrulatua 
b^.  X  C.  rugosus 


ames  Roof- 


-3-^-1 


".  griaeus  25» 

3.  cordulatus  14. 

Tolutlnu3  1. 

C.    sninosiis  11. 

j^Ji^-C.   oli^anthus  21. 

"          C,   8orediatU9  22. 

~-  -  -C.   thyrslflorus  26. 


exemplified  in  the  intersectional  cross  between  C.  prostratus  and  C.  cordulatus 
which  has  been  given  the  designation  x  C.  scrrulatus.  This  hybrid  was  first 
collected  in  1926  from  a  small  colony  of  not  more  than  four  or  five  plants  in 
a  shallow  draw  on  a  ridge  between  Emerald  Bay  and  Cascade  Lake,  Eldorado 
County,  California.  To  my  knowledge  it  has  never  been  found  at  any  other 
locality.  It  appears  that  a  single  hybrid  plant  may  have  been  formed  and  that 
the  other  plants  have  resulted  from  natural  layering.  The  evidence  for  con- 
sidering the  two  parents  of  this  hybrid  as  belonging  to  different  species  com- 
plexes would  be  more  conclusive  if  more  sterile  hybrid  plants  were  known. 
Associated  with  this  hybrid  is  C.  prostratus,  C.  cordulatus,  and  C.  velutinus. 
The  prostrate  habit  and  the  presence  of  sunken  stomatal  pits  on  the  underside 
of  the  leaves  relate  it  to  C.  prostratus,  but  the  predominantly  thinner  and 
alternate  leaves,  the  small  deciduous  stipules,  and  short  racemose  flower  clus- 
ters relate  it  to  C.  cordulatus.  The  presence  of  both  opposite  and  alternate 
leaves  and  the  very  pale  blue  to  nearly  white  flowers  seem  to  be  intermediate 
characters.  The  plants  have  never  been  known  to  set  fruit. 

Two  other  field  hybrids  and  six  garden  hybrids  have  been  observed  by  the 
writer  between  members  of  the  two  sections  (Chart  1).  Only  one  of  these 
hybrids,  the  garden  hybrid  C.  James  Roof,  has  ever  been  known  to  set  fruit. 
The  seeds  of  this  hybrid  when  tested  failed  to  germinate.  The  parents  were 
C.  purpureus  and  probably  one  of  the  following :  C.  oliganthus,  C.  spinosus. 


326  CALIFOBNIA  ACADEMY  OF  SCIENCES        [Proc.  4th  ser. 

C.  thyrsiflorus,  or  C.  sorediatus.  The  two  other  observed  field  hybrids  between 
members  of  the  two  sections  are  x  C.  rugosns  and  x  C.  Veitchianus.  x  Ceano- 
thus  rugosns  appears  to  be  a  hybrid  between  C.  prostratns  and  C.  velntinus. 
Specimens  have  been  obtained  from  two  localities,  one  from  the  lava  beds  near 
Egg  Lake,  Modoc  County,  California,  and  the  other  from  near  Truckee,  Ne- 
vada County,  California.  M.  S.  Baker  and  F.  P.  Nutting  collected  the  speci- 
mens from  INIodoc  County  in  1894.  Baker  in  a  note  states  that  the  plant  was 
a  ''decumbent  form  of  C.  vehitinus — probably  a  hybrid  of  C.  prostratns  and 
C.  vehitinus."  The  specimens  from  Truckee  were  collected  by  C.  F.  Sonne  in 
1890.  Sonne  reported  that  the  plant  sets  flowers  sparingly  and  that  it  does 
not  fruit  at  all.  It  would  seem  then,  that  the  two  parents,  C.  ijrostratus  and 
C.  velntinus,  definitely  belong  to  different  species  complexes. 

X  Ceanothus  Veitchianus  appears  to  be  a  hybrid  between  C.  grisens  and 
probably  C.  rigidus.  To  my  knowledge  it  was  first  collected  by  William  Lobb 
in  California,  probably  near  Monterey  where  C.  grisens  and  C.  rigidus  grow. 
Lobb  probably  collected  seeds  from  C.  grisens  and  C.  rigidus  and  from  this 
collection  appeared  the  hybrid  x  C.  Veitchianus  in  the  Veitch  and  Sons  nurs- 
eries of  Exeter  and  Chelsea,  England.  I  have  seen  no  specimens  collected  from 
the  field.  The  few  garden  hybrids  of  x  C.  Veitchianus  which  I  have  observed 
have  never  set  fruit.  It  would  appear  from  the  sterility  test  that  C.  rigidus 
and  C.  griseus  belong  to  different  species  complexes. 

The  genetic  barrier  between  the  two  sections  must  be  something  more  than 
chromosome  number,  because  the  diploid  number  for  all  species  examined  in 
the  two  sections  by  Nobs  (1941)  is  24  (Plates  29  and  30) .  Nobs  also  examined 
six  hybrid  plants  of  different  parentage  and  all  were  found  to  have  the  same 
number  of  chromosomes  as  the  parents,  namely  2n  =  24. 

INTRASECTIONAL  HYBRIDS 

If  obser\'ed  hybrid  plants  are  partially  sterile  and  set  fruit  which  produce 
seeds  that  germinate  poorly  or  whose  seeds  produce  a  weak  second  generation, 
we  may  assume  that  the  hybrid  plants  were  produced  by  the  crossing  of  dis- 
tinct species  (eeospecies  of  the  experimentalist).  If  the  hybrids  proved  to  be 
fertile  and  the  second  generation  vigorous,  we  may  then  assume  that  their 
parents  belonged  to  distinct  subspecies  (ecotypes  of  the  experimentalist)  or 
that  they  were  local  variations  of  one  species  (biotypes  of  the  experimentalist ) , 
depending  upon  whether  they  occurred  in  different  environments  or  in  the 
same  environment.  Examples  of  such  hybrids  have  been  observed  many  times 
in  the  field  between  members  of  the  same  section  of  the  genus  wherever  tliey 
overlap  in  their  distribution. 

Section  Euceanothus 

In  the  section  Euceanothus,  the  evidence  from  hybridization  points  toward 
the  existence  of  more  cenospecies  and  eeospecies  than  in  the  section  Cerastes. 
In  tliis  section    {Euceanothu.s)   hybrids  liave  been  observed  between  most 


Vol.  XXV]        McMIXX:   FIELD  EYBBIDS   IX    CEAXOTHUS 


327 


of  the  species  that  occupy  the  same  area  and  that  flower  at  the  same  time. 
In  some  cases  where  two  species,  as  C.  cordulatus  and  C.  diversifolius  or  as 
C.  thyrsiflorus  and  C.  incanus,  occur  together  only  an  occasional  hybrid  plant 
is  formed  and  that  plant  usually  appears  to  be  totally  sterile.  In  these  cases 
we  may  assume  that  C.  cordulatus  and  C.  diversifolius  belong  to  different 
species  complexes  (cenospecies)  and  also  that  C.  thyrsiflorus  and  C.  inca- 
nus belong  to  different  species  complexes.  Since  neither  C.  cnrdulatiis  nor 


1.  C.  velutinus 
la.    var.  laevigatu: 

2.  C,  enericanus 

3.  C.  ovatUB 

4.  C.   sangulneaB 

5.  C.   serpyllifoliua 

6.  C.  microphjllus 

7.  C.   integerrimu? 
(riB.   white  or  "D 

8.  C,   parrlfolius.      \ 
(FlB.  ■blue) 

9.  C.  Kartlnil 

10.  C.  Faljneri 

11.  C.   spinosus 
(TIb.  pale  blue)^ 

1?.  C.   1-eucoderiiis 

13.  C.   iBcanus 

14.  C.   cordulatus 

15.  C.  buxlfclius 

16.  C.  rendleri 

17.  C.  dopressua 


CHAET   2 

K„f„rpl   I'yhri.^^..  '?r»---^':"  ^^'■cie-   of  the  Sectlog  F.u:canothu3 

yio/rerg  blu9 
__— C.  arboreua 


,   diTersifoliuj  33» 


C.  diversifolius  are  ever  found  in  nature  growing  with  either  C.  thyrsiflorus 
or  C.  iyicanus,  no  evidence  from  field  hybrids  has  been  obtained  regarding 
their  interrelationships.  From  morphological  characters  it  would  seem  that 
G.  cordulatus  and  C.  incanus  belong  to  the  same  species  complexes,  and  that 
that  complex  is  different  from  the  one  to  which  C.  thyrsiflorus  or  the  one  to 
which  C.  diversifolius  belongs. 

In  the  natural  hybrids  observed  between  C.  cordulatus  and  C.  velutinus 
and  between  C.  thyrsiflorus  and  C.  papillosus  on  the  other  hand,  many  varia- 
tions occur  which  appear  to  be  only  partially  sterile  and  which  appear  to  be 
members  of  a  hybrid  swarm,  thus  indicating  that  probably  these  two  entities 
are  distinct  species  (ecospecies).  If  these  hybrids  were  quite  fertile  and  their 
second  generation  vigorous,  more  local  variations  would  be  expected  than  are 
now  found,  and  then  these  four  entities  would  be  considered  subspecies  (eco- 


328  CALIFOEXIA  ACADEMY   OF  SCIEXCES        [Proc.  4th  ser. 

types)  or  local  variations  of  one  species  (biotypes).  depending  upon  whether 
they  normally  occurred  in  different  environments  or  in  the  same  environment. 
Most  of  the  other  crosses  observed  in  the  Euceanothus  section,  and  recorded 
in  chart  2,  appear  to  follow  the  same  general  patterns  as  given  in  the  examples 
just  cited. 

A  summary  of  the  recorded  natural  hybrids  in  this  section  shows  a  total  of 
thirty,  involving  twenty-one  species.  Ceanothus  thyrsiflorus,  which  in  its  dis- 
tribution contacts  more  species  than  any  other  entity  in  the  Euceanothus 
section,  has  entered  into  nine  crosses,  whereas  the  other  species  with  fewer 
contacts  have  been  involved  in  from  one  to  five  crosses.  Three  of  the  thirty 
crosses  have  been  given  hybrid  binomial  designations  because  they  have  been 
considered  as  species  or  varieties  by  other  workers.  One  other  hybrid  has  been 
named  because  of  its  presence  in  the  nursery-  trade.  Since  much  confusion 
exists  in  the  literature  about  these  hybrid  entities,  it  seems  worth  while  to  give 
a  brief  account  concerning  them. 

Named  Hybrids  of  the  Section  Enccanoihus 

X  Ceanothus  mendocinensis  appears  to  be  a  hybrid  between  C  thyrsiflorus 
and  C.  velutinus  var.  laevigatus.  Plants  of  this  cross  have  been  collected  in 
several  scattered  locations  in  Mendocino  County,  California,  and  in  adjacent 
counties  where  the  two  parents  overlap  in  their  distribution.  They  usually 
occur  as  individual  plants  with  one  or  both  parents,  never  in  colonies.  The 
fertility  of  the  seed  has  not  been  tested,  but  since  so  few  plants  have  been 
found  where  the  parents  occur  together,  and  since  the  hybrids  appear  to  be 
rather  uniform  in  those  characters  which  are  intermediate  between  those  of 
the  parents,  it  would  appear  that  the  plants  were  first-generation  hybrids  and 
that  the  parents  belong  to  distinct  species. 

X  Ceanothus  Lorenzenii  has  been  considered  to  be  a  variety  of  C.  velutinus 
by  some  botanists.  It  occurs  in  the  Sierra  Nevada  of  California  from  Kern 
County  northward  to  Siskiyou  County  and  eastward  in  Nevada.  Apparently 
this  hybrid  is  the  result  of  a  cross  between  C.  velutinus  and  C.  cordulatus 
(Plate  31).  Field  studies  made  about  five  miles  north  of  Markleeville,  Alpine 
County,  California,  in  the  summers  of  1941  and  1942,  furnished  the  most 
convincing  evidence.  On  an  extensive  old  burned-over  area  there  occurred 
C.  velutinus,  C.  cordulatus,  and  C.  prostratus.  The  hybrid  plants  were  found 
only  in  areas  where  C.  velutinus  and  C.  cordulatus  were  closely  associated. 
In  areas  where  one  species  occurred  to  the  exclusion  of  the  other  no  hybrids 
were  observed.  Most  of  the  hybrids  resembled  the  species  C.  velutinus  more 
than  C.  cordulatus.  The  stiff  branches  and  smaller  leaves  without  varnisli 
above,  however,  gave  evidence  of  the  part  C.  cordulatus  had  in  its  origin. 
A  few  plants  were  found  which  resembled  C.  cordulatus  more  than  C.  velu- 
tinus. It  appeared  from  the  field  studies  that  the  hybrid  jilants  were  mostly 
sterile.  If  they  were  quite  fertile  one  would  expect  more  individuals  inter- 
mediate between  the  two  parents  and  the  hybrid  (Histogram  1 ) .  I  believe  most 


Vol.  XXV]        McMINX:   FIELD  HYBRIDS   IN    CEANOTHUS 


329 


of  the  specimens  of  x  C.  Lorerizenii  are  first  generation  hybrids.  Nobs,  after 
field  and  eytological  studies,  also  concluded  that  C.  velutinus  var.  Lorenzenii 
Jepson  is  a  hybrid  between  C.  velutinus  and  0.  cordulatus  (Plates  32  and  33). 
X  Ceanothus  Lohhianus  is  probably  the  result  of  a  cross  between  C.  griseus 
and  C.  dcntatus.  It  was  first  grown  in  England  from  seed  collected  by  William 


no.  of 
plants 


Histogram  1 


0      1      2     3     4     5 
C.  velutinus  Dougl. 


index 
6    7     8    9    10        ^^'^^^ 

C.  cordulotus  Kellogg 


X  C.  Lorenzenii  (Jepson)  McMinn 

Histogram  1  represents  the  distribution  of  jDlants  into  11  groups  based  upon  an  index 
value  determined  by  studying  10  characters  of  C.  cordulatus.  The  plants  were  taken  from 
a  random  sampling  in  an  area  where  C.  cordulatus  and  C.  velutinus  overlapped  in  their  dis- 
tribution. Plants  A\ith  all  10  characters  were  classified  as  C.  cordulatus;  plants  with  none 
of  these  characters  were  classified  as  C.  velutinus;  those  with  4,  5,  and  6  characters  were 
classified  as  x  C.  Lorenzenii;  and  those  with  1,  3,  7,  or  8  characters  are  unnamed  hybrids. 
No  plant  in  the  sampling  was  found  with  2  or  9  characters.  As  the  groups  approach  0  the 
plants  more  closely  resemble  C.  velutinus. 

(x  =  hybrid) 

Lobb  in  California,  probably  in  the  region  of  Monterey  where  C.  griseus  and 
C.  dentatus  occur.  A  few  plants  collected  as  natural  hybrids  in  the  Monterey 
region  and  now  growing  in  California  gardens  approximate  the  type  specimen 


330 


CALIFOEXIA  ACADEMY  OF  SCIENCES        [Proc.  4th  ser. 


of  the  original  C.  Lohhianns.  Seeds  have  been  collected  from  these  hybrids 
but  their  fertility  is  not  known.  Since  so  few  plants  of  this  hybrid  occur  either 
in  the  wild  or  in  any  known  garden  collection  it  would  seem  that  the  hybrid 
is  only  partially  fertile  and  therefore  the  parents,  C.  griseus  and  C.  dentatus, 
belong  to  distinct  species. 

X  Ceanothus  regius  is  a  hybrid  between  C.  ihyrsiforus  and  C  papillosus. 
It  has  been  observed  several  times  wherever  the  two  parents  overlap  in  their 


no.  of 
plants 


Histogram  2 


40- 


0      12      3      4 
0.  papillosus  T  a  G. 


6     7      8     9     10 
C.  thyrsiflorus  Esch, 


index 
value 


X  C.  regius 

Histogram  2  represents  the  distribution  of  plants  into  11  groups  based  upon  an  index 
value  detorniined  by  studying  10  characters  of  C.  tliyi'sifloni.'^.  The  plants  were  counted  from 
a  random  sampling  in  an  area  Avhere  C.  thyrnflorufi  and  C.  papillosus  overlapped  in  their 
distribution.  Plants  with  10  characters  were  classified  as  C.  thyrsiflorus;  those  with  none 
of  these  characters  were  classified  as  C.  j^opillosus ;  those  with  4  or  5  characters  were  classi- 
fied as  X  C.  rcyius ;  and  those  with  2,  6,  or  8  characters  were  classified  as  unnamed  hybrids. 
No  plant  in  this  sampling  was  found  with  1,  3,  7,  or  9  characters. 


distribution  in  San  Mateo  and  Santa  Cruz  counties,  California.  The  hybrid 
give  evidence  of  being  only  partially  sterile,  consequently  the  parents  belong 
to  distinct  species  (Plate  34,  and  Histogram  2). 

Section  Cerastes 

In  the  section  Cerastes  (those  plants  with  typically  opposite  leaves)  there 
appears  to  be  more  evidence  for  a  greater  number  of  subspecies  and  fewer 


I 


Vol.  XXV]        McMINN:   FIELD  HYBRIDS  IN    CEANOTHUS  331 

species  and  species  complexes.  In  this  section,  C.  ciineatus,  C.  Greggii,  and 
C.  prostratus  have  Avider  ranges  of  distribution  than  any  of  the  other  entities 
and  they  have  entered  into  more  crosses  than  any  of  the  others.  Wherever 
C.  cuneatus  and  C.  Greggii  and  its  vars.  vestitus  and  perplexans  overlap  in 
their  distribution,  many  intermediates  occur  which  appear  to  be  fertile.  But 
since  their  degree  of  fertility  is  not  known  it  is  not  possible  to  state  whether 
the  parents  are  in  the  nature  of  subspecies  or  species.  In  the  crosses  observed 
between  C.  cuneatus  and  C.  prostratus  the  hybrid  plants  appear  to  be  partially 
sterile,  wherefore,  their  parents  appear  to  be  distinct  species.  At  the  present 
time  no  localities  are  known  to  the  writer  where  C.  prostratus  and  C.  Greggii 
occur  together.  However,  there  is  evidence  in  the  southern  middle  Sierra 
Nevada  of  California  that  they  may  have  overlapped  in  their  distribution 
sometime  in  the  past.  In  this  area,  particularly  in  Tuolumne  and  Calaveras 
counties,  entities  occur  which  seem  to  possess  characters  originating  from 
both  species.  Two  of  these  entities  appear  in  some  manuals  as  species  binomials 
and  are  worthy  of  further  consideration. 

The  apparently  fertile  entity  known  as  C.  fresnensis  (Fresno  Mat),  occurs 
on  the  western  slope  of  the  Sierra  Nevada  between  3000  and  6500  feet  eleva- 
tion, from  Fresno  County  northward  to  Tuolumne  County  or  probably  to 
Eldorado  and  Placer  counties,  mostly  at  the  southern  extension  of  the  range 
of  C.  prostratus.  In  1939  the  writer  gave  the  distribution  of  this  rather  baffling 
entity  as  extending  from  Fresno  County  northward  to  Plumas  County.  How- 
ever, after  becoming  more  aware  of  the  presence  of  hybrids  in  this  genus, 
extensive  field  studies  were  carried  on  during  the  summers  of  1940,  1941,  and 
1942,  and  I  am  now  of  the  opinion  that  some  of  the  herbarium  specimens 
studied  from  the  more  northerly  localities  had  been  taken  from  hybrid  plants 
of  C  cuneatus  x  C.  prostratus  and  its  variety  Jaxus,  and  from  C.  arcuatus. 
Ceanothus  fresnensis  is  entirely  prostrate,  while  C.  arcuatus  and  most  hybrids 
between  C.  cuneatus  and  C.  prostratus  are  usually  intermediate  between  the 
two  in  habit  of  growth.  That  C.  Greggii  var.  vestitus  has  entered  into  the 
origin  of  C.  fresnensis  is  evidenced  by  the  character  of  the  pubescence  and 
the  nature  of  the  fruit.  Ceanothus  Greggii  var.  vestitus  now  occurs  in  Kern, 
Inyo,  and  Mono  counties  and  in  the  past  may  have  overlapped  in  its  distribu- 
tion with  C.  prostratus.  John  Thomas  Howell  (1940)  also  considers  the  prob- 
ability that  C.  fresnensis  "is  descended  from  the  offspring  of  a  fertile  cross 
between  C.  prostratus  and  C.  vestitus"  (C.  Greggii  Yar.  vestitus).  If  we  accept 
an  early  hybrid  origin  of  C.  fresnensis,  then  C.  prostrat\is  and  G.  Greggii  var. 
vestitus  must  have  belonged  to  distinct  subspecies. 

The  other  apparent  fertile  entity  previously  referred  to  is  C.  arcuatus.  It 
occurs  in  scattered  localities  from  about  3000  to  7600  feet  elevation  on  the 
western  slope  of  the  Sierra  Nevada  of  California  from  Madera  County  north- 
ward to  Plumas  County.  Near  the  summit  on  the  south  slope  of  Robb's  Peak, 
Eldorado  County,  elevation  6500  to  6700  feet,  it  is  one  of  the  dominant  shrubs 
associated  with  Arctostaphylos  patula  and  Ceanothus  prostratus.  A  few  pros- 


332  CALIFORNIA  ACADEMY  OF  SCIENCES        [Proc.  4th  ser. 

trate  plants  Avliieli  appear  to  be  Ceanothus  fresnensis  also  occur  in  this  asso- 
ciation. These  may  be  hybrids  of  C.  prostratus  and  C  arcuatus. 

Ceanothus  arcuatus  seems  to  have  segregated  out  of  a  hybrid  population 
formed  by  the  crossing  of  C.  cuneatus  with  C.  fresnensis.  The  habit  of  growth, 
character  of  fruit,  and  color  of  flowers  are  intermediate  between  the  two 
probable  parents.  The  gi-ayish  bark  resembles  that  of  C.  cuneatus  while  the 
leaves  resemble  more  those  of  C.  fresnensis.  In  the  Sugar  Pine  Ridge  region 
along  Highway  108,  Tuolumne  County,  occur  plants  of  C.  cuneatus  and 
C.  fresne7isis  as  well  as  hj^brids  between  these  two  entities.  At  higher  elevations 
along  Highway  108  the  hybrid  form  alone  occurs.  In  Eldorado,  Nevada, 
Placer,  and  Plumas  counties,  plants  scarcely  distinguishable  from  these  hy- 
brids occur  in  the  Upper  Transition  and  Canadian  Life  Zones  far  above  the 
distribution  of  C.  cuneatus.  Since  these  plants  are  scarcely  distinguishable 
from  the  hybrids  of  Tuolumne  County  and  since  they  occur  in  a  different 
geographical  area  in  the  absence  of  both  C.  fresnensis  and  C.  cuneatus,  I 
believe  there  are  sufficient  reasons  for  regarding  them  as  belonging  to  a  dis- 
tinct subspecies  or  probably  to  a  distinct  species.  It  seems  to  me  we  have  here 
a  subspecies  or  a  species  which  has  "just  arrived"  and  is  now  migrating  north- 
Avard  from  its  center  of  hybrid  origin. 

The  group  of  entities  including  C.  pinetoruni,  C.  postratus  and  its  vars. 
laxns  and  occidentulis,  C.  pmnilus,  C.  confusus,  C.  divergens,  and  C.  purpii- 
reus  has  so  many  forms  which  appear  to  grade  into  each  other  that  the  entities 
may  well  exist  as  subspecies  of  one  large  polymorphic  species.  These  entities 
are  not  only  very  closely  related  to  each  other  but  also  to  another  smaller 
group  composed  of  C.  rigidus,  C.  Masonii,  C.  gloriosus  and  its  var.  exaltatus. 
The  two  groups  differ  slightly  from  one  another  in  the  character  of  the  leaf 
margins.  The  former  normally  has  leaves  with  serrate  or  spinulose  margins, 
and  the  latter  has  leaves  with  dentate  to  quite  entire  margins.  In  the  foothills 
of  the  Hood  Mountain  Range  in  Sonoma  County,  California,  members  of  these 
two  groups  overlap  in  their  distribution  and  have  produced  numerous  hybrids, 
many  of  which  appear  to  be  fertile.  Since  the  barrier  between  most  of  these 
entities  appears  to  be  that  of  geographic  or  ecologic  isolation  rather  than 
genetic,  it  would  seem  that  the  entities  are  probably  in  the  nature  of  subspe- 
cies. They  seem  to  hybridize  freely  wherever  they  meet,  but  appear  quite 
distinct  in  their  own  geographic  ranges. 

Ceanothus  sonomensis  of  the  Hood  Mountain  Range,  Sonoma  County,  Cali- 
fornia, may  be  the  result  of  the  crossing  of  one  of  these  hybrids  with  C.  cunea- 
tus, which  occurs  in  the  same  area.  This  unit  now  occupies  a  small  area 
somewhat  isolated  from  the  other  forms  o^  Ceanothus  growing  in  that  general 
region.  When  C.  cuneatus  is  brought  into  the  picture  it  brings  in  another 
series  of  forms  which  seems  to  hybridize  with  the  others,  and  until  experi- 
mental methods  are  used  it  would  be  unwise  to  make  predictions  as  to  the 
exact  nature  of  the  entities.  That  C.  sonomensis,  C.  arcuatus,  and  several  other 
somewhat  similar  entities  in  the  section  Cerastes  might  be  the  results  of  inter- 


Vol.  XXV]        McMINN:  FIELD  HYBRIDS  IN    CEANOTEUS 


333 


specific  hybridization  cannot  be  entirely  ruled  out.  G.  Ledyard  Stebbins 
(1942a)  lias  pointed  out  in  his  paper  on  the  "Role  of  Isolation  in  the  Differen- 
tiation of  Plant  Species"  that  occasionally  recombinations  should  occur  in 
the  progeny  of  a  partially  sterile  interspecific  hybrid  involving  "small  units 
genetically  independent  of  morphological  differences."  He  further  states  that 
"Some  of  these  new  recombinations  could  be  fertile  types  which  would  be 
different  in  appearance  from  either  of  their  original  parents  and  exhibit 
partial  intersterility  with  both  of  them.  They  would  thus  become  the  progeni- 
tors of  new  species." 

CHAET  3 
Katviral  Hybrlda  'betwcBn  gpeclea  of  the  Section  Cerastes 
FlowerB  white  Flowers  blue 

34.  C.  verrucosus -> — -> > —   ^.  ri(;idu3  ''.5, 

35.  C.   insularia  ,C.  Masonll  46. 
26.      C.   megacarpus • 
07.     C.   crassifollus^ 


/    p.  gloriosus         47. 
/  -^      var.   exaltftua 


38.  C.    Oreggii 

yar.   vestltus 
yar.  perplexano 

39.  C.    lanuginosus^ 

40.  C.    cuneatu 

41.  C.    Ferrisae  —   -^ 
yiowere  tlua  cr  lavender  ^ 

4S.      C.    arcuatu; 

43.  C.    fresnensls'^  1 

■?  -»(t.   Vision 

44.  C.      T-«nni1nel1c y 


A  summary  of  the  recognized  natural  hybrids  in  the  section  Cerastes,  as 
given  in  the  book  Ceanothus,  gives  a  total  of  twenty-one,  of  which  four  are 
named  and  seventeen  unnamed  (Chart  3).  The  four  named  hybrids  represent 
binomials  which  have  appeared  as  distinct  species  in  some  publications.  Eight 
so-called  species  of  this  section  have  entered  into  the  hybrid  complex.  Ceano- 
thus cuneatus,  C.  prostratus,  and  C.  Greggii  are  the  most  widely  distributed 
named  entities  and  they  have  entered  into  crosses  more  often  than  any  of  the 
other  named  entities.  Ceanothus  verrucosus,  a  species  with  relatively  few 
variations,  to  my  knowledge  has  not  entered  into  any  natural  cross.  This 
species  has  a  rather  limited  range  of  distribution  in  San  Diego  County,  Cali- 
fornia, which  does  not  overlap  with  the  distributional  ranges  of  other  species 
in  the  same  section.  According  to  the  concept  of  genetic  homogeneity  as 
reviewed  by  Stebbins  (1942),  we  might  consider  that  C.  verrucosus,  because 
of  its  limited  distribution,  is  genetically  homogeneous  and  that  it  consists  of 
relatively  few  biotypes  which  in  turn  are  relatively  homozygous.  On  the  other 
hand  we  might  consider  C.  cuneatus,  C.  prostratus,  and  C.  Greggii,  siuce  they 


334  CALIFORNIA  ACADEMY  OF  SCIENCES        [Proc.  4th  ser. 

are  widespread  species,  as  being  heterogeneous  in  their  genetic  make  up.  This 

heterogeneity  is  well  exemplified  by  the  numerous  biotypes  which  occur  in 

these  species. 

Named  Hybrids  of  the  Section  Cerastes 

X  Ceanothus  otayensis  in  known  only  from  Otay  Mountain  in  San  Diego 
County,  California,  and  is  here  considered  to  have  originated  as  the  result  of 
a  cross  between  C.  crassifolius  and  C.  Greggii  var.  perplexans.  Both  of  these 
entities  occur  together  on  Otay  Mountain.  The  hj^brid  plants  bear  good  fruit 
and  the  seeds  appear  to  be  fertile.  If  they  are  found  to  produce  strong  second 
generation  plants,  then  the  two  parents,  C.  ci-assifolius  and  C.  Greggii  var. 
perplexans,  must  be  considered  to  belong  to  different  subspecies.  One  evidence 
for  their  fertility  is  the  amount  of  variability  which  occurs  among  the  hybrid 
plants. 

The  name  x  Ceanothus  flexilis  has  been  given  to  those  hybrid  plants  occur- 
ring in  northern  Shasta,  Lassen,  and  southwestern  Modoc  counties  of  Cali- 
fornia. In  these  localities  C.  cuneatus  and  C.  prostr-at^is  var.  laxus  overlap  in 
their  distribution  and  they  are  considered  to  be  the  parents  of  x  C.  flexilis. 
The  habit  of  growth,  many  leaf  characters,  size  of  fruit,  and  flower  color  are 
usually  intermediate  between  the  two  parents.  The  fertility  of  the  hybrids 
is  not  known,  but  from  field  observations  I  would  suspect  partial  sterility. 
If  the  suspicion  is  proved  to  be  a  fact  by  experimental  methods,  then  the 
parents,  C.  cuneatus  and  C.  prostratus,  belong  to  distinct  species. 

X  Ceanothus  Bakeri  is  another  named  hybrid  which  appears  to  have  re- 
sulted from  a  cross  between  C.  prostratus  var.  laxus  and  either  C.  Greggii 
var.  vest  it  us  or  one  of  the  many  varieties  of  C.  cuneatus.  The  single  plant 
known  to  me  of  this  hybrid  was  collected  by  C.  F.  Baker  from  Kings  Canyon, 
Ormsby  County,  Nevada,  in  1902.  The  three  fragmentary  specimens  resemble 
somewhat  specimens  of  x  C.  flexilis  and  x  C.  connivens,  both  of  which  prob- 
ably have  genes  of  C.  prostratus  in  their  makeup.  This  small  bit  of  evidence 
would  favor  the  assumption  that  C.  prostratus  and  C.  Greggii  or  C.  cuneatus 
belong  to  distinct  species. 

Specimens  representing  the  named  hybrid  x  C.  connivens  were  collected 
from  near  Sheep  Ranch,  Calaveras  County,  California,  by  E.  L.  Greene  and 
by  T.  S.  Brandegee.  Field  studies  in  the  Calaveras-Sheep  Ranch  area  in 
1941  failed  to  disclose  any  specimens  which  quite  resembled  the  specimens 
of  the  earlier  collectors.  However,  in  the  Calaveras-Tuolumne  area,  where 
C.  cuneatus,  C.  prostratus,  and  C.  fresnensis  grow,  many  h.ybrid  jilants  occur, 
some  of  which  appear  much  like  x  C.  connivens.  It  is  probable  that  x  C.  con- 
nivens represents  a  cross  between  C.  cuneatus  and  either  C.  prostratus  or 
C.  fresnensis. 

In  concluding  this  paper  mention  should  be  made  of  the  numerous  other  hy- 
brid plants  which  occur  between  C.  cuneatus  and  C.  jrrostratus  wherever  these 
two  species  occupy  the  same  local  area  (Plates  35  and  86  and  Histogram  3). 
Ceanothus  cuneatus  is  primarily  a  component  of  hard  chaparral  in  the  Upper 


Vol.  XXV]        McMIXX:   FIELD  HYBRIDS  IX    CEAXOTHUS 


335 


Sonoran  Life  Zone,  while  C.  prostratus  occurs  chiefly  in  the  open  pine  forests 
of  the  Transition  and  Lower  Canadian  Life  Zones.  However,  along  the  none- 
too-well-defined  borders  of  these  life  zones  these  two  entities  overlap  in  their 
distribution  and  also  in  their  characteristics.  Some  of  the  hybrids  observed 
appear  to  have  good  fruits  and  seeds ;  whether  the  seed  produce  strong  and 


no.  of 
plants 


Histogram  3 


40- 


0  '    I   ■  2  ■  3  ■  4 
C.  prostratus  Benth, 


6  ■  7     8 

C.  cuneatus  Nutt. 


index 
value 


Hybrids 


Histogram  3  represents  the  distribution  of  plants  into  9  groups  based  upon  an  index 
value  determined  by  studying  8  characters  of  C.  cuneatus.  The  plants  were  taken  from  a 
random  sampling  in  an  area  where  C.  cuneatus  and  C.  prostratus  overlapped  in  their  dis- 
tribution. Plants  with  8  characters  Avere  classified  as  C.  cuneatus;  plants  with  none  of  these 
characters  were  classified  as  C.  prostratus;  and  those  with  1,  2,  3,  4,  5,  6,  or  7  characters 
were  classified  as  unnamed  hybrids.  Plants  with  a  decreasing  number  of  C.  cuneatus  charac- 
ters possessed  an  increasing  number  of  C.  prosti-atus  ones.  ^ 


fertile  offspring  is  not  known.  Since  the  hybrid  forms  do  not  vary  as  much  as 
F,  and  F3  offspring  usually  do  from  heterozygous  parents,  and  since  there 
are  relatively  few  individuals  compared  to  the  number  of  individuals  of  both 
parents,  I  believe  the  hybrids  are  partially  sterile  and  that  their  parents  are 
distinct  species. 


336  CALIFORNIA  ACADEMY  OF  SCIENCES        [Proc.  4th  ser.     i 

LITERATURE  CITED 
Brandegee,  K. 

1894,    Studies  in  Ceanothus.  Proceedings  California  Academy  of  Sciences  (2),  4 :  173-222. 

Clausen,  J.  C,  D.  D.  Keck,  and  W.  M.  Hiesey 

1939,  The  Concept  of  Species  Based  on  Experiment.  American  Journal  of  Botany,  26 : 

] 03-106. 

HowEi.i..  J.  T. 

1940,  Studies  in  Ceanothus.  Leaflets  of  "Western  Botany,  2 :232. 
1940a,  Studies  in  Ceanotlius.  Leaflets  of  Western  Botany,  2 :  259-262. 

McMiNN,  H.  E. 

1939,    An  Illustrated  Manual  of  California  Shrubs.  307-309.  J.  W.  Stacey,  Inc.,  San 
Francisco. 

Nobs,  M.  A. 

1941,  Personal  Communications. 

Stebbins,  G.  L. 

1942,  The  Genetic  Approach  to  Problems  of  Eare  and  Endemic  Species.  Madrono,  6: 
241-258. 

1942a,  The  Eole  of  Isolation  in  the  Differentiation  of  Plant  Species.  Biological  Symposia, 
6:217-232. 

VAX  Eensselaer,  M.  and  H.  E.  McMinn 

1942,    Ceanothus.  Santa  Barbara  Botanic  Garden,  Santa  Barbara. 


I  wisli  to  express  my  sincere  appreciation  to  Mr.  M.  A.  Nobs  and  Mr.  John  Poindextei 
for  f urjiishing  the  data  and  photographs  for  plates  29  to  36  and  text  figures  1  to  3. 


EXPLANATION  OF  PLATES 


PLATE  28 
Section  Euceanotltus 

A.  Branchlet  with  alternate  leaves,  thin,  early  falling  stipules,  and  unihel-like  clusters 
of  flowers  in  racemes. 

B.  Fruit  without  apical  horns. 

C.  Cross  section  of  leaf  showing  stomata  protected  by  guard  cells  on  lower  surface. 

Section  Cerastes 

D.  Branchlet  with  opposite  leaves,  thick,  corky,  persistent  stipules,  and  umbel-like 
fruit -cluster. 

E.  Fruit  udth  apical  horns. 

F.  Cross  section  of  leaf  showing  stomata  in  sunken  pits  on  the  lower  surface. 


[338  I 


PROC.  CALIF.  ACAD.  SCI..  4TH  SERIES.  VOL.  XXV,    NO.    14  [McMINN]   PLATE    28 


[  339  ] 


PLATE  29 

Section  Euceanothus 
Fig.  1.  C.  arboreus.  Fig.  10.  C.  Parri/i. 

Fig.  2.  C.  cordulatu.s.  Fig.  11.  C.  Parryi. 

Fig.  3.  C.  cyaneus.  Fig.  12.  C.  thijrsiflonis. 

Fig.  4.  C.  foliosus.  Fig.  13.  C.  Ihymiflorus. 

Fig.  5.  C  incanu^.  Fig.  14.  C.  tomentosus. 

Fig.  6.  C  integerrimus  var.  californicus.      Fig.  15.  C.  velutiniis. 
Fig.  7.  C.  Lemmonii.  Fig.  16.  C.  veluti)iu.s  var.  laevigatus. 

Fig.  8.  C  leucodermis.  Fig.  17.  C.  spinosus. 

Fig.  9.  C.  papillosus. 


i 


[  340  ] 


PROC     CALIF,  ACAD.  SCI..  4TH  SERIES.   VOL.  XXV.   NO.    14 


[MCMINN]   PLATE   29 


^1 

Fig.    1 


Fig.    ^ 


Fig.    12 


Fig.    2 


Fig.    7 


4^- 

Fi£^.    13 


Fig.    3 


Fig.    8 


Fig.   9 


Fig.   10 


Fig.    5 


Fig.    11 


Fig.    14 


1^ 

Fig.    15 


Fig.   16 


Fig,    17 


[  341  ] 


PLATE  3U 

Section  Cerasies 


Fig.  1.  C.  proNfratiis  var.  occidentalis. 

Fig.  2.  C.  cuneatus. 

Fig.  3.  C.  cnneatus. 

Fig.  4.  C.  cuneatus. 

Fig.  5.  C.  cuneatus. 

Fig.  6.  C.  Ferrisae. 

Fig.  7.  C.  gloriosus. 


Fig.    8.  C  prost rains  var.  laxus. 

Fig.    9.  C.  prostratus  var.  laxus. 

Fig.  1(1.  C  pro.strafus. 

Fig.  11.  (7.  ramulosus. 

Fig.  12.  C.  rigidus. 

Fig.  13.  C.  rigidus. 

Fig.  14.  C.  rerr«co.s«.s. 


342 


PROC.   CALIF.  ACAD.  SCI    .  4TH  SERIES,  VOL.  XXV.   NO.    14 


[MCMINN]   PLATE    30 


4^ 


Fig.   2 


Fig.    6 


Fie.  11 


Fig.    7 


Fig.   12 


Fig.   3 


Fig.    8 


.0, 


Fig,   13 


Fig.    5 


Fig.   9 


^' 


Fig.    14 


Fig.  4 


Fig.   10 


[  343  ] 


PLATE  31 


Diagrams  of  the  growth  forms  of  C.  velutinus,  C.  cordulatus, 
and  the  hybrid  x  C.  Lorensenii. 


[344] 


r 


PROC.   CALIF.   ACAD.  SCI..  4TH  SERIES.   VOL.  XXV,    NO.    14 


[McMINNJ   PLATE    31 


X  C.Lorenzenii 
(  hybrid) 


£•    coraxxlatus 


[345] 


PLATE  32 


Photographs  of  specimens  of  C.  velutinus,  C.  cordulatus,  and  a  graded  series  of  five 
of  their  hybrids  taken  from  an  area  where  the  two  species  occur  together. 


[  346  ] 


J 


N 

<0 

UJ 
h 
< 

J 

a. 

i—i 

z 
z 

i 

u 

S 


0 

z 
> 

X 
X 

J 

o 
> 

lli 
i 

I 
I- 


u 

d 
< 
u 
< 


_i 
< 
u 

u 

0 
o: 


PLATE  33 

Chromosomes  of  C.  cordidatus,  C.  velutinus,  and  of  the  hybrid  x  C.  Loremenii. 
C.  corduJatus,  Nobs  76,  Snow  Flat,  elevation  4500  feet,  Nevada  County,  California  ; 
12  regular  pairs.  C.  veluti7ius,  Nobs  79,  Agate  Bay,  Lake  Tahoc,  Placer  County,  Cali- 
fornia; 12  regular  pairs,  x  C.  Lorenzenii,  Nobs  117,  Observation  Point,  Lake  Tahoe, 
Eldorado  County,  California;  12  regular  pairs,  x  C.  Lorenzenii,  Nobs  118,  Observation 
Point,  Lake  Tahoe,  Eldorado  County,  California ;  12  regular  pairs,  one  very  small 
pair,  and  3  fragments. 


348 


PROC.  CALIF.  ACAD.  SCI..   4TH  SERIES.  VOL.   XXV.   NO.   14  [McMINN]   PLATE    33 


ui::ijfiut 


C,  cordulatus 


No,   117 


uiaiiutii 


X  C.   Lorenzenii 


No.   118 


ittmiim-' 


C.   velutinus 


««Ji:::i:i 


[349] 


PLATE  34 

Photographs  of  specimens  of  C.  thyrsiflorus,  C.  papillosiis,  and  a  graded  series  of  five 
of  their  hybrids  taken  from  an  area  where  the  two  species  occur  togetlier. 


[  350  ] 


Ui 

H 
< 

Q. 

I— 1 

Z 
Z 

i 

u 

2 


O 

z 
> 

X 
X 

J 

o 
> 

ui 

hi 

5 

LlI 
0) 

I 


U 

d 
< 

u 
< 


< 

u 

u 
o 

a. 


'      / 


PLATE  35 

Photographs  of  specimens  of  C.  prostratus,  C.  cuneatus,  and  a  graded  series  of  four  of 
their  hybrids  taken  from  an  area  where  the  two  species  occur  together. 


[352] 


10 

(n 

UJ 

H 
< 

_J 

0. 

I — 1 

z 
z 

s 

o 

5 


o 

z 

> 

X 
X 


o 
> 

en 

q: 

UJ 

tn 

I 
I- 


u 

d 
< 
u 
< 


< 
u 

u 
o 

q: 
a. 


PLATE  36 

Diagrams  of  the  growth  forms  of  C.  prostratus,  C.  cuneatus,  and 
one  of  their  hybrids. 


[354] 


ID 

n 
u 

< 

J 

D. 

I — I 
Z 
Z 

u 

S 


o 

z 

> 

X 
X 

J 
0 

> 

ui 

E 
111 

U) 

I 

H 


u 

d 
< 

u 
< 


_i 
< 
u 

d 

0 

(T 
Q. 


-^^ 


O 

c 
u 


3 

a 

O 

a 

^1 


I 


« 


PROCEEDINGS  ^^ 

OF  THE  V 

CALIFORNIA  ACADEMY  OF  SCIENCES      ^ 

Foil  nil  Series 
Vol.  XXV,  No.  15,  pp.  357-376  December  27,  1944 


ALICE  EASTWOOD 
SEMI-CENTENNIAL  PUBLICATIONS 

No.  15 
A  REVISION  OF  PHACELIA  SECTION  MILTITZIA 

BY 

JOHN  THOMAS  HOWELL 

Assistant  Curator,  Department  of  Botany 
California  Academy  of  Scie?ices 

INTRODUCTION 

THE  SPECIES  which  are  here  presented  as  the  section  MUtitzia  of  the  genus 
Phacelia  have  formerly  been  treated  either  as  constituting  the  genus  MU- 
titzia A.DC.  or  as  forming  the  more  numerous  part  of  the  genus  Emmenanthe 
Benth.  {Emmenanthe  §  MUtitzia  A.  Gray).  I  have  recently  expressed  my 
belief  that  there  is  definitely  a  generic  break  between  MUtitzia  and  Emmenan- 
the but  that  MUtitzia  exhibits  both  morphologically  and  habitally  a  relation- 
ship so  close  to  Phacelia  sect.  Microgenetes  that  it  should  be  treated  as  a  section 
in  Phacelia  (1944,  pp.  12-16).  The  plump,  corrugated  seeds  found  in  sect. 
MUtitzia  are  like  those  in  sect.  Microgenetes,  while  the  compressed,  reticulate 
seeds  of  Emmenanthe  are  different  from  any  in  Phacelia.  Moreover  the  lateral 
attachment  of  the  ovules  to  fleshy  placentae  in  MUtitzia  is  characteristic  of 
Phacelia,  while  in  E^nmenanthe  the  pendent  ovules  are  basally  attached  to  the 
wing-like  margins  of  membranous  placentae.  The  marcescent  persistent  co- 
rolla, by  which  the  species  of  sect.  MUtitzia  may  be  distinguished  from  those 
of  sect.  Microgenetes,  was  the  character  by  which  MUtitzia  was  separated 
from  Phacelia  although  the  character  was  not  finally  diagnostic  since  several 
species  in  Phacelia  sect.  Eutoca  also  have  persistent  corollas.  Although  it  is 
this  corolla  character  which  MUtitzia  and  Emm.enanthe  have  in  common,  all 
other  structural  characters  of  MUtitzia  are  entirely  Phacelia-like  and  indicate 
a  relationship  to  Phacelia  that  must  be  interpreted  as  sectional  rather  than 
generic. 

The  section  MUtitzia  consists  of  only  nine  species  and  these  are  confined  to 
the  desert  areas  of  the  western  United  States  between  the  Rocky  Mountains 

.     [ 357  ] 


M.-si  CALIFOIiXIA  ACADEMY   OF  SCIENCES        [Proc.  4th  Ser. 

un  the  east  and  the  Sierra  Nevada  and  Cascade  Range  on  the  west,  extending- 
northward  to  latitude  45°  N.  or  thereabouts  and  southward  to  about  35°  N. 
In  this  relatively  limited  area,  the  northern  part  of  Nevada  is  not  only  the 
geographic  center  for  the  distribution  of  the  species,  but  is  also  the  center  for 
species  concentration.  Here  are  found  seven  of  the  nine  species,  while  in  adja- 
cent southeastern  Oregon  are  five  and  in  eastern  California  four.  To  the 
eastward  not  more  than  two  species  are  found  in  any  one  state.  Three  of  the 
nine  species  are  relatively  common  and  widespread,  P.  adenophora,  P.  scopu- 
lina,  and  P.  tetramera;  two  are  more  localized  in  distribution  and  less  common, 
P.  inundata  and  P.  lutea;  four  are  to  be  counted  among-  the  rarest  of  western 
plants:  P.  glaherrima,  knowai  only  from  three  collections;  P.  inyocnsis  and 
P.  suhmutica  known  from  a  few  collections  from  restricted  areas;  and  P. 
salina,  known  from  five  stations  in  three  different  states. 

Even  to  the  present,  there  has  been  considerable  confusion  and  not  a  little 
difficult}'  in  limiting  species,  part  of  this  being  due  without  doubt  to  the 
meagerness  of  collections  available  for  study.  A  critical  examination  of  numer- 
ous specimens  assembled  from  various  collections  and  institutions  has  revealed 
a  series  of  mostly  clear-cut  and  readily  defined  entities.  Habit,  foliage,  and 
vesture  have  been  found  quite  variable  within  species  so  that  plants  unlike  in 
aspect  are  sometimes  referable  to  the  same  species  because  of  the  critical  but 
dependable  characters  in  flowers  and  fruits.  Because  so  little  is  known  about 
the  field  occurrence  of  the  subspeeific  variants,  only  a  single  variety  has  been 
recognized  taxonomically,  but  undoubtedly  others  will  be  recognized  as  the 
variants  become  better  known.  Phacelia  adenophora,  P.  lutca,  and  P.  scopulina 
are  the  only  species  presenting  facies  so  polymorphic  that  they  may  be  some- 
times confused  with  related  species.  The  first,  M'hich  may  be  considered  the 
most  primitive  species  in  the  group  because  of  its  larger  flower-parts  and 
hairy  corolla-tube  and  filaments,  is  sometimes  difficult  to  distinguish  from  P. 
Jiitea  aiid  P.  inundata.  Phacelia  scopidina,  the  most  widely  distributed  species 
in  the  section,  is  also  the  most  variable ;  to  the  west  there  are  intergrades  wnth 
P.  lutea,  in  the  southeast  P.  suhmutica  is  very  closely  related,  and  P.  salina 
seems  to  be  merely  an  ecologically  specialized  variant  with  smaller  parts. 

From  a  consideration  of  the  environments  in  which  these  phacelias  are 
found,  it  is  very  apparent  that  speeiation  in  the  group  has  produced  entities 
which  occupy  several  of  the  most  highly  specialized  situations  in  our  Avestern 
deserts.  Thus  P.  inundata  is  nearly  confined  to  play  as  of  evanescent  ponds 
and  shrunken  lakes.  Phacelia  tetramera  and  P.  salina  are  commonly  restricted 
to  highly  saline  flats.  Although  P.  scopulina  is  generally  rather  common  in 
sandy  or  gravelly  soil  of  juniper  slopes  or  sagebrush  flats,  it,  too,  sometimes 
occurs  locally  in  more  alkaline  situations.  Phacelia  adenophora  grows  on  sage- 
brush slopes,  but  in  certain  forms  it  is  characteristic  of  clay  flats  of  high 
plains.  The  closely  related  P.  lutca  seems  to  be  characteristic  of  wash  slopes 
of  clay  hills  and  valleys  of  eastern  Oi-egon;  and  the  highly  localized  P.  inyo- 
ensis  seems  to  be  confined  to  boggy  spots  among  granitic  sands  bordoi-ing  the 


Vol.  XXV]  HOWELL:  PEACELIA  SECTION  MILTITZIA  359 

western  Sierra.  In  no  other  group  of  like  size  in  the  genus  are  there  so  many 
forms  edaphically  specialized  and  localized,  rare  creations  to  thrill  the  soul 
of  the  botanical  traveler  who  might  chance  upon  one  flourishing  amid  the  vast 
loneliness  and  seeming  desolate  sterility  of  a  saline  desert. 

ACKNOWLEDGMENTS 

This  revision  has  been  prepared  in  the  Herbarium  of  the  California  Acad- 
emy of  Sciences  (CAS),  and,  although  the  collection  there  has  furnished  the 
basis  for  research,  my  study  would  have  been  far  from  complete  if  I  had  not 
been  able  to  borrow  abundant  collections  from  other  institutions.  These  insti- 
tutions are  here  listed,  together  with  the  symbol  used  in  citing  specimens : 

Chicago  Natural  History  Museum  (CM) 

Dudley  Herbarium,  Stanford  University  (DS) 

Gray  Herbarium,  Harvard  University  (G) 

Intermountain  Herbarium,  Utah  State  Agricultural  College  (lU) 

Missouri  Botanical  Garden  (M) 

Greene  Herbarium,  University  of  Notre  Dame  (ND) 

New  York  Botanical  Garden  (NY) 

Pomona  College  (P) 

Eocky  Mountain  Herbarium,  University  of  "Wyoming  (EM) 

University  of  California  (UC) 

United  States  National  Herbarium  (US) 

Willamette  University  (W) 

Washington  State  College  (WS) 

For  the  privilege  of  borrowing  specimens,  I  am  very  grateful;  and  again  I 
wish  to  express  my  gratitude  and  appreciation  to  many  friends  who  have 
furnished  me  with  field  data  and  specimens  in  this  group  over  a  period  of 
years.  Especially  am  I  grateful  to  H.  D.  Kipley  and  R.  C.  Barneby  for  a  com- 
plete set  of  their  numerous  excellent  collections  which  they  have  obtained 
during  the  course  of  their  critical  field  work  in  western  American  deserts 
from  1940  to  1944.  The  number  of  specimens  cited  from  their  collections  in 
the  present  revision  is  indicative  of  the  important  contribution  they  have 
made  to  my  study. 

Also  it  should  be  noted  that  because  type  specimens  in  the  United  States 
National  Herbarium  and  in  the  Herbarium  of  the  New  York  Botanical  Garden 
are  not  available  for  study  during  wartime,  some  collections  that  are  undoubt- 
edly in  those  institutions  are  not  cited  in  this  paper. 

TAXONOMY 

Phacelia  sect.  Miltitzia  (A.  DC.)  J.  T.  Howell,  Leafl.  West.  Bot.  4 :  15  (IdU) .  Miltitzia 
A.  DC.  in  DC.  Prodr.  9:  296  (1845).  Emmenanthe  §  Miltitzia  (A.  DC.)  A.  Gray,  Proe. 
Amer.  Acad.  10:  328  (1875). 

Flowers  5-merous  or  occasionally  4-merous;  corolla  mareescent,  yellow  or 
frequently  tinged  with  lavender  or  violet ;  corolla-scales  present  or  lacking ; 
filaments  glabrous  except  in  one  species;  ovary  generally  more  or  less  pubes- 


360  CALIFORNIA   ACADEMY   OF  SCIENCES        [Proc.  4th  Ser. 

cent,  the  style  pubescent  at  least  at  base  except  in  one  species,  ovules  4-26, 
laterally  attached  to  fleshy  placentae ;  capsule  apiculate  or  muticous ;  seeds 
transversely  corrugated. — Annual  herbs  of  saline  or  subsaline  habitats  in 
western  North  American  deserts;  herbage  frequentlj^  fleshy,  more  or  less 
pubescent  and  glandular,  occasionally  glabrous;  stems  branching  and  gen- 
erally widely  spreading ;  leaves  entire  to  deeply  pinnately  divided. 

KEY  TO  THE  SPECIES 

A.  Corolla-tube  hairy  inside  between  stamens;  filaments  liairy 1.  P.  adenophora 

A.  Corolla-tube  glabrous  inside;  filaments  glabrous. 

B.  Seeds  faintly  but  definitely  transversely  striate,  the  striations  about  12;  ovules 

20-26 3.  P.  inundata 

B.  Seeds   prominently   transversely  corrugated,  the  corrugations   6-12 ;    ovules  4-20. 

C.  Corolla  5-8  mm.  long;  corolla-scales  generally  conspicuous  and  united  to  base 

of  filaments  about  0.5  mm.;   style  and  branches   (1.75  or)   2-3.75  mm. 

long;   hypogynous  disk  conspicuous  Avith  prominent  rounded  or  deltoi<l 

lobes  alternating  with  the  stamens 2.  P.  lutea 

C.  Corolla  1.3-4.5  mm.  long;  corolla-scales  inconspicuous,  obsolescent,  or  lacking, 
free  from  the  filaments  at  base  if  present ;  style  and  branches  0.25-2  mm. 
long. 
D.  Plants  glabrous  or  nearly  so ;  corolla  subrotate  in  anthesis,  the  lobes  often 
about  as  long  as  the  tube;  ovary  and  style  glabrous  or  nearly  so. 

4.  P.  glaberrima 

D.  Plants  more  or  less  hirsutulous  or  villous  and  capitate-glandular  or  viscidu- 

lous;  corolla  eampanulate,  the  lobes  shorter  than  the  tube  (or  about 

equaling   the   tube   in  P.   tetramera)  ;    ovary   generally   sparsely  to 

densely  hairy,  the  style  hairy  at  least  at  base. 

E.  Flowers  mostly  5-mcrous;  corolla  3-4.5  mm.  long. 

F.  Style  and  branches  1.25-2  mm.  long;   hypogynous  disk  incon- 
spicuous; ovules  10—15. 
G.  Style  hairy  to  above  middle;  capsule  apiculate. 

5.  P.  scopulina 
G.  Style  glabrous  except  at  very  base;  capsule  nearly  or  quite 

without  apiculation 6.  P.  submutica 

F.  Style  and  branches  1  mm.  long;  hypogynous  disk  relatively  con- 
sj)icuous. 
H.  Ovules  18-20;   seeds  nearly  1  mm.  long,  the  corrugations 

6  or  7 7.  P.  inyoensis 

H.  Ovules  7-9;  seeds  1.5-2  long,  the  corrugations  about  9-11. 

8.  P.  salina 
E.  Flowers  commonly  4-merous;  corolla  1.3-1.8  mm.  long. 

9.  P.  tetramera 

1 .  Phacelia  adenophora  J.  T.  Howell 

Leafl.  West.  Bot.  4:15  (1944) 

Emvienanthe  gJandulifcra  Torr.  ex  S.  Wats.,  Bot.  U.S.  Geol.  Explor.  40th  Paral.  (King's 

Exped.)  257  (1871). 
Miltitsia  glandulifera  (Torr.)  Heller,  Muhlenbergia  8:20  (1912). 
Miltitsia  glanduUfera  (Torr.)  Heller  var.  calif ornim  Brand,  Univ.  Calif.  Publ.  Bot.  4:224 

(1912). 
Not  PlKicelia  (jhnuhilifcm  Piper,  Contrib.  U.S.  Nat.  Herb.  11 :472  (1906). 


Vol.  XXV]  HOWELL:    PHACELIA  SECTION  MILTITZIA  361 

Stems  few  or  several,  spreading'  from  the  top  of  the  annual  taproot,  pros- 
trate or  strongly  ascending,  slender  or  rather  stout,  mostly  1-3  dm.  long, 
capitate-glandular  and  villous-puberulent  or  hirsutulous;  basal  leaves  not 
very  numerous  and  the  upper  leaves  rather  sparse,  to  3  em.  long  and  1.7  cm. 
wide,  oblongish  to  subovate,  pinnately  lobed  or  divided,  the  lobes  oblongish, 
obtuse,  entire  or  rarely  dentate-lobed,  the  uppermost  leaves  sometimes  entire, 
hirsutulous  and  little  glandular,  petioles  hirsutulous  and  glandular,  to  2.5  cm. 
long;  racemes  mostly  exceeding  the  leafy  part  of  the  plant,  to  1  dm.  long, 
sessile  or  nearly  so,  flowers  somewhat  distant  below,  congested  above  or  the 
flowers  on  spreading  or  recurved  pedicels,  pedicels  stout  or  slender,  1-3  mm. 
long;  flowers  5-merous;  calyx-segments  in  flower  2—4.5  mm.  long,  0.4—1  mm. 
wide,  in  fruit  5-6.5  mm.  long,  0.5-1.25  mm.  wide,  linear  to  narrowly  oblong- 
oblanceolate,  sometimes  a  little  spatulate,  obtuse,  hirsutulous  and  more  or  less 
glandular ;  corolla  marcescent  and  a  little  accrescent,  tubular  to  campanulate, 
yellow  or  the  lobes  more  or  less  tinged  with  lavender,  4r-l  mm.  long,  lobes  1.25- 
2  mm.  long,  outside  of  corolla  hairy,  inside  spareely  to  densely  hairy  in  the 
tube  between  the  stamens;  corolla-scales  1-2  mm.  long,  linear  to  narrowly 
semilinear-lanceolate,  attached  at  base  to  base  of  filaments  about  0.5  mm.; 
stamens  3-5  mm.  long,  somewhat  unequal,  the  filaments  hairy  below  the  mid- 
dle, anthers  0.5-0.75  mm.  long  (or  rarely  shorter)  ;  styles  and  branches  2-4 
mm.  long,  the  style  hairy  to  the  middle,  branches  0.5-0.75  mm.  long ;  ovary  1 
mm.  long,  ovules  8-14 ;  hypogynous  disk  relatively  prominent,  the  lobes  be- 
tween the  stamens  broad  and  rounded ;  capsule  4-6  mm.  long,  oblong  to  oblong- 
ovate,  subacute  to  truncate  below  the  terminal  apiculation,  longitudinally 
sulcate  on  backs  of  valves,  sparsely  puberulent,  sometimes  a  little  glandular 
above ;  seeds  oblongish,  1-1.5  mm.  long,  black-brown,  prominently  transversely 
corrugated,  corrugations  8-12,  rounded  and  mostly  broad. 

Alkaline  soil  of  high  plains  and  sagebrush  slopes,  from  middle  western  Ne- 
vada northward  to  northeastern  California  and  southeastern  Oregon. 

Eeprescntative  collections.  Nevada  :  Virginia  Mts.,  5000  ft.,  8.  Watson 
No.  885  (G,  type  collection  of  Emmenanthe  gtandulifera)  ;  about  6  miles  south 
of  Gardnerville,  Douglas  Co.,  Cantetoiv  in  1941  (CAS,  CM,  lU,  US) ;  Carson 
City,  Ormsby  Co.,  Anderson  in  1865  (G,  NY) ;  Reno,  Washoe  Co.,  M.  E.  Jones 
No.  3986  (CAS,  DS,  G,  lU,  M,  NY,  P,  RM,  Vi^),  Eastivood  No.  14773  (CAS, 
CM,  P)  ;  5  miles  east  of  Sparks,  Washoe  Co.,  Eastwood  and  Howell  No.  30 
(CAS,  US)  ;  Pyramid  Lake,  Washoe  Co.,  Kennedy  No.  1037  (CM,  DS,  G,  M, 
NY,  RM,  UC,  US)  ;  12  miles  north  of  Reno  toward  Pyramid  Lake,  Washoe 
Co.,  Ripley  a7id  Barne'by  No.  4491  (CAS,  G).  In  Graj'-  Herbarium  is  a  speci- 
men labeled  "Winnemucca  Valley"  collected  by  Lemmon  in  1875,  but  the  cor- 
rectness of  the  locality  is  to  be  questioned. 

California:  Near  Viewland,  Lassen  Co.,  Ripley  and  Barnehy  No.  5755 
(CAS);  12  miles  north  of  Susanville,  Lassen  Co.,  J.  T.  Howell  No.  11884 
(corolla-limb  pale  lavender  fading  to  lavender-rose;  CAS,  US,  W,  WS)  ; 
Horse  Lake,  Lassen  Co.,  J.  T.  Howell  No.  11892  (corolla  bright  yellow;  CAS, 


362  CALIFORNIA  ACADEMY  OF  SCIENCES        [Proc.  4th  Ser. 

CM,  G,  lU,  M,  ND),  J.  r.  Howell  No.  11898  (corolla-limb  lavender;  CAS, 
NY,  P,  RM) ;  Madeline  Plains,  Lassen  Co.,  Bnice  No.  2135  (UC,  type  of  3///- 
titzia  glandulifera  var.  calif oniica) ,  Peirson  No.  6851  (CAS,  P),  J.  T.  Howell 
No.  11909 A  (CAS,  DS,  W);  canyon  below  Jess  Valley,  Modoc  Co.,  Payne 
No.  549  (CAS). 

Oregon,  Harney  Co. :  Catlow  Valley,  Henderson  No.  8690  (CAS)  ;  Blitzen, 
Train  No.  90  (US)  ;  base  of  Steens  Mts.,  Train  (Univ.  Colo.).  These  collec- 
tions from  Oregon  are  not  typical  and  show  characters  that  vary  toward 
P.  lutea. 

Phacelia  adenophora  is  a  rather  variable  entity,  but  it  is  usually  well  marked 
by  the  conspicuous  pubescence  on  the  filaments  and  in  the  corolla-tube.  In 
typical  form  as  it  occurs  in  middle  western  Nevada,  it  is  a  slender  and  some- 
what elegantly  branched  plant  with  the  upper  parts  rather  finely  hirsutulous 
and  copiously  capitate-glandular.  In  the  vicinity  of  Reno,  Nevada,  and  to  the 
northward,  this  typical  variant  passes  by  numerous  intergrades  into  a  coarser, 
more  densely  pubescent  plant  with  flowers  a  little  larger,  the  form  which 
Brand  called  Miltitzia  glandulifera  var.  calif ornica.  From  southeastern  Ore- 
gon relatively  few  collections  of  P.  adenophora  are  known,  but,  because  the 
plants  that  grow  there  frequently  exhibit  characters  intermediate  to  P.  lutea 
which  also  grows  in  the  same  region,  it  is  suspected  that  the  intermediates 
may  be  of  hybrid  origin. 

Although  P.  adenophora  is  closely  related  to  P.  lutea  and  perhaps  should 
be  considered  only  a  variety  of  that  species,  I  regard  the  differences  between 
the  two  specific  in  value.  It  is  true  that  Asa  Gray  remarked  that  P.  adenophora 
{Emmenanthe  glandidifera)  was  ''probably  a  mere  form"  of  P.  lutea  (1878. 
p.  171) ,  but  an  examination  of  specimens  in  Gray  Herbarium  discloses  that  at 
that  time  he  had  no  specimen  of  P.  lutea,  and  that  he  mistook  for  that  species 
the  more  robust  form  of  P.  adenophora  which  Brand  has  called  Miltitzia 
glandulifera  var.  calif  ornica.  Quite  recently  Jepson  has  treated  P.  adenophora 
and  P.  lutea  as  the  same  species  (1943,  p.  281,  as  Miltitzia  lutea),  but  each  is 
characterized  by  a  distinctive  appearance  and  each  by  its  own  peculiar  set  of 
variations  which  do  not  approach  each  other  except  in  instances  of  suspected 
hybridization.  I  do  not  believe  that  there  should  be  a  change  in  the  status 
of  P.  adenophora  until  its  relation  to  P.  lutea  is  clearly  understood  through 
more  detailed  study  and  critical  field  work. 

Within  P.  adenophora  no  variant  seems  sufficiently  distinguished  to  be 
recognized  taxonomically.  Although  Brand's  var.  calif  ornica  of  northeastern 
California  has  considerable  right  to  varietal  recognition,  I  have  concluded 
that  it  is  not  clearly  enough  defined  either  geographically  or  morphologically 
to  be  accepted  at  this  time.  Also  I  have  seen  several  collections  of  a  peculiar 
glandular-villous  form  with  large  floAvers  and  slender  fruits  from  southern 
Oregon  which  should  perhaps  be  accorded  taxonomic  recognition,  but.  be- 
cause of  a  possible  connection  which  this  plant  may  have  with  the  imperfectly 
understood  relations  between  P.  adenophora  and  P.  lutea  in  tliat  region. 


VoL.XXV]  HOWELL:  PHACELIA  SECTION  MILTITZIA  363 

nominal  recognition  of  this  entit}^  too,  should  await  more  numerous  collections 
and  further  field  work. 

A  collection  intermediate  between  P.  adenophora  and  P.  lutea  is  Coville 
and  Leiherg  No.  68  (US),  from  the  road  to  Coleman  Valley,  Harney  or  Mal- 
heur Co.,  Oregon.  In  it  the  corolla-tube  is  pubescent  only  at  the  base  of  the 
stamens,  or  glabrous,  and  the  filaments  vary  from  subglabrous  to  distinctly 
hairy.  Another  intermediate  collection  from  Oregon  looking  quite  like  P. 
adenophora,  but  with  the  technical  floral  characters  of  P.  lutea,  is  Willits 
No.  169  (DS)  from  Hart  Lake,  Lake  Co.  A  collection  of  P.  adenophora  which 
resembles  P.  lutea,  but  which  has  heavily  hairy  corolla-tube  and  filaments, 
has  been  collected  near  Canby,  Modoc  Co.,  California,  /.  T.  Howell  No. 

12215  (CAS). 

2.  Phacelia  lutea  (H.  and  A.)  J.  T.  Howell 

Leafl.West.  Bot.  4:15  (1944) 

Stems  one  to  several  from  the  top  of  an  annual  taproot,  spreading  or 
ascending  to  substrictly  erect,  2-8  cm.  long,  hirsutulous  and  scarcely  gland- 
ular to  hirsutulous  and  capitate-glandular  or  almost  entirely  glandular- 
villous ;  leaves  entire  or  coarsely  serrate,  rarely  serrate-lobed,  oblanceolate 
and  oblong  to  elliptic  and  ovate,  mostly  1-2.5  cm.  long,  0.3-1  cm.  wide, 
hirsutulous  and  a  little  glandular  to  sparsely  glandulai'-villous,  obtuse  at 
apex,  cuneate  at  base,  petiole  to  2.5  cm.  long,  usually  equaling  or  shorter 
than  the  blade ;  racemes  to  5  cm.  long,  sometimes  more  or  less  exceeded  by 
and  concealed  by  the  leaves  but  generally  exceeding  the  leaves  and  showy, 
pedicels  1-2  mm.  long  or  the  loAvest  to  5  mm.  long,  the  lowest  sometimes 
reflexed  in  fruit ;  calyx-segments  in  flower  3.5-5  mm.  long,  in  fruit  6.5-9 
mm.  long,  1-1.75  mm.  wide,  linear-oblong  to  narrowly  oblong-oblanceolate, 
subacute,  hirsutulous  and  more  or  less  glandular  to  rather  densely  capitate- 
glandular  ;  corolla  yellow,  sometimes  becoming  lavender-tinged  in  age,  5-8 
mm.  long,  campanulate  or  tubular,  the  lobes  erect  or  spreading  in  anthesis, 
1-2  mm.  long,  denticulate,  corolla  sparsely  hairy  outside,  glabrous  inside ; 
corolla-scales  2-3  mm.  long  or  sometimes  obsolescent,  semilinear-lanceolate, 
attached  at  base  about  0.5  mm.  to  the  filaments ;  stamens  3-5.5  mm.  long, 
filaments  glabrous,  anthers  0.4-0.75  mm.  long;  style  and  branches  1.75-3.75 
mm.  long,  branches  0.25-0.75  mm.  long,  style  hairy  only  near  the  base  or 
to  above  the  middle;  ovary  1.25-1.75  mm.  long,  ovules  10-17;  hypogynous 
disk  prominent  with  conspicuous  rounded  or  deltoid  lobes  between  the 
stamens ;  capsule  5.5-7  mm.  long,  oblong  to  elliptic,  subobtuse,  apiculate, 
the  beak  to  1  mm.  long,  sparsely  hairy  and  a  little  glandular ;  seeds  1-1.5 
mm.  long,  elliptic-oblong,  subobtuse  to  subacute,  brown,  prominently  cor- 
rugated, the  corrugations  about  9  or  10,  conspicuously  reticulate. 

Clayey  and  sandy  slopes  and  flats  in  southeastern  Oregon,  occurring 
locally  in  western  Idaho  and  northwestern  Nevada. 

The  type  of  P.  lutea  was  part  of  a  collection  of  plants  "gathered  in  the  sum- 
mer of  1837  by  a  friend  of  Mr.  Tolmie"  at  "Snake  Fort,  Snake  Country."  In 


364  CALIFORNIA  ACADEMY  OF  SCIENCES        [Proc.  4th  Ser. 

the  introduction  to  the  Supplement  of  Botany  of  Captain  Beeehey's  Voyage 
(p.  315),  it  is  stated  that  "Snake  Fort  is  built  at  the  junction  of  Reed's  River 
with  the  Snake  [River],  the  position  of  which  is  in  N.  lat.  44°  20',  long.  116° 
W."  On  modern  maps  the  name  Reed's  River  does  not  appear  and  the  lati- 
tude and  longitude  indicate  a  point  considerably  east  of  the  Snake  RiA'er. 
On  the  map  in  Hooker's  Flora  Boreali-Americana,  vol.  1,  however,  Reed's 
River  is  given  and  from  its  position  it  is  clearly  the  Boise  River  of  the  present 
day,  the  chief  affluent  of  the  Snake  River  from  the  east  between  the  Payette 
River  and  the  great  bend  of  the  Snake.  Hence,  collections  that  have  been 
available  from  northwestern  Owyhee  Countj^  Idaho,  may  be  regarded  as  topo- 
types  of  this  interesting  species  that  was  first  collected  more  than  a  hundred 
years  ago. 

As  has  been  pointed  out  in  the  discussion  of  P.  adenopliora ,  Asa  Gray  had 
no  specimens  of  P.  lutea  at  the  time  he  prepared  his  account  of  Emmenmifhe 
§  Miltitzia  for  the  Synoptical  Flora  (1878) .  As  is  indicated  by  his  annotations 
on  specimens  in  Gray  Herbarium,  he  did  receive  collections  of  P.  lutea  from 
Cusick  by  the  time  the  second  edition  was  published  (1886),  but  there  is 
nothing  in  the  supplement  of  the  second  edition  concerning  these  specimens. 
Cusick's  plants,  collected  from  three  different  stations  under  varying  environ- 
ments, were  probably  more  confusing  than  helpful  to  Gray.  One  part  of  the 
collection.  No.  1233a,  from  "sandy  hillsides  of  Malheur  and  vicinity"  is  typical 
P.  lutea,  but  No.  12331),  "same  locality  but  on  heavy  clay  soil,"  and  No.  1233c, 
"barren  valley  in  rather  heavy  soil,"  are  quite  different  in  appearance  and 
were  originally  noted  as  probably  Emmenanthe  parviflora  (P.  inundata)  by 
Gray,  and  were  more  recently  identified  as  Miltitzia  gland^difcra  (i.e.,  P.  ade- 
nophora)  by  Macbride.  Although  these  puzzling  plants  and  others  like  them 
from  Professor  Peck's  collections  are  definitely  referable  to  P.  lutea,  they 
vary  toward  P.  scopidina  and  present  a  problem  of  relationship  that  can  only 
be  indicated  at  this  time.  These  variants  are  discussed  further  under  P.  .scopu- 
lina,  and  under  P.  adenopliora  the  problem  of  intermediates  between  that 
species  and  P.  lutea  is  discussed. 

One  variant  in  P.  lutea,  howeATr,  seems  quite  distinct  and  well  deserving 
taxonomic  recognition,  a  small-flowered  plant  with  glandular-villous  pubes- 
cence. This  variety  may  be  distinguished  as  follows : 

Key  to  Varieties  of  Phacelia  liutea 

Corolla  6-8  mm,  long;  style  hairy  to  the  middle  or  above var.  typica 

Corolla  5-6  mm.  long;  style  hairy  only  near  the  base var.  purpurascens 

2a.  Phacelia  lutea  (H.  and  A.)  J.  T.  Howell 

var.  typica,  nom.  nov. 

P.  lutea  (H.  and  A.)  J.  T.  Howell,  1.  c. 

Euioca  lutea  H.  and  A.  Bot.  Beech.  Voy.  373  (1840)  ;  Hook.  Icon.  tab.  354  (1841). 

Miltitzia  lutea  (H.  and  A.)  A.  DC.  in  DC.  Prodr.  9:296  (1845). 

Emmertanthe  lutea  (11.  and  A.)  A.  Gray,  Proc.  Amer.  Acad.  10:328  (1875). 


Vol.  XXV]  HOWELL:  PEACELI A   SECTION  MILT ITZI A  365 

Plants  hirsntulous  and  scarcely  glandular  to  moderately  glandular- 
liirsutulous,  generally  yellowish-green ;  racemes  mostly  exceeding  the  leaves ; 
corolla  6-8  mm.  long,  showy ;  style  sparsely  hairy  to  the  middle  or  above, 
2.5-3.75  mm.  long. 

Representative  collections.  Idaho:  Squaw  Creek  near  Marsing,  Owyhee 
Co.,  Tucker  No.  1029  (CAS  ex  Univ.  Ida.  So.  Br.). 

Oregon:  Harper,  Malheur  Co.,  Peck  No.  20553  (CAS,  W),  Henderson  No. 
8689  (CAS)  ;  5  miles  south  of  Harper,  Malheur  Co.,  Peck  No.  16104  (W, 
approaching  var,  purpurascens)  ;  Roekville,  Malheur  Co.,  Henderson  No. 
8691  (CAS,  depauperate) ;  Sucker  Creek,  30  miles  north  of  Jordan  Valley, 
Malheur  Co.,  Holmgren  No.  2206  (CAS,  lU)  ;  Malheur  Butte,  Malheur  Co., 
Leiierg  No.  2043  (CM,  G,  NY,  UC,  US)  ;  15  miles  southeast  of  Blitzen,  Harney 
Co.,  Applegate  No.  5591  (DS,  W)  ;  near  Burns,  Harney  Co.,  Lilla  Luch  in 
1927  (W) ;  base  of  SteensMts.,  Harney  Co.,  HoweZZmiSSo  (CM,  G,  NY,  US)  ; 
Unity,  Baker  Co.,  Peck- No.  16186  (W)  ;  Paulina  Creek  northeast  of  Lapine, 
Deschutes  Co.,  WUted  No.  1679  (ND). 

Nevada:  Virgin  Valley,  Humboldt  Co.,  L.  Kellogg  in  1909  (P). 

An  unsual  variant,  entirely  glabrous  except  for  an  occasional  hair  in  the 
inflorescence  and  for  a  few  scattered  hairs  and  glands  at  the  top  of  the  ovary 
and  on  the  lower  part  of  the  style,  has  been  collected  by  Ripley  and  Barneby 
in  Owyhee  County,  Idaho,  where  it  grew  on  a  "friable  alkaline  clay  bank  24 
miles  southwest  of  Marsing"  (No.  6153,  CAS,  G,  US).  A  few  of  the  corollas 
were  noted  by  the  collectors  as  being  tetramerous. 

2b.  Phacelia  lutea  (H.  and  A.)  J.  T.  Howell 
var.  purpurascens  J.  T.  Howell,  var.  nov. 

Glanduloso-villosa  vix  hirsutula,  caulibus  foliis  rhachidibus  et  segmentis 
calycis  valde  purpurascenti-tinctis;  foliis  racemos  plerunique  superantibus ; 
corolla  5-6  mm.  longa;  stylo  1.75-2.5  mm.  longo,  pilifero  solum  prope  basin. 

Type:  Herb.  Calif.  Acad.  Sci.  No.  127399,  collected  in  "chocolate  soil"  on 
high  hills  opposite  Humphrey's,  Grant  County,  Oregon,  by  L.  F.  Henderson, 
No.  5092,  April  30,  1925.  There  is  a  duplicate  in  Gray  Herbarium.  Only  one 
other  collection  entirely  typical  of  var.  purpurascens  has  been  seen :  Cusick 
No.  2634,  which  was  obtained  "in  red  clay  soil  of  Crook  Co."  (CM,  G,  M,  ND, 
NY,  P,  RM,UC,US). 

3.  Phacelia  inundata  J.  T.  Howell 

Leafl.  West.  Bot.  4 :  15  (1944) 

Emmenanthe  parviflora  A.  Gray,  U.S.  Pacif.  ER.  Reports  6:84,  tab.  15  (1857). 
Miltitzia  parviflora  (A.  Gray)  Brand,  Das  Pflanzenr.  IV.  251:131  (1913). 
Not  Phacelia  parviflora  Pursli,  Fl.  Amer.  Sept.  1 :140  (1814)  ;  nor  P.  parviflora  Phil.,  Anal. 
Univ.  Chile  90:226  (1895). 

Stems  several,  spreading  or  ascending  from  the  top  of  the  annual  taproot, 
mostly  1-4  dm.  long,  but  the  plants  sometimes  depauperate  and  the  stems  only 
0.1-0.3  dm.  long,  finely  glandular  and  somewhat  hirsutulous;  leaves  rosu- 


366  CALIFOSNIA  ACADEMY  OF  SCIENCES        [Proc.  4th  Ser. 

late  and  also  rather  numerous  up  to  the  ends  of  the  branches,  mostly  1-3  cm. 
long  and  0.4r-1.5  em.  wide,  oblong  to  oblong-obovate,  subentire  to  pinnately 
parted  and  divided,  the  divisions  obtuse,  mostly  entire,  oblong  or  deltoid- 
oblong,  the  terminal  division  largest  and  generally  lobed,  the  basal  divisions 
frequently  small  and  discrete,  petioles  0.5-2  cm.  long,  finely  glandular- 
hirsutulous,  the  blades  finely  hirsutulous  and  a  little  glandular  beneath ; 
racemes  mostly  exceeded  by  the  leaves,  to  12  cm.  long,  the  lowest  flowers  rather 
distant,  the  uppermost  somewhat  crowded,  pedicels  1-2  mm.  long  or  the  lowest 
to  4  mm.  long,  erect  or  somewhat  spreading  in  fruit ;  flowers  5-merous ;  calyx- 
segments  in  flower  narrowly  oblongish,  4  mm.  long,  0.6-1  mm.  wide,  in  fruit 
6-10  mm.  long,  1-1.5  mm.  wide,  linear-oblanceolate  to  linear-oblong,  some- 
times a  little  spatulate,  obtuse,  hirsutulous  and  a  little  glandular;  corolla 
yellow,  4—5  mm.  long,  the  lobes  rotately  spreading  in  anthesis,  becoming  erect 
later,  lobes  about  2  mm.  long,  pilose-hairy  outside,  glabrous  within;  corolla- 
scales  narrowly  semilinear -lanceolate,  1.25  mm.  long,  at  base  free  from  fila- 
ments, or  the  scales  obsolete ;  stamens  quite  unequal,  2-3  mm.  long,  filaments 
glabrous,  anthers  roundish,  0.3  mm.  long;  style  and  branches  1-1.25  mm.  long, 
style  hirsutulous  nearly  to  the  branches,  branches  0.25  mm.  long;  ovary  1.5 
mm.  long,  densely  hirsutulous,  ovules  20-26 ;  hypogynous  disk  rather  promi- 
nent with  rounded  lobes  between  the  stamens ;  capsule  5.5-7  mm.  long,  oblong, 
abruptly  narrowed  above  into  a  prominent  deltoid  apiculation  and  persistent 
style,  sparsely  hairy  below,  sparsely  hirsutulous  above  and  on  the  beak,  not 
glandular;  seeds  1.5-1.75  mm.  long,  subdeltoid,  irregularly  angled,  mostly 
acute  at  the  ends,  tending  to  be  strongly  flattened,  reticulate-pitted,  finely  but 
definitely  transversely  striate,  the  striations  about  12-14. 

Summer-dried  playas  of  desert  lakes  and  sinks  from  middle  southern  and 
southeastern  Oregon  south  to  northeastern  California  and  middle  western 
Nevada. 

Collections.  Oregon:  Klamath  Lake,  Klamath  Co.,  Xewhcrry  (G,  type); 
Silver  Lake,  Harney  Co.,  Cusick  No.  2727  (CM,  G,  M,  ND,  NY,  P,  RM,  UC, 
US;  plant  noted  as  having  a  "very  disagreeable  odor") ;  Steens  Mts.  above 
Alberson,  Harney  Co.,  Teck  No.  14217  (NY,  W). 

California  :  east  side  of  Clear  Lake,  Modoc  Co.,  J.  T.  Howell  No.  12357 
(CAS,  US)  ;  between  Malin  and  Canby,  Modoc  Co.,  M.  S.  Baker  No.  8307 
(CAS)  ;  west  side  of  Eagle  Lake  at  Spaldings,  Lassen  Co.,  /.  T.  Howell  No. 
12510  (CAS,  CM,  DS,  P,  W). 

Nevada  :  Swan  Lake,  Washoe  Co.,  Train  No.  3006  (lU,  NY) ;  Washoe  Lake, 
Washoe  Co.,  M.  E.  Jones  in  1897  (CAS,  P)  ;  Franktown,  Washoe  Co.,  Ripley 
and  Barnehy  No.  5947  (CAS) . 

Phacelia  inundata  is  one  of  the  more  rarelj'-  collected  species  in  the  section 
Miltitzia.  Although  three  of  the  nine  collections  that  have  been  examined  have 
come  from  northea-stern  California,  before  this  it  has  been  reported  only 
from  southeastern  Oregon  and  Nevada  (Tidestrom,  1925,  p.  448;  Peck,  1941, 
p.  593).  Only  a  single  specimen  has  been  seen  (Peck  No.  14217)  in  which  the 


VoL.XXV]  HOWELL:  PHACELIA  SECTION  MILTITZIA  367 

collection  data  indicate  an  environment  away  from  a  desert  lake  or  sink,  while 
the  Californian  collections  which  I  made  came  from  lake-bottom  areas  that 
had  been  covered  with  water  in  the  earlier  part  of  the  season.  Because  of  its 
preference  for  so  peculiar  a  habitat,  P.  inimdata  appears  to  be  one  of  the  most 
highly  specialized  and  advanced  species  in  the  genus  from  a  physiological 
point  of  view.  Morphologically  it  is  quite  distinct,  too,  its  nearest  relative 
perhaps  being  P.  lutea. 

4.  Phacelia  glaberrima  (Torr.)  J.  T.  Howell 

Leafl.  West.  Bot.  4:15  (1944) 

Emmenanthe  glaberrima  Torr.  ex  S.  Wats.,  Bot.  U.S.  Geol.  Explor.  40th  Paral.   (King's 

Exped.)  257  (1871). 
Miliiisia  glaberrima  (Torr.)  Brand,  Das  Pfianzenr.  IV.  251:131  (1913). 

Stems  several,  spreading  from  top  of  annual  taproot,  3-20  cm.  long  and 
probably  longer,  first  internodes  above  the  base  very  long,  entirely  glabrous ; 
leaves  1-1.5  cm.  long,  0.4-1  cm.  wide,  glabrous  or  with  a  few  hirsutulous  hairs 
on  the  margins  and  near  the  apex,  broadly  oblanceolate  and  ovate  to  broadly 
elliptic,  entire  or  with  1  to  few  coarse  serrations,  obtuse  at  apex,  narrowly 
cuneate  at  base  and  frequently  a  little  decurrent  along  the  petiole,  petiole  to 
2  cm.  long  below,  very  short  above ;  racemes  to  8  cm.  long,  flowers  approximate 
or  rather  distantly  spaced,  pedicels  mostly  1-3  mm.  long ;  flowers  5-merous ; 
calyx-segments  in  flower  very  unequal,  2.5  mm.  long,  0.25-1  mm.  wide,  in 
fruit  4—5  mm.  long,  0.3-2  mm.  wide,  oblanceolate  to  obovate,  fleshy  and  with 
a  prominent  midrib  near  the  base,  glabrous  or  with  a  few  short  scattered 
hirsutulous  hairs  near  the  apex ;  corolla  3-4  mm.  long,  subrotate  in  anthesis, 
the  lobes  1-3  mm.  long,  becoming  erect  later,  entirely  glabrous  within  and 
without;  corolla-scales  obsolete;  stamens  2.5-3  mm.  long,  filaments  glabrous, 
flattened  at  base,  anthers  0.4  mm.  long ;  style  and  branches  1-2  mm.  long,  the 
style  glabrous,  deltoid-subulate,  compressed,  the  branches  0.5  mm.  long, 
ascending ;  ovary  glabrous  or  sometimes  with  a  few  scattered  hairs  near  the 
top,  1  mm.  long,  ovules  4-10 ;  hypogynous  disk  prominent  about  uniformly 
wide  all  around  and  not  conspicuously  lobed;  capsule  4-5  mm.  long,  ovate  to 
elliptic,  glabrous  or  nearly  so,  somewhat  rounded  above,  prominently  apicu- 
late,  valves  coriaceous  with  conspicuous  thickened  margins,  longitudinally 
sulcate;  seeds  1.5-2  mm.  long,  oblong  or  ovatish,  prominently  corrugated, 
corrugations  rounded,  reticulate,  about  10  or  11. 

Phacelia  glaberrima  is  a  remarkable  and  localized  endemic  in  the  saline 
desert  of  north-central  Nevada  and,  until  it  was  rediscovered  in  the  Keese 
River  Valley  in  June,  1944,  by  H.  D.  Ripley  and  R.  C.  Barneby,  it  was  known 
only  from  the  collections  of  Sereno  Watson  made  in  1868  on  King's  Expedi- 
tion. The  three  presently  known  collections  are:  Humboldt  Sink,  4300  ft., 
Watson  in  Maij,  1868  (G,  UC  frag.)  ;  Reese  Valley,  5000  ft.,  Watson  in  July, 
1868  (G,  UC  frag.)  ;  Reese  River  Valley  32  miles  north  of  Austin,  Lander 
Co.,  Eipley  and  Barnehy  No.  6186  (CAS,  G,  US).  Concerning  the  highly 


368  CALIFORNIA  ACADEMY  OF  SCIENCES        [Proc.  4th  Ser. 

specialized  liabitat  of  P.  glaherrima,  Mr.  Barneby  writes  as  follows :  "The 
Reese  River  Valley  is  not  verj^  exciting  for  the  most  part,  one  of  those  broad 
sterile  valley's  covered  with  sagebrush  and  with  long  fans  of  gravel  leading 
into  the  hills,  but  at  one  point  about  60  miles  south  of  Battle  Mountain  the 
mountains  close  in  and  the  river  has  to  negotiate  a  short  canyon.  Here  the 
mountains  are  mostly  volcanic,  but  there  are  localized  occurrences  of  lime- 
stone, and  these  have  been  broken  down  by  weathering  into  white  bluffs  of  a 
stiff  gravel  and  clay  mixture  impregnated  with  alkali.  On  the  sides  of  these 
bluffs,  on  which  nothing  else  can  exist,  Phacelia  glaherrima  is  abundant,  its 
yellowish  herbage  very  succulent  and  fragile  and  its  abundant  racemes  of 
golden-yellow  visible  from  afar.  Occasionally  it  seeds  down  into  the  washes 
and  there  was  one  spot  where  it  was  associated  with  P.  scopulma  and  P. 
gymnoclada,  the  three  species,  so  similar  and  yet  so  different,  sometimes 
enlaced  into  small  "trispecific"  mats.  The  corolla  of  P.  glaherrima  is  very 
open-campanulate,  almost  rotate  when  fully  expanded,  and  the  leaves  are 
occasionally  crenate." 

Although  P.  glaherrima  might  be  related  to  P.  scopulina  because  of  its 
smaller  corollas  which  are  devoid  of  corolla-scales,  this  rare  Nevada  plant  is 
probably  much  nearer  P.  lufea  and,  in  its  subrotate  corollas,  it  is  reminiscent 
of  the  equally  highly  differentiated  P.  imindata,  a  species  which  I  have  also 
related  to  P.  lutea.  Among  the  numerous  and  varied  Miltitzia  collections  of 
Ripley  and  Barneby  are  remarkable  glabrous  forms  of  both  P.  lutea  and  P. 
scopulina  but  from  these  P.  glaherrima  may  be  readily  distinguished  by  the 
rotately  spreading  and  deeply  lobed  corolla,  the  glabrous  style,  and  the  thick- 
rimmed  valves  of  the  capsule.  From  Watson's  collections  I  concluded  that  the 
ovary  was  glabrous  as  originally  described  but  in  the  ample  material  of 
Ripley  and  Barneby  it  is  apparent  that  the  ovary  is  sometimes  sparsely  and 
minutely  hairy  near  the  top. 

Phacelia  glaherrima  has  been  reported  from  the  Flax  River  (Little  Colorado 
River)  in  northern  Arizona,  where  it  was  supposed  to  have  been  collected  by 
Newberry,  but  I  have  located  no  Arizona  collection  (cf.  Kearney  and  Peebles, 

1942,  p.  737).  J 

5.  Phacelia  scopulina  (A.  Nels.)  J.  T.  Howell  | 

Leafl.  West.  Bot.  4:16  (1944) 

Emmenanthe  scopulina  A.  Nels.,  Bull.  Torr.  Bot.  Club  25  : 380  (1898). 

Miltitzia  lutea  (H.  and  A.)  A.  DC.  var.  scopulina  (A.  Nels.)  Brand,  Das  Pflanzenr.  IT. 

251:131  (1913). 
Miltitzia  scopidina  (A.  Nels.)  Eydb.,  Bull.  Torr.  Bot.  Club  40  : 479  (1913). 
Emmenanthe  dispar  M,  E.  Jones  ex  Eydberg,  Fl.  Eky.  Mts.  and  Adj.  Plains  708  (1917),  as 

synonym  of  Miltitzia  salina. 

Stems  several  from  the  top  of  an  annual  taproot,  2-10  cm.  long,  spreading 
or  ascending  and  tending  to  be  erect,  sparsely  to  rather  densely  hirsutulous,    » 
more  or  less  cinereous,  eglandulose  or  minutely  viscidulous  above;  leaves 
oblanceolate  to  oblong  or  oblong-obovate,  1-3  cm.  long,  0.5-1.5  cm.  wide, 
entire,  serrate-  or  crenate-lobed,  or  rather  deeply  pinnately  lobed,  the  lobes 


VoL.XXV]  BOWELL:  FRACELIA  SECTION  MILTITZIA  3(59 

oblong,  deltoid,  or  elliptic,  the  blade  obtuse  at  apex,  cuneate  to  rounded  at 
base,  hii-sutulous,  petioles  to  4  cm.  long,  frequently  longer  than  the  blades ; 
racemes  rather  few-flowered  and  more  or  less  concealed  by  the  leaves,  to 
4.5  cm.  long,  pedicels  1-6  mm.  long,  erect  or  sigmoid-spreading,  or  the  lowest 
recurved  in  fruit ;  floAvers  5-merous ;  calyx-segments  2.5-4.5  mm.  long,  about 
0.5  mm.  wide,  linear-oblong,  in  fruit  becoming  6-7  mm.  long,  0.5-1  mm.  wide, 
linear-oblong  or  linear-oblanceolate,  obtuse,  usually  Avithout  a  prominent 
raised  midvein  except  at  the  base,  hirsutulous,  rarely  capitate-glandular 
and  perhaps  sometimes  a  little  viseidulous;  corolla  tubular-campannlate, 
somewhat  accrescent  in  age,  3-5  mm.  long,  the  lobes  0.75-1  mm.  long,  yellow 
or  strongly  tinged  with  lavender  or  violet,  the  outside  hairy,  the  inside 
glabrous;  corolla-scales  inconspicuous  or  obsolete,  filiform-linear,  about 
1  mm.  long,  free  from  the  base  of  the  filaments ;  stamens  2.5-3  mm.  long,  the 
longest  frequently  equaling  the  lobes  of  the  corolla,  filaments  glabrous, 
anthers  0.3-0.4  mm.  long;  style  and  branches  1.3-2  mm.  long,  the  branches 
0.2-0.3  mm.  long,  the  style  hirsutulous-hairy  to  the  middle  or  above ;  ovary 
1-1.25  mm.  long,  ovules  11-15  ;  hypogynous  disk  inconspicuous  and  scarcely 
lobed  between  the  stamens ;  capsule  5-6  mm.  long,  oblong,  obtuse  or  sub- 
acute at  top  below  the  prominent  apiculation,  hirsutulous-puberulent  espe- 
cially above;  seeds  oblong  or  oblong-ovate,  1.25-1.75  mm.  long,  brown, 
prominently  corrugated,  corrugations  rounded,  9-11,  reticulate. 

Loose  sandy  or  gravelly  soil  of  desert  flats  and  slopes,  from  southwestern 
Montana  and  Wyoming  southward  to  western  Utah  and  northern  Nevada, 
and  westward  to  eastern  Oregon. 

Representative  collections.  Montana  :  Melrose,  Silver  Bow  Co.,  Rydherg 
No.  2771  (NY,  US). 

Wyoming  :  Green  River,  Sweetwater  Co.,  A.  Nelson  No.  3056  (RM,  type  of 
EmmenantJie  scopulina;  G,  M,  US),  A.  Nelson  No.  4712  (CAS,  G,  M,  NY,  P, 
RM,  US,  WS),  M.  E.  Jones  in  1896  (CAS,  lU,  M,  NY,  P,  RM,  US)  ;  between 
Eden  and  Big  Piney,  Sublette  Co.,  Pay  son  and  Pay  son  No.  2576  (CM,  G,  M, 
NY,  P,  RM,UC,US). 

Utah  :  Dutch  Mt.,  Tooele  Co.,  M.  E.  Jones  in  1891  (P). 

Nevada:  Wells,  Elko  Co.,  31.  E.  Jones  in  1930  (P),  Eastwood  &  Howell 
No.  330  (CAS,  CM,  DS,  ND,  US),  Ripley  and  Barnehy  No.  5524  (CAS)  ;  25 
miles  north  of  Wells,  Elko  Co.,  Holmgren  No.  955  (CAS,  lU,  UC)  ;  foothills 
of  Independence  Mts.  southeast  of  Tuscarora,  Elko  Co.,  Ripley  and  Barnehy 
No.  5551  (CAS) ;  near  Wendover,  Elko  Co.,  M.  E.  Jones  in  1929  (CAS,  P)  ; 
Cobre,  Elko  Co.,  M.  E.  Jones  in  1906  (CAS,  P),  in  1907  (P) ;  foothills  of 
Schell  Creek  Range  west  of  Connors  Pass,  White  Pine  Co.,  Ripley  and 
Barnehy  No.  6305  (CAS)  ;  Monitor  Valley  south  of  Lone  Mt.,  Eureka  Co., 
Ripley  and  Barnehy  No.  6203  (CAS)  ;  11  miles  west  of  Eureka,  Eureka  Co., 
Ripley  and  Barnehy  No.  6235  (CAS)  ;  Reese  River  Valley  32  miles  north  of 
Austin,  Lander  Co.,  Ripley  and  Barnehy  No.  6187  (CAS). 

Oregon  :  Harney  Lake,  Harney  Co.,  Henderson  No.  8688  (CAS)  ;  24  miles 


370  CALIFORNIA   ACADEMY  OF  SCIENCES        [Proc.  4th  Ser. 

north  of  Wagontire,  Harney  Co.,  Peck  No.  19394  (W,  simulating  atypical 
P.  lutea)  ;  8  miles  west  of  Lakeview,  Lake  Co.,  Peck  No.  15243  (W,  more  glan- 
dular than  typical) ;  south  of  Narrows,  Harney  Co.,  Bipley  and  Barneby  No. 
6088  (depauperate,  CAS). 

Although  Brand  considered  Emmenanthe  scopulina  a  variety  of  Miltitzia 
lutea,  I  can  see  no  reason  for  following  so  conservative  a  procedure.  Phacelia 
scopulina  has  a  very  distinctive  appearance  and  is  marked  by  excellent  char- 
acters over  a  wide  range;  and,  in  a  much  more  restricted  range,  Phacelia  lutea 
is  usually  quite  as  distinct.  In  eastern  Oregon,  however,  where  the  ranges  of 
the  two  species  overlap,  plants  have  been  collected  which  are  confusingly 
intermediate  in  appearance,  although  from  the  critical  characters  of  the 
flower-parts,  such  specimens  have  been  definitely  referable  to  one  species  or 
the  other.  A  pair  of  such  specimens  has  been  collected  by  Prof.  M.  E.  Peck, 
P.  lutea  coming  from  7  miles  east  of  Buchanan,  Harney  County  {No.  20536, 
W),  P.  scopulina  from  25  miles  north  of  McDermitts,  Malheur  County  {No. 
20600,  W) .  A  series  of  collections  of  P.  lutea  accompanied  by  field  notes  made 
by  Cusick  in  eastern  Oregon  and  preserved  in  the  Gray  Herbarium  would 
seem  to  indicate  that  some  of  the  variations  noted  may  be  due  to  environmental 
conditions,  but  it  seems  likely  that  much  of  the  variability  results  from  the 
genetic  makeup  of  the  group  and  may  be  due  to  hybridization  between  P.  lutea 
and  P.  scopulina.  Cusick's  collections  are  more  fully  described  under  P.  lutea. 

Another  collection  of  P.  scopulina  that  exhibits  noteworthy  variability  was 
made  by  Ripley  and  Barneby  in  Humboldt  County,  Nevada  {No.  4557,  CAS). 
In  this  collection  there  are  two  kinds  of  plants,  one  that  is  normally  hirsutu- 
lous-pubescent  throughout,  the  other  that  is  quite  glabrous  except  for  a  few 
scattered  hirsutulous  hairs  on  some  of  the  leaves  and  for  a  few  minute  capitate- 
glands  on  the  stems,  calyx-segments,  and  upper  part  of  the  ovary.  Because 
the  only  known  collections  of  P.  glaherrima  were  made  in  the  northwestern 
part  of  Nevada  not  very  far  from  the  station  where  Ripley  and  Barneby  found 
this  odd  variant,  their  plants  were  compared  with  specimens  of  P.  glaherrima. 
The  floral  morphology  of  the  two  was  found  to  be  so  unlike  that  it  would  be 
quite  wrong  even  to  suggest  that  the  subglabrous  individuals  are  intermediate 
between  the  two  species. 

M.  E.  Jones  distributed  his  1896  collection  of  P.  scopulina  made  at  Green 
River  with  the  name  Emmenanthe  dispar,  hut  apparently  he  never  described 
the  plant.  Although  Rydberg  published  the  name  as  a  synonym  of  Miltitzia 
salina,  as  indicated  above  in  the  synonymy,  he  had  correctly  determined  and 
annotated  Jones'  specimen  in  Herb.  N.Y.  Bot.  Gard.  as  Miltitzia  scopulina. 

6.  Phacelia  submutica  J.  T.  Howell,  spec.  nov. 

Herba  annua  plus  minusve  glanduloso-hirsutula;  caulibus  paucis  vel  mul- 
tis  ex  ba.si,  patentibus  vel  suberectis,  2-8  cm.  longis,  purpurascenti-tinctis, 
sparse  vel  subdeiise  vestitis  pilis  et  hirsutulis  albis  gracilibus  et  paucis  capitato- 
glandulosis;  foliis  plerumque  dispersis,  0.5-1.5  cm.  longis,  0.2-0.8  cm.  latis,. 


VoL.XXV]  HOWELL:  PHACELIA  SECTION  MILTITZIA  371 

elliptico-oblongis  ovatis  obovatisve,  integris  vel  tenuiter  creiiulatis  vel  promi- 
nenter  serrato-lobatis,  apice  obtusis,  cuneatis  basi,  plus  minusve  hirsutulis, 
petiolo  ad  2  cm.  longo,  saepe  quam  lamina  longiore;  racemis  circa  2  cm.  longis, 
saepe  f  oliis  brevioribus,  floribiis  subcongestis  vel  infimis  distantibus,  pedicellis 
plerumque  erectis  vel  adscendentibus,  ad  6  mm.  longis ;  floribus  5-meris ;  seg- 
mentis  calycis  ad  anthesin  3.5-5  mm.  longis,  0.3--0.5  mm,  latis,  in  fructu  valde 
accrescentibus,  6-10  mm.  longis,  0.3-1  mm.  latis,  hirsutulis  et  capitato-glandu- 
losis,  basi  calcarif ormi-angulata,  costa  prominenti ;  corolla  tubulata,  3.5-4.5 
mm.  longa,  dilute  lutea,  baud  purpureo-tincta,  pubescenti  extus,  intus  glabra, 
lobis  1  mm.  longis,  crenulatis ;  squamis  ad  lineas  reductis  vel  nullis ;  sta- 
minibus  inclusis,  2.2-2.75  mm.  longis,  filamentis  glabris,  antlieris  0.3  mm. 
longis,  latioribus  quam  longioribus  explanatis;  stylo  et  ramis  1.25-1.5  mm. 
longis,  stylo  paulum  pilifero  tantum  prope  basin,  ramis  0.2-0.25  mm.  longis; 
ovario  0.75-1  mm.  longo,  hirsutulo  supra,  ovulis  10  vel  11 ;  eapsula  3-5  mm. 
longa,  elliptica  vel  elliptico-ovata,  apice  rotundata,  mutica  vel  brevissime 
apiculata,  longitudinaliter  dorso  valvarum  sulcata,  sparse  hirsutula  superne; 
seminibus  1.5-2  mm.  longis,  subovatis  vel  ovato-oblongis,  obtusis,  nigro-b run- 
nels, paulo  angulatis,  prominenter  transverse  corrugatis,  rugis  circa  12,  pro- 
funde  alveolato-reticulatis. 

Type:  No.  163032  in  Rocky  Mountain  Herbarium,  University  of  Wyoming, 
collected  on  May  19, 1911,  at  DeBeque,  Mesa  County,  Colorado,  by  George  E. 
Osterliout,  No.  4458.  There  is  a  duplicate  in  Herb.  N.Y.  Bot.  Gard.  Phacelia 
suhmutica  is  known  only  from  Osterhout's  collections  made  at  DeBeque,  others 
being :  No.  4726  (RM,  frag.  CAS)  and  No.  5978  (RM,  frag.  CAS) . 

Although  P.  suhmutica  is  obviously  related  to  P.  scopulina  from  which  it 
was  undoubtedly  derived,  it  may  be  readily  differentiated  by  a  number  of 
distinctive  characters  that  indicate,  each  in  a  small  way,  a  separate  specific 
entity.  Phacelia  suhmutica  has  a  rather  distinctive  appearance  which  comes 
from  its  less  leafy  and  more  fiorif erous  habit  and  which  is  also  due  to  the  less 
cinereous  character  of  the  pubescence.  In  P.  suhmutica  the  fruiting  calj^x- 
segments  are  usually  much  more  elongate ;  and  while  the  pedicels  of  the  lower 
flowers  are  reflexed  or  sigmoid-spreading  in  fruit  in  P.  scopulina,  they  are 
mostly  strictly  erect  or  ascending  in  P.  suhmutica.  There  is  a  noteworthy 
difference  in  the  coloration  of  the  corolla  in  the  two  species,  too,  that  of  P. 
suhmutica  being  yellowish,  that  of  P.  scopulina  being  more  or  less  strongly 
tinged  in  age  with  lavender  or  violet.  The  stamens  in  P.  scopulina  equal  or 
exceed  the  corolla-throat,  sometimes  in  anthesis  even  exceeding  the  corolla- 
lobes,  while  in  P.  suhmutica  the  stamens  are  shorter  than  the  corolla-throat. 
Although  all  these  characters  indicate  real  differences  between  P.  suhmutica 
and  P.  scopulina,  they  are  not  so  important  either  specifically  or  diagnostically 
as  the  characters  that  are  found  in  the  style,  capsule,  and  seed.  In  P.  suhmutica 
the  style  is  glabrous  except  for  a  few  hairs  near  the  base,  but  in  P.  scopulina 
the  style  is  hirsutulous-hairy  to  the  middle  or  above,  generally  densely  so. 
The  capsule  of  P.  suhmutica  differs  from  the  capsules  of  other  species  in  this 


■M2  CALIFORNIA   ACADEMY   OF  SCIENCES        [Proc.  4th  Ser. 

section  except  P.  tetramera,  in  that  it  is  either  entirely  devoid  of  a  terminal 
apiculation  or  has  a  very  insignificant  one.  By  contrast,  the  capsule  of  P. 
scopulina  is  prominently  apiculate.  The  differences  in  the  seeds,  although 
small,  are  nevertheless  appreciable:  the  seeds  of  P.  suhmutica  are  usually 
larger  and  more  finely  and  numerously  corrugated,  the  corrugations  are  not  so 
i-ounded  and  are  more  deeply  alveolate-pitted,  and  the  seed-cover  is  blackish- 
brown  and  tends  to  be  iridescent. 

Hence,  although  P.  suhmutica  is  undoubtedly  closely  related  to  P.  scopulina, 
I  believe  that  it  is  an  entity  sufficiently  distinguished  to  be  accorded  specific 
recognition. 

7.  Phacelia  inyoensis  (Macbr.)  J.  T.  Howell 

Leali.  West.  Bot.  4:16  (1944) 
Miltitzia  inyoensis  Macbr.,  Contrib.  Gray  Herb.,  n.  ser.,  49:41  (1917). 

Stems  erect  or  spreading,  slender,  one  to  several  from  the  top  of  the  taproot, 
2-10  cm.  tall,  hirsutulous  and  finely  capitate-glandular ;  leaves  few  and  scat- 
tered, not  concealing  the  flowers,  0.5-1.5  cm.  long,  0.3-1  cm.  wide,  oblong  to 
elliptic  or  obovate,  entire  to  coarsely  and  pinnately  few-lobed,  subobtuse  at 
apex,  euneate  at  base,  hirsutulous  and  more  or  less  finely  capitate-glandular, 
petioles  of  basal  leaves  to  2  cm.  long,  those  of  the  uppermost  leaves  only  about 
3  mm.  long ;  racemes  lax,  loosely  few-flowered,  to  7  em.  long,  the  lowest  flowers 
distant,  the  uppermost  approximate,  pedicels  slender  and  spreading,  2—6  mm. 
long,  hirsutulous  and  glandular;  flowers  5-merous;  calyx-segments  in  flower 
a  little  unequal,  2-3  mm.  long,  0.4-0.75  mm.  wide,  linear  to  linear-oblanceolate, 
in  fruit  3.75-6  mm.  long,  0.5-1.3  mm.  wide,  linear-oblong  to  oblanceolate, 
somewhat  spatulate,  narrowed  below  to  claw-like  base,  subacute,  glandular 
and  hirsutulous ;  corolla  pale  yellow  becoming  tinged  with  violet,  tubular-  or 
cylindric-campanulate,  3-3.5  mm.  long,  lobes  erect,  0.6-1  mm.  long,  corolla- 
tube  hairy  outside  to  the  base,  glabrous  inside;  corolla-scales  obsolete  or  pres- 
ent as  a  mere  line  about  1  mm.  long,  free  from  the  base  of  the  filaments; 
stamens  1.5-2.5  mm.  long,  filaments  glabrous,  anthers  about  0.2  mm.  long, 
wider  than  long  after  dehiscence;  style  and  branches  1  mm.  long,  branches 
0.2-0.25  mm.  long,  style  glabrous  except  at  the  very  base ;  ovary  1  mm.  long, 
hairy  above,  ovules  18-20;  hypogynous  disk  relatively  prominent  with  round- 
ish lobes  between  the  stamens ;  capsule  3.3-4  mm.  long,  elliptic,  rounded- 
obtuse,  shortly  apiculate,  sparsely  hirsutulous,  valves  shallowly  longitudinally 
sulcate  on  the  back;  seeds  nearly  1  mm.  long,  ovatish  or  deltoid-ovate,  acute 
at  apex,  rounded  at  base,  brownish,  corrugations  rather  i^rominent,  reticulate, 
6  or  7. 

Known  only  from  Owens  Valley,  Inyo  Count3^  California. 

Collectio')Ks.  Foothills  west  of  Bislioi),  Heller  No.  8324  (G,  type;  CAS,  DS, 
ITS)  ;  Big  Pine,  K.  Brandegee  in  1913  (P)  ;  Lone  Pine,  K.  Brandegee  in  1913 
(CAS,  P). 

According  to  data  accompanying  Heller's  collection,  the  habitat  in  which 


I 


» 


VOL.XXV]  HOWELL:  PHACELIA  SECTION  MILTITZIA  373 

this  rare  species  grows  is  "a  small  dry  peat  bog  among  the  granite  sands." 
.Vlthoiigh  it  is  quite  a  distinct  species,  it  is  probably  most  closely  related  to 
P.  saliiia  and  through  it  to  P.  scojmlina. 

8.  Phacelia  salina  (A.  Nels.)  J.  T.  Howell 

Leafl.  West.  Bot.  4:16  (1944) 

Emmenanthe  foliosa  M.  E.  Jones,  Zoe  4:278  (1893). 

Emmenanthe  salina  A.  Nels.,  Bull.  Torr.  Bot.  Club  25:381  (1898). 

Miltitzia  foliosa  (M.  E.  Jones)  Brand,  Das  Pflanzenr.  IV.  251 :131  (1913). 

Miltitzia  salina  (M.  E.  Jones)  Eydb.,  Bull.  Torr.  Bot.  Club  40  :479  (1913). 

Xot  Phacelia  foliosa  Phil.,  Anal.  Mus.  Nac.  Chile  53  (1891). 

Xot  Fliacelia  salina  M.  E.  Jones  ex  Brand,  Das  Pflanzenr.  IV.  251 :  119  (1913),  as  synonym. 

Stems  short  and  stout  or  slender  and  spreading,  2  to  several  from  the  top 
of  a  slender  annual  taproot,  0.5-3  cm.  or  even  to  6  cm.  long,  sparsely  hirsutu- 
lous,  eglandulose  or  minutely  and  sparsely  capitate-glandular  above  and  in 
the  inflorescence ;  leaves  entire,  sinuate-wavy,  serrate  or  pinnately  f ew-lobed, 
0.5-1.5  cm.  long,  0.3-0.9  cm.  wide,  oblanceolate  to  elliptical  or  obovate, 
sparsely  hirsutulous,  obtuse,  narrowly  to  broadly  cuneate  at  base,  frequently 
a  little  asymmetrical  at  base,  petioles  0.5-3  cm.  long;  racemes  exceeded  by 
the  leaves,  laxly  few-flowered  and  only  slightly  scorpioid,  the  flowers  distant 
or  tending  to  be  congested  near  the  ground  at  the  base  of  the  stems,  pedicels 
1-5  mm.  long,  straight  or  spreading;  flowers  5-merous;  calyx-segments  in 
flower  2.5-4  mm.  long,  in  fruit  4.5-6  mm.  long,  0.5-0.75  mm.  wide,  linear  to 
linear-oblong  or  linear-oblanceolate,  sparsely  hirsutulous  and  sometimes  a 
little  glandular ;  corolla  yellow  or  tinged  with  lavender,  tubular,  3-4  mm.  long, 
the  lobes  erect,  0.9-1.25  mm.  long,  the  corolla  hairy  outside,  glabrous  within ; 
corolla-scales  generally  lacking,  if  present  filiform  and  about  1.5  mm.  long, 
at  base  free  from  the  filaments ;  stamens  1.5-2.5  mm.  long,  filaments  glabrous, 
anthers  about  0.25  mm.  long,  roundish ;  style  and  branches  1  mm.  long,  the 
branches  0.2-0.25  mm.  long,  style  hairy  only  near  the  base  or  to  near  the 
middle;  ovary  0.75-1  mm.  long,  ovules  7-9 ;  hypogynous  disk  with  relatively 
prominent  rounded-deltoid  lobes,  the  lobes  alternating  with  the  stamens  and 
about  0.2  mm.  long ;  capsule  3-4  mm.  long,  broadly  elliptical  or  elliptic-ovate, 
obtuse  and  apiculate,  sparsely  hairy,  the  valves  thin-chartaceous  with  thick 
conspicuous  margins,  back  of  the  valves  shallowly  sulcate;  seeds  brown,  1.5--2 
mm.  long,  acute  or  subacute  at  both  ends,  oblongish  to  ovatish,  prominently 
corrugated,  corrugations  9-11,  broad  and  rounded. 

Of  rare  and  local  occurrence  in  alkaline  soil  in  southern  Wyoming,  western 
Utah,  and  northern  Nevada. 

Collections.  Bitter  Creek,  Sweetwater  Co.,  Wyoming,  A.  Nelson  No.  3105 
(RM,  type  of  Emmenanthe  salina;  CAS,  G,  M,  US),  A.  Nelson  No.  4786 
(NY)  ;  Deep  Creek  Valley  above  Furber,  Tooele  Co.,  Utah,  M.  E.  Jones  in 
1891  (P,  type  of  Emmenanthe  foliosa;  CAS,  DS,  G,  M,  UC,  US)  ;  in  loose 
red  sand,  west  end  of  White  Pine  Mts.  below  Little  Antelope  Summit,  White 
Pine  Co.,  Nevada,  Ripley  and  Barnehy  No.  3592  (CAS)  ;  Reese  River  Valley  6 


374  CALIFORNIA  ACADEMY  OF  SCIENCES        [Proc.  4th  Ser.     > 

miles  nortliAvest  of  Austin,  Lander  Co.,  Eipley  and  Barneby  No.  5926  (CAS) ; 
ea!?t  base  of  Railroad  Pass,  Lander  Co.,  Ripley  and  Barnehy  No.  5932  (CAS). 

There  is  considerable  variation  in  the  data  accompanying  the  original  col- 
lection of  Emmenanthe  foliosa.  The  specimen  in  the  Jones  Herbarium  at 
Pomona  College  which  I  am  accepting  as  the  tj'^pe  gives  "Furber,  Nevada," 
June  8, 1891,  while  a  second  sheet  gives  "Deep  Creek,  alkaline  flats  above  Fur- 
ber,  Nevada."  Data  on  specimens  in  Herb.  Calif.  Acad.  Sci.  and  U.S.  Nat. 
Herb,  are  "On  Deep  Creek,  above  Furber,"  Utah,  June  8,  1891 ;  and  in  Gray 
Herbarium  in  Jones'  handwriting  are  "Deep  Creek,  Utah,"  June  4,  1891.  The 
data  given  in  the  original  description  are  still  different :  "Deep  Creek,  Utah, 
June  6,  1891,  altitude  5000  feet."  After  a  careful  study  of  the  plants  consti- 
tuting these  specimens,  I  have  concluded  that  all  are  parts  of  one  collection ; 
and  by  reference  to  the  "Botanical  Exploration  of  Marcus  E.  Jones  from  1875 
to  1919"  (Ms.),  I  believe  the  correct  data  for  the  collection  to  be:  Alkaline 
flats.  Deep  Creek  Valley  above  (i.e.  south  of)  Furber,  Tooele  Co.,  Utah,  June  8, 
1891.  According  to  his  journal  (I.e.,  p.  19),  Jones  was  at  Fish  Springs  on 
June  4,  at  Ibapah  on  June  6,  and  at  Furber  on  June  8. 

Phacelia  salina  is  intermediate  between  P.  scopulina  and  P.  tetramera,  the 
three  species  apparently  representing  a  distinct  phylogenetic  line.  Morpho- 
logically P.  salina  seems  more  closely  related  to  P.  scopulina,  differing  chiefly 
in  the  smaller  size  of  most  parts;  but  ecologically  it  is  much  nearer  P.  tetra- 
mera, both  species  being  usually  restricted  to  highly  alkaline  or  mineralized 
situations  in  desert  regions.  Phacelia  salina,  P.  inyocnsis,  P.  glaherrima,  and 
P.  suhmutica  are  to  be  counted  among  the  rarest  of  western  plants. 

9.  Phacelia  tetramera  J.  T.  Howell 

Leafl.  West.  Bot.  4:16  (1944) 

Emmenanthe  pusilla  A.  Gray,  Proc.  Amer.  Acad.  11:87  (1876). 
Miltitzin  pusilla  (A.  Gray)  Brand,  Das  Pflanzenr.  IV.  251:132  (1913). 
Miltitzia  pusilla  (A.  Gray)  Brand  var.  flagellaris  Brand,  I.e. 

Not  Phacelia  pusilla  Buckl.,  Amer.  Jour.  Sci.  45:172   (1843);  nor  P.  pusilla  Torr.  ex  S. 
Wats.,  Bot.  U.S.  Geo].  Explor.  40th  Paral.  (King's  Exped.)  253  (1871). 

Stems  several  from  the  top  of  the  taproot,  spreading  or  ascending,  2-10  cm. 
long,  slender,  finely  glandular- villous  and  near  the  base  sparsely  hirsutulous ; 
leaves  narrowly  elliptic-oblong  to  oblong-oblaneeolate,  0.4-1.5  em.  long,  0.2- 
0.7  cm.  wide,  entire  or  saliently  few-serrate-lobed,  obtuse,  narrowly  cuneate  at 
base,  sparsely  hirsutulous,  petiole  0.4-1.5  cm.  long,  glandular-villous ;  racemes 
to  8  cm.  long,  very  laxly  flowered,  the  lowest  flowers  distant,  pedicels  to  4  mm. 
long,  slender,  recurved  or  spreading  in  the  lower  flowers ;  flowers  usuallj^ 
4-merous,  sometimes  5-merous ;  calyx-segments  in  flower  2.5  mm.  long,  oblong 
or  narrowly  oblong-ovate,  in  fruit  unequal  in  width,  4r-5  mm.  long,  0.5-1  mm. 
wide,  narrowly  to  broadly  oblong,  hirsutulous  and  sparsel.y  and  minutely 
glandular,  obtuse,  united  at  base  sometimes  as  much  as  2  mm.,  the  base 
cuneate;  corolla  whitish,  1.3-1.8  mm.  long,  eampanulate,  the  lobes  erect,  0.5-1 
mm.  long,  the  lobes  sparselj^  hairy  outside,  glabrous  inside;  corolla-scales 


Voi..XXV|  nOWELL:  PHACELIA   SECTION  MILTITZIA  375 

none ;  stamens  1  mm.  long-,  filaments  glabrous,  anthers  ronndish,  nearly  0.2 
mm.  across ;  style  0.25-0.4  mm.  long,  the  branches  indicated  only  by  an  emar- 
gination  at  the  top  of  the  style;  ovary  roundish,  0.75  mm.  long,  hirsutulous  at 
top,  ovules  20;  hypogynous  disk  narrow  and  inconspicuous;  capsule  oblong 
to  oblong-obovate,  3-4  mm.  long,  very  obtuse,  shortly  apiculate  or  muticous, 
sparsely  hirsutulous,  the  verj^  base  sometimes  a  little  inferior ;  seeds  1  mm. 
long,  narrowly  ovate  to  quadrate,  round  at  base,  acute  at  apex,  light  brown, 
finely  and  sometimes  indistinctly  corrugated,  corrugations  about  7-9. 

Alkaline  flats  and  washes  of  high  valleys,  eastern  Oregon  south  to  central 
Nevada  and  eastern  California. 

B.epreseyitative  collections.  Oregon:  Union  Co.,  Cnsick  No.  758  (CM,  G, 
US  ;  first-cited  collection  of  Miltitzia  pusilla  var.  flagellaris)  ;  Malheur  River, 
Harney  or  Malheur  Co.,  Cusick  No.  1946  (CM,  G,  M,  ND,  RM,  UC,  US,  WS) ; 
Wagontire,  Harney  Co.,  Peck  No.  20844  (W)  ;  8  miles  west  of  Lakeview,  Lake 
Co.,  Peck  No.  15243  (CAS,  NY,  W). 

Nevada  :  8  miles  north  of  Twin  Bridges,  Elko  Co.,  Holmgren  No.  871  (lU, 
UC);  Steamboat  Springs,  Washoe  Co.,  8.  Watson  (G,  type  of  Emmenanthe 
pusilla;  NY)  ;  Washoe  Lake,  Washoe  Co.,  M.  E.  Jones  in  1897  (CAS,  CM, 
DS,  P,  US)  ;  Empire  City,  Ormsby  Co.,  M.  E.  Jones  No.  3987  (CAS,  DS, 
lU,  M,  NY,  P,  RM,  UC,  US);  foothills  of  Toquima  Range  near  Belmont, 
Nye  Co.,  Ripley  and  Barnehy  No.  3696  (CAS)  ;  northwestern  Nevada,  Lem- 
mon  in  1875  (G)  ;  Grass  Valley,  Lander  Co.,  Ripley  and  Barnehy  No.  5920 
(CAS). 

California  :  "Sierra  Nevada  Mts.,"  Lemmon  in  1875  (US)  ;  Chat,  Plumas 
Co.,  M.  E.  Jones  in  1897  (CAS,  P)  ;  Long  Valley,  Mono  Co.,  J.  T.  Howell  No. 
14382  (CAS,  CM,  G,  lU,  P,  US)  ;  valley  east  of  Cedarville,  Modoc  Co.,  Ripley 
and  Barnehy  No.  6005  (CAS) . 

The  earliest  collection  of  this  species,  Watson  No.  878  from  Steamboat 
Springs,  was  included  by  Torrey  in  hLs  Phacelia  pusilla;  and,  although  Gray 
cited  Watson's  collection  in  liis  original  description  of  Emmenanthe  pusilla, 
Brand  annotated  it  in  the  U.S.  National  Herbarium  as  Phacelia  pusilla  Torr. 
and  cited  it  as  that  species  in  his  monograph  (1913,  p.  122) . 

The  most  remarkable  feature  of  P.  tetramera  is  the  tendency  for  the  flowers 
to  be  4-merous  instead  of  5-merous  as  is  usual  in  Phacelia.  The  reduction  in 
the  number  of  parts  seems  to  result  from  the  suppression  of  one  part  of  the 
calyx,  corolla,  and  androecium,  and  not  to  be  due  to  the  union  of  adjacent 
parts.  Although  most  of  the  flowers  that  have  been  examined  are  truly  4- 
merous,  occasionally  they  are  either  partly  or  complete! j^  5-merous.  In  some 
5-merous  flowers,  the  fifth  calyx-segment  is  much  smaller  than  are  the  other 
four ;  in  one  case  two  calyx-segments  were  united  nearly  throughout,  while 
the  other  three  were  deeply  divided.  It  has  been  noted  that  5-merous  flowers 
iu-e  more  numerous  in  specimens  from  Oregon  than  in  those  from  California 
(vr  Nevada.  Certainly  the  reduction  of  flower-parts  exhibited  by  P.  tetramera 
is  a  most  unusual  tendency  in  Hydrophyllaceae . 


;576  CALIFORNIA   ACADEMY  OF  SCIENCES        [Proc.  4tii  Skk. 

REFERENCES 

Brand,  A. 

1913.  Hjdrophyllaceae  in  A.  Engler,  Das  Pflanzenreich  IV.  251  (Heft  59).  210  pp.  Wil- 
helm  Engelmann,  Leipzig. 
Gray,  A. 

1878.  Synoptical  Flora  of  North  America,  vol.  2,  pt.  1,  393  pp.  Ivisoii,  Blakeinan,  Taylor, 
and  Co.,  New  York. 
Howell,  J.  T. 

1944.  A  reconsideration  of  the  genus  Miltitzia.  Leafl.  West.  Bot.  4:12-16. 
Jepson,  W.  L. 

1943.  Flora  of  California,  vol.  3,  pt.  2,  pp.  129-464.  Associated  Students  Store,  University 
of  California,  Berkeley. 
Kearney,  T.  H.,  and  E.  H.  Peebles 

1942.  Flowering  plants  and  ferns  of  Arizona.  U.S.  Dept.  Agric.  Miscell.  Publ.  No.  42:>. 
1069  pp.  Government  Printing  Office,  Washington,  D.C. 
Peck,  M.  E. 

1941.  Manual  of  the  higher  plants  of  Oregon.  866  pp.  Binfords  and  Mort,  Portland, 
Oregon. 

TiDESTROM,  I.  I 

1925.  Flora  of  Utah  and  Nevada.  Contrib.  U.S.  Nat.  Herb.  25.  665  pp.  Government  Print- 
ing Office,  Washington,  D.C.  ^ 


PROCEEDINGS 

OF  THE 


Marine  Biological  LabordWiy  | 

X*  I  33  K.  -A-  R  "ST 

JAN  3  1 1947 

WOODS  HOLE,  MASS. 


CALIFORNIA  ACADEMY  OF  SCIENCES 

Fourth  Series 
Vol.  XXV,  No.  i6,  pp.  377-420,  pis.  37-46  November  15,  1946 


ALICE  EASTWOOD 
SEMI-CENTENNIAL  PUBLICATIONS 

No.  16 

ENDEMISM  IN  THE  HAWAIIAN  FLORA,  AND  A 

REVISION  OF  THE  HAWAIIAN  SPECIES  OF 

GUNNERA  (HALORAGIDACEAE) 

HAWAIIAN  PLANT  STUDIES  U' 

BY 

HAROLD  ST.  JOHN 

Professor  of  Botany,  University  of  Hawaii 


E 


ENDEMISM  IN  THE  HAWAIIAN  FLORA 

NDEMisM  in  its  flora  has  long  made  the  plants  of  the  Hawaiian  Islands 


famous.  Of  the  islands  or  floristic  regions  of  the  world  having  a  flora  of 
more  than  a  few  species,  they  are  said  to  have  the  world's  largest  percentage 
of  endemics. 

Prof.  A.  Guillaumin  (Proc.  3rd.  Pan.  Pacif.  Sci.  Congr.  Tokyo  1 :  930, 1926) 
in  his  review  of  the  Pacific  island  floras  calculates  that  endemism  for  the 
Hawaiian  Phanerogams  is  66  per  cent.  He  does  not  state  how  he  arrived  at 
this  figure  but  his  totals  of  the  species  are  evidently  those  of  Wm.  Hillebrand. 
This  early  botanist,  Hillebrand  (PI.  Hawaiian  Ids.  p.  XVII,  1888),  calculated 
that  of  the  native  Phanerogams,  574  were  endemic,  while  246  only  were  in- 
digenous, and  24  of  early  aboriginal  introduction,  and  the  total  of  native 
Phanerogams  and  Vascular  Cryptogams  was  860.  It  is  apparent  that  Guil- 
laumin used  this  total  of  860  indigenous  Vascular  Plants  and  Hillebrand's 
figure  of  574  indigenous  Phanerogams  in  computing  the  percentage  of  en- 

^  This  is  the  eleventh  of  a  series  of  papers  designed  to  present  descriptions,  revisions,  and 
records  of  Hawaiian  plants.  The  preceding  papers  have  been  published  as  Occasional  Papers 
Bishop  Museum  10(4),  1933;  10(12),  1934;  11(14),  1935;  12(8),  1936;  14(8),  1938; 
15(1), 1939;  15(2), 1939;  15(22), 1940;  15(28), 1940;  17(12), 1943;  Lloydia  7:265-274, 
1944;  Bull.  Torrey  Bot.  Club  72:22-30,  1945. 

[377] 


378  CALIFORNIA  ACADEMY  OF  SCIENCES  [Proc.  4th  Ser. 

demism,  which  he  calculated  as  66  per  cent.  Had  he  used  705  native  species 
and  574  endemics,  which  are  Ilillebrand's  truly  comparable  figures  for  the 
Phanerogams,  the  endemism  would  figure  out  as  81  per  cent,  which  was  the 
estimate  of  Hillebrand,  previous  to  his  death  in  1886.  All  species  published 
since  that  date  or  unknown  to  Hillebrand,  Guillaumin  seems  to  reject  by  his 
phrase  that  the  area  presents  an  extraordinary  number  of  varieties  which  are 
sometimes  considered  as  species  of  narrow  geographic  localization  ("presente 
un  nombre  extraordinaire  de  varietes,  considerees  parf ois  des  comme  especes  et 
etroitement  localisees.").  D.  H.  Campbell  used  the  same  figures  from  Hille- 
brand, but  he  correctly  calculated  the  endemism  of  Phanerograms  as  81.42  per 
cent  (The  Derivation  of  the  Flora  of  Hawaii,  Univ.  Ser.  Stanford  Univ.,  Publ. 
15,  1919).  Also  R.  Kanehira  apparently  used  the  same  figures  as  he  lists  the 
Hawaiian  endemism  as  81  per  cent  (Proc.  6th  Pacif.  Sci.  Congr.  4  :610, 1940). 
Guillaumin  concludes  (pp.  921,  931)  that  in  the  Hawaiian  flora  the  Australian, 
New  Zealand,  and  even  Polynesian  elements  are  almost  completely  lacking, 
that  the  flora  has  strong  relationships  with  the  American  flora  and  should  be 
considered  a  part  of  the  IMexican  floristic  region.  The  present  writer  disputes 
all  of  these  conclusions  of  Guillaumin. 

The  interpretations  of  C.  Skottsberg  seem  more  truly  in  accord  with  the 
facts.  He  divided  the  Hawaiian  species  of  Phanerogams  (Bull.  Bishop  Mus. 
16  :  6,  1925)  as  32.3  per  cent  Old-Pacific ;  29  per  cent  Australian-Polynesian ; 
26.5  per  cent  Indo-Malayan ;  7.5  per  cent  American;  3.3  per  cent  widespread ; 
1.2  per  cent  Subantarctic ;  and  0.2  per  cent  Boreal.  The  widespread  or  Pan- 
tropic  and  the  Subantarctic  may  well  have  immigrated  by  the  same  route  as 
the  other  groups  from  Oceania.  If  these,  then,  are  all  added,  the  total  is  92.3 
per  cent  for  the  plants  with  a  probable  southwestern  origin  that  came  or  whose 
ancestors  came  as  a  stream  of  plant  immigrants  through  Polynesia.  Later 
Skottsberg  discusses  the  genera  (Proc.  Linn.  Soc.  151 :  182-186,  1939),  their 
relationship,  and  probable  route  of  immigration.  Of  more  than  200  genera, 
18  are  Pantropic,  15  Paleotropic,  49  Malesian,  12  littoral  are  widespread,  57 
Austral,  22  Boreal,  9  aquatics  apparently  Boreal,  and  13  Neotropical.  This 
review  of  the  Hawaiian  genera  gives  the  same  picture  as  with  the  species :  a 
large  majority  Indo-Malayan  or  Polynesian,  small  minorities  Boreal  and 
American. 

The  same  early  source,  Hillebrand,  was  used  by  J.  C.  Willis  (Proc.  Linn. 
Soc.  London  148:86-91,  1936)  in  his  contemporary  discussion  of  regional 
endemism.  Hillebrand's  figures  should  not  be  discounted.  They  were  based 
on  his  excellent  flora  and  were  correct  for  his  time,  that  is  58  years  ago.  Still,  to 
discuss  the  endemism  of  the  Hawaiian  flora  today  and  to  use  only  Hillebrand's 
figures  is  not  the  modern  scientific  method,  which  should  use  all  available 
evidence.  Since  1886  in  Hawaii  there  has  been  a  great  deal  of  arduous  explora- 
tion of  the  rain  forests  and  the  precipitous  mountains.  Here  and  elsewhere 
a  large  amount  of  floristic  and  monographic  work  has  been  published.  To  over- 
look and  reject  all  the  work  of  C.  B.  Clarke,  Heller,  Rock,  C.  N.  Forbes, 


Vol.  XXV]       ST.  JOHN:  ENDEMISM  IN  HAWAIIAN  FLORA.  GUNNEBA         379 

Beccari,  Leveille,  Skottsberg,  Cliristensen,  Degener,  St.  John,  Sherff ,  Yuncker, 
Christophersen,  Caum,  Hitchcock,  Hochreutmer,  Pilger,  von  Poelhiitz,  Radl- 
kofer,  Oliver,  Danser,  Heimerl,  Keck,  Krajina,  Whitney,  Hosaka,  Fosberg, 
Egler,  Lam,  Croizat,  Sleumer,  and  others,  is  not  the  way  to  arrive  at  a  modern 
estimate  of  the  known  Hawaiian  flora.  Skottsberg  used  the  more  sound  method 
of  including  subsequent  publications  (Bull.  Bishop  Mus.  16:5,  1925)  when 
he  gave  a  tentative  estimate  of  the  indigenous  Hawaiian  Phanerogams  as  about 
900  species.  Later  he  revised  this  figure  to  between  1,000  and  1,100  (Proc.  6th 
Pacif.  Sei.  Congr.  Berkeley  4 :  685,  1940) .  The  writer  keeps  a  check  list  of  the 
Hawaiian  flora.  There  are  quite  a  number  of  the  recently  proposed  segregate 
species  which  he  has  not  included,  awaiting  time  to  verify  their  status,  but 
many  will  doubtless  be  accepted.  When  eventually  added  in,  they  will  increase 
rather  than  diminish  the  percentage  of  endemism.  At  present  the  writer's 
figures  for  endemism  of  Hawaiian  Phanerogams  is  90  per  cent,  based  on  totals 
of  1795  indigenous,  and  1614  endemic  species  and  their  subdivisions.  Esti- 
mates by  some  others  exclude  all  subdivisions  of  species,  but  here  each  is  in- 
cluded as  a  unit. 

Some  Hawaiian  genera  contain  species  that  are  homogeneous  and  wide- 
spread, occurring  unmodified  on  all  or  nearly  all  of  the  large  islands.  Ex- 
amples are  given  in  the  two  following  lists. 

I.  Homogeneous  Widespread  Species  in  Genera  Monottpic  in  the  Hawaiian  Islands 

Freycinetia  arhorea  Gaud. 

Joinvillea  GaudicJiaudiana  Brogn.  and  Gris 

Dianella  sandwicensis  H.  and  A.^ 

AnoectocMlus  sandwicensis  Lindl. 

Liparis  Jiawaiiensis  Mann 

Hahenaria  holochila  Hbd. 

Pilea  peploides  H.  and  A. 

Argemone  alba  Lestib.  var.  glauca  Prain 

Osteomeles  anthyllidi folia  Lindl. 

Erythrina  sandwicensis  Degener 

Strongylodon  lucidus  Seem. 

Plumiago  seylanica  L. 

Osmanthus  sandwicensis  (Gray)  B.  and  H. 

Nertera  depressa  Banks  and  Soland. 

II.  Homogeneous  Widespread  Species  in  Genera  with  Several  Hawaiian  Species 

Smilax  sandwicensis  Kunth 
Chenopodium  oahuense  (Meyen)  Aellen^ 
Peperomia  leptostachya  H.  and  A. 

There  are  other  Hawaiian  species  widespread  among  the  larger  islands,  but 
which  are  not  homogeneous,  having  a  tendency  to  vary.  These  variations  are  in 
part  recognized  as  described  varieties  but  in  large  part  are  unrecognized, 

2  The  recently  described  Dianella  lavarum  Degener  and  D.  multipedicellata  Degener  are 
considered  exact  synonyms. 

3  Chenopodium  sandwicheum  Moq.  forma  macrospermum  Aellen  and  forma  microspermum 
Aellen  are  not  separable,  as  the  larger  and  smaller  seeds  occur  on  a  single  plant. 


380  CALIFORNIA  ACADEMY  OF  SCIENCES  [Proc.  4th  Ser. 

forming  a  part  of  the  heterogeneous  population  now  included  in  the  species. 

Such  ones  are  listed  below. 

TIT.  Polymorphic  Widespread  Species 
Eumex  giganteus  Ait. 
Charpentiera  oiovata  Gaud. 
Charpentiera  ovata  Gaud. 
Broussaisia  arguta  Gaud. 
Acacia  Koa  Gray 
Ilex  anomala  H.  and  A. 
Cheirodendron  Gaudichaudii  (DC.)  Seem. 
Vacci7iium  calycinum  Sm. 
Vacciniutn  dentatum  Sm. 
Styphelia  Tameiameiae  (Cham.)  F.  Muell. 
Myrsine  Lessertiana  A.DC. 
Gouldia  terminalis  (H.  and  A.)  Hbd. 

Other  genera,  usually  with  many  species,  show  a  conspicuous  segregation 

into  species,  usually  each  species  being  restricted  to  a  single  island  or  even  to 

a  single  mountain  range  or  valley  on  the  particular  island.  Such  genera  are 

the  following : 

IV.  Genera  with  Numerous  Species  that  are  Narrow  Endemics 

Astelia,  Schiedea,  Pelea,  Fagara,  HibisctirS,  Viola,  Labordia,  Haplostachys,  Phyllos- 
tegia,  Stenogyne,  Cyrtandra,  Coprosma,  Hedyotis,  Clermontia,  Cyanea,  Delissea, 
Lobelia,  Bidens,  Bubauiia,  and  Lipochaeta. 

The  Hawaiian  plants  here  listed  illustrate  a  progressive  series :  1,  genera 
monotypic  in  Hawaii  occurring  as  invariable  wides  on  all  the  principal 
islands ;  2,  species  that  are  invariable  wides,  belonging  to  genera  with  several 
Hawaiian  species;  3,  species  that  are  wides,  but  showing  variabilitj^  on  the 
various  islands ;  and  4,  general  with  many  Hawaiian  species,  typically  with 
different,  distinct  species  on  each  island.  These  species  represent  degrees  of 
increasing  differentiation  and  presumably  of  decreasing  age.  Thej'  show  evi- 
dences of  speciation.  The  results  of  this  species  formation  can  be  seen  in  the 
percentages  of  endemism  here  presented  and  discussed. 

REVISION  OF  THE  HAWAIIAN  SPECIES  OF  GUNNERA 

One  of  the  most  famous  of  the  genera  occurring  in  the  Hawaiian  Islands  is 
Gunnera,  called  by  the  natives  "apeape."  It  is  a  giant  herb  with  a  stem  12  to 
15  cm.  or  more  in  diameter,  fleshy,  2  to  6  meters  tall,  the  base  decumbent  and 
prostrate.  The  paired  stipules  are  large,  pale,  and  conspicuous.  From  near  the 
summit  are  produced  several  alternate  leaves  on  fleshy  petioles  6  to  13  dm. 
long.  The  blades  are  rounded,  shallowly  lobed,  like  great  parasols  4  to  20  dm. 
in  diameter.  Axillary  panicles  8  to  10  dm.  long  bear  numerous  minute  green 
flowers  and  later  tiny  yellow,  orange,  red,  or  purplish  drupes.  The  plant  itself 
is  so  bizarre  as  to  hold  the  attention  and  interest  of  all  observers,  whether 
scientific  or  not.  Thus  its  occurrence  is  shown  even  on  the  tourist  maps. 

The  habitat  of  the  "apeape"  is  also  noteworthy.  It  occurs  on  precipitous 
slopes  with  the  soil  saturated  from  the  run-off  of  the  boggy  mountain  summits. 


Vol.  XXV]        ST.  JOHN:  ENDEMISM  IN  HAWAIIAN  FLORA.  GUN  NEB  A         381 

It  is  usually  on  the  face  or  the  foot  of  a  precipice  ("pali"  in  Hawaiian)  and 
is  often  difficult  of  access.  It  forms  a  definite  vegetational  belt  between  the 
Middle  Forest  and  the  Upper  Forest,  at  anywhere  from  2,500  to  5,000  feet 
altitude.  Because  of  the  distinctive,  giant  leaves,  this  belt  of  "apeape,"  though 
narrow,  is  often  so  conspicuous  that  it  can  be  seen  from  a  distance  of  two  or 
three  miles. 

The  "apeape"  was  discovered  in  1819  by  Charles  Gaudichaud,  who  published 
it  in  1830  as  Gunnera  petalo'idea.  Subsequent  collections  have  revealed  that 
Gunnera  occurs  on  Kauai,  Oahu,  Molokai,  Maui,  and  Hawaii.  Hillebrand  in- 
terpreted these  as  different  locality  records  for  the  single  species.  This  has 
been  the  generally  accepted  view,  that  the  same  species  occurred  on  all  of 
the  islands. 

The  taxonomic  history  since  the  original  publication  by  Gaudichaud  has 
been  brief.  Alphonse  De  CandoUe  described  a  specimen  collected  on  Kaui 
[=  Kauai]  by  the  United  States  Exploring  Expedition  as  G.  petaloidea  Gaud. 
13  Kauensis  A.DC.  (De  Candolle,  Prodr.  Reg.  Veg.  16  (2)  :  597, 1868) .  His  four 
word  description  included  only  two  characters,  almost  glabrous,  the  bracts 
shorter.  We  now  find  no  differences  in  the  Kauai  plants  in  pubescence,  and 
the  bracts  appear  to  be  alike.  This  plant  was  treated  later  as  an  unnamed 
variety,  G.  petaloidea  Gaud.  /?  (Hillebrand,  Fl.  Haw.  Ids.  124,  1888).  In  1930 
it  was  again  distinguished  by  two  botanists  in  almost  simultaneous  publi- 
cations. Gunnera  kauaie^isis  Rock  in  Caum  (Occ.  Pap.  B.  P.  Bishop  Mus. 
9(5)  :  3^,  pi.  I,  1930,  Sept.).  Rock  described  it  from  a  new  type  specimen. 
Rock,  5,053  from  Mt.  Waialeale.  The  second  author,  Krajina,  redescribed  it 
as  G.  Dominii  Krajina  (Acta  Bot.  Bohemica  9 :  50,  ill.  p.  51,  1930,  without 
precise  date),  citing  as  synonyms  the  varieties  of  De  Candolle  and  of  Hille- 
brand, but  listing  only  his  own  collection  from  "montis  Waialeale."  It  is 
apparent  from  the  characters  given  below  that  this  Kauai  plant  with  peltate 
blades  and  red  to  purplish  drupes  is  a  distinct  species.  Several  attempts  to 
learn  from  Prague  the  exact  month  and  day  of  publication  of  Krajina's  paper 
have  failed.  A  new  search  is  being  made  for  the  date.  In  the  meantime,  the 
name  given  by  Rock  is  accepted  as  it  is  dated  to  the  month. 

Krajina  also  described  and  illustrated  from  his  own  collections  G.  peta- 
loidea. He  published  as  two  new  varieties:  var.  kaalensis  (p.  50)  from  his 
collection  on  Puu  Kaala,  Oahu ;  and  var.  mauiensis  from  his  collection  by 
Waikamoi  Stream,  east  Maui.  Krajina  apparently  did  not  examine  Gaudi- 
chaud's  type  specimen,  nor  did  he  indicate  whether  or  not  either  of  his  two 
varieties  was  to  be  considered  var.  typica  though  with  a  different  name.  Dupli- 
cates of  Krajina's  collections  have  not  yet  reached  the  Bishop  Museum,  nor 
has  the  writer  seen  them  in  other  herbaria.  Still,  there  is  no  difficulty  in  match- 
ing them.  Dr.  V.  Krajina  was  a  fellow  at  the  Bishop  Museum  in  1929-30, 
a  keen,  energetic  botanist  who  soon  became  my  good  friend  and  I  took  him  on 
many  field  trips.  His  t^'pe  of  var.  kaalensis  was  collected  on  my  class  trip  to 
Puu  Kaala  at  the  well-known  locality  above  the  fire-brick  trail  from  Schofield 


382  CALIFOBNIA  ACADEMY  OF  SCIENCES  [Proc.  4™  Ser. 

Barracks.  My  collection,  St.  John  10,072,  was  made  at  the  same  spot  and  time 
as  Krajina's  and  there  are  numerous  other  collections  from  this  locality.  His 
type  of  var.  mauiensis  was  collected  at  another  often  visited  spot  where  the 
Olinda  Pipe  Line  Trail  crosses  Waikamoi  gulch.  There  are  several  other  col- 
lections from  the  same  spot. 

That  recounts  the  known  taxonomic  history  of  this  genus  in  Hawaii.  The 
plant  is  known  to  all  collectors  and  there  are  several  standard,  fairly  easily 
accessible  localities.  The  writer  has  collected  it  on  Kauai,  Oahu,  Maui,  and 
Haw^aii,  but  any  new  localitj'  is  always  noteworthy.  On  Oahu  it  has  been  re- 
peatedly collected  on  the  wet,  windward  side  of  Mt.  Kaala  in  the  Waianae 
Mountains.  There  are  two  or  three  collections  from  the  drier,  leeward  side  of 
the  same  mountain.  Also,  Miss  Marie  Neal  of  the  Bishop  Museum  tells  the 
writer  that  she  has  observed  Giomera  in  the  same  mountain  range  on  the  adja- 
cent peak,  Puu  Kalena,  on  its  northeast  or  windward  side. 

The  preferred  habitat  is  steep,  water-soaked  slopes  on  the  windward  side 
of  a  mountain  range.  It  has  long  puzzled  the  writer  that  the  "apeape"  was 
abundant  in  the  Waianae  Mts.  and  not  in  the  Koolau  Range,  the  more  easterly 
mountain  chain  on  Oahu.  The  Waianae  Mts.  are  rather  arid,  lying  in  the  lee 
of  the  Koolau  Range  which  gets  the  impact  of  the  moist  northeast  trade  winds 
and  receives  most  of  their  moisture,  while  but  little  of  it  reaches  the  Waianae 
Mts.  Only  the  highest  peak,  Mt.  Kaala  (called  Puu  Kaala  on  the  earlier,  1917, 
official  topographic  map)  gets  a  considerable  rainfall.  It  is  just  far  enough  to 
the  north  to  receive  the  impact  of  the  northeast  trade  winds  which  sweep  over 
or  past  the  descending,  low^  north  end  of  the  Koolau  Range.  This  is  only  about 
1,000  feet  high  at  this  point,  not  high  enough  to  cause  the  precipitation  of  all 
the  moisture.  Hence,  Mt.  Kaala  with  its  large  flat  summit  at  4,025  feet  altitude 
and  its  upper  ridges  get  enough  rainfall  to  support  a  luxuriant  rain  forest. 
The  summit  has  a  swampy  forest  except  for  some  boggy  openings  near  its 
center.  The  average  annual  rainfall  is  now  known  to  be  91  inches  (Feldwisch, 
W.  E.,  First  Progress  Rept.,  Hawaii  Territorial  Planning  Board  121,  1939). 

The  Koolau  Range,  Oahu,  running  from  southeast  to  northwest,  forms  the 
backbone  of  the  eastern  side  of  the  island.  It  rises  abruptly  close  to  the  shore, 
and  the  section  above  1,500  feet  in  height  extends  for  about  25  miles, 
making  a  barrier  almost  exactly  at  right  angles  to  the  prevailing  northeast 
trades.  There  are  numerous  sharp  peaks  that  rise  from  the  range,  the  highest 
being  Puu  Konahuanui  with  twin  peaks  3,105  and  3,150  feet  high.  The  several 
rain  gauging  stations  show  a  high  rainfall,  as  this  mountain  range  catches 
much  of  the  moisture  from  the  clouds.  The  rainfall  is  more  than  100  inches 
annually  for  most  of  this  ridge  and  at  the  wettest  station  is  311  inches.  There 
are  miles  of  wind-swept  precipices  ("pali")  and  steep  slopes  with  much  water 
seepage  from  the  higher  peaks,  just  the  kind  of  habitat  that  usually  supports 
Gunnera.  The  writer  had  botanized  these  brinks  and  declivities  at  many  locali- 
ties along  the  Koolau  Range,  and  had  never  found  the  "apeape."  Yet,  this 
Gunnera  is  a  plant  that  one  cannot  easily  overlook.  With  its  bizarre  leaves 


Vol.  XXV]       ST.  JOHN:  ENDEMISM  IN  HAWAIIAN  FLORA.  GUNNEBA         383 

like  huge  iTmbrellas,  it  is  always  the  biggest  and  most  conspicuous  plant  in  its 
area.  There  were  no  specimens  from  the  Koolau  Range  in  the  Bishop  Museum, 
yet  the  plant  has  been  reported  from  there.  Prof.  Vaughan  MacCaughey  listed 
it  as  rare  in  the  rain  forests  of  Manoa  Valley  (Am.  Journ.  Bot.  4 :  600,  1917). 
In  the  same  year  in  a  short  article  about  Gunnera  petaloidea  (Am.  Journ. 
Bot.  4 :  38,  1917,  the  reprints  incorrectly  printed  vol.  1)  he  stated  that  among 
its  typical  habitats  were  "Ka-ala  and  Kona-hua-nui  summit  ridges  on  Oahu 
(2,500-4,000  ft.) ."  The  4,000  foot  record  is  certainly  that  for  the  often  visited 
station  of  Mt.  Kaala  in  the  Waianae  Mts.  Hence,  we  infer  that  his  station  on 
the  3,150  foot  peak  Puu  Konahuanui  was  at  about  2,500  feet  altitude.  Mac- 
Caughey did  exploring  and  original  research  in  Manoa  Valley  which  heads 
on  the  slope  of  Puu  Konahuanui.  The  plant  is  unmistakable,  so  there  would 
seem  no  reason  to  question  this  record,  yet  corroborating  specimens  were  much 
to  be  desired. 

Some  years  ago  the  writer  had  an  enterprising  young  man  as  a  student, 
John  R.  Coleman.  He  was  enlisted  in  the  United  States  Marine  Corps  and  was 
stationed  in  Honolulu  with  a  company  doing  guard  duty  at  the  docks.  At  his 
request  he  was  given  continuous  and  regular  night  guard  duty.  This  left  his 
day  time  free  and  enabled  him  to  attend  the  University  of  Hawaii.  It  was  not 
obvious  when  he  slept,  but  he  did  not  sleep  in  class.  He  completed  the  course 
on  elementary  botany  and  on  week  ends  he  often  joined  advanced  classes  on 
collecting  trips  to  the  mountains.  One  Sunday  he  induced  two  other  marines 
to  join  him,  and  from  Nuuanu  Pali  they  started  to  climb  the  knife-edged  ridge 
that  culminates  in  Puu  Konahuanui.  As  he  led  the  way  and  scrambled  up  one 
precipitous  slope,  the  bushes,  soil,  and  all  caved  away.  He  managed  to  make 
headway  over  the  landslide  and  surmounted  the  ledge.  All  the  footholds  were 
destroyed,  so  his  companions  could  not  follow,  and  turned  back.  Coleman  went 
on,  climbed  to  the  summit,  and  descended  by  a  more  southerly  route.  The  next 
day  he  brought  me  a  10  cm.  tip  from  a  fruiting  inflorescence  with  half  ripe 
drupes,  of  a  specimen  that  seemed  to  be  Gunnera.  He  reported  collecting  it 
near  the  summit  on  Oct.  19,  1930.  This  specimen  gave  partial  proof  of  the 
occurrence  in  the  Koolau  Range,  but  it  was  incomplete.  At  last,  in  June  1942 
the  proof  was  furnished  by  two  students,  L.  Eubank  and  A.  D.  Conger,  who,  on 
a  wet  day  amid  swirling  clouds  and  rain,  ascended  Puu  Konahuanui.  They  ob- 
served and  collected  abundant  and  complete  material.  They  reported  the  three 
small  colonies  to  be  on  the  Manoa  or  leeward  slope.  A  second  visit  on  Sept. 
13, 1942  by  A.  D.  Conger,  H.  St.  John,  and  R.  P.  St.  John  w^as  also  in  the  clouds 
and  intermittent  rain,  but  thrice  the  clouds  lifted  to  reveal  the  slopes,  valleys 
and  the  shore  line.  All  three  patches  proved  to  be  on  the  windward  or  Kailua 
side.  Any  mountaineer  will  understand  how  in  following-  forking,  intricate 
ridges  in  dense  clouds,  one  can  lose  track  of  the  directions.  One  locality  was 
a  single  plant  some  250  feet  below  the  lower  or  south  peak  which  is  mapped 
and  triangulated  as  Puu  Konahuanui.  The  two  others  were  on  the  east  face  of 
a  northeast  ridge  of  the  north  or  higher  peak.  There  were  numerous  plants  in 


384  CALIFOENIA  ACADEMY  OF  SCIENCES  [Proc.  4th  Ser. 

a  steep,  brushy,  hanging  valley  at  2,950  feet  altitude.  Though  both  Miss 
Eubank  and  Mr.  Conger  previously  slid  down  the  cloud-blanketed  slope  to 
the  plants,  then  scrambled  up  again,  on  the  second  trip  no  one  wanted  to  try  | 
it  again.  It  was  very  dangerous,  and  when  clear  of  clouds  it  was  obvious  that 
an  uncontrolled  slip  on  the  steep,  slippery  clay  would  give  one  a  flying  start 
over  the  1,000  foot  precipice.  Later  a  fourth  colony  was  located.  On  May  9, 
1943  the  writer  conducted  a  student  party  of  nine  from  Tantalus  to  the  sum- 
mit of  Puu  Konahuanui.  Clouds  blanketed  the  precipices  on  the  east  slope 
hiding  the  single  plant  on  the  South  Peak.  From  the  North  Peak  a  descent  was 
made  down  the  secondary  ridge  towards  Kailua.  The  middle  and  inaccessible 
colony  of  Gunnera  was  again  seen  far  below  in  the  cloud.  Neither  the  writer 
nor  his  son  Robert  could  locate  the  third  colony,  though  this  time  a  rope  was 
at  hand  for  a  descent  to  it.  The  clouds  hid  it  completely.  Climbing  up  again, 
the  descent  was  made  along  the  main  divide  to  the  road  at  Nuuanu  Pali.  On 
this  divide  a  new  colony  was  found,  also  on  the  east  face,  in  a  moist,  precipi- 
tous, hanging  valley  at  2,300  feet  altitude.  By  the  use  of  a  rope  the  colony  was 
visited  and  specimens  collected.  There  were  more  than  12  plants,  up  to  4 
meters  tall,  some  exposed,  others  half  hidden  by  the  thicket  of  Pipturus  alhidus 
and  Cyrtandra  paludosa.  The  specimens  collected  have  been  deposited  in  the 
Bishop  Museum.  These  plants  from  the  Koolau  Range  were  critically  com- 
pared with  those  from  the  east  side  of  Mt.  Kaala  in  the  Waianae  Mts.  and 
were  found  to  be  identical  in  structure  and  quite  inseparable,  though  the 
blades  are  smaller,  the  length  along  the  midrib  not  exceeding  4  dm. 

While  investigating  this  material  the  writer  took  occasion  to  assemble  all 
available  specimens  and  to  evaluate  the  species  and  varieties  described  from 
the  other  Hawaiian  Islands  by  Rock  and  by  Krajina.  Its  massive  fleshy  stems, 
giant  fruit  clusters,  and  rounded  leaves,  1  to  2  meters  in  diameter,  cannot  well 
be  compressed  upon  a  standard  herbarium  sheet,  so  that  the  collector  is  forced  f 
to  take  sample  fragments  which  are  inevitably  incomplete,  and  the  leaves 
selected  are  often  the  smaller  ones.  Hence,  all  taxonomists  revising  the  group 
and  writing  descriptions  from  the  available  herbarium  material  present  meas- 
urements which  are  true  as  far  as  they  go,  but  minimize  the  size  of  the  giant 
stems,  leaves,  and  inflorescences.  Several  of  the  writer's  whole  numbers  in- 
cluding many  unmounted  duplicates  are  available,  but  still  there  are  gaps. 
From  east  Maui  there  are  no  mature  stipules,  and  from  west  Maui  there  are 
no  petals,  and  mature  drupes  and  stones  are  needed  from  Molokai  and  from 
Makaha  Valley,  Oahu.  Still  the  collections  are  numerous  and  have  proven 
sufficient.  At  a  glance  they  all  look  alike,  but  on  close  study  differences  are^ 
noted  in  the  floral  and  fruiting  morphology.  It  is  curious  that  most  of  the 
characters  are  minute,  a  difference  in  shape  to  be  seen  with  a  lens  or  in  size 
measured  in  fractions  of  a  millimeter.  Seedlings  are  often  collected.  Since  they  i 
produce  smaller  stipules  and  blades  that  are  much  smaller  and  more  renif  orm,^ 
they  have  not  been  included  in  our  descriptions.  The  only  large  parts  furnish- 
ing characters  are  the  stipule.s  and  the  leaves  as  to  lobing  and  toothing  and! 


Vol.  XXV]       ST.  JOHN:  ENDEMISM  IN  HAWAIIAN  FLORA.  GUNNEBA         385 

pubescence.  When  contrasted  throughout,  several  correlated  characters  mostly 
of  floral  and  fruiting  anatomy  are  found.  When  separated  on  these  characters, 
the  segregates  are  found  to  agree  in  leaf  and  stipule  characters  and  to  have 
good  geographic  correlations.  Each  island  has  one  local  kind,  except  Maui  and 
Oahu  which  have  two  each.  When  evaluated,  the  number  and  sort  of  char- 
acters would  indicate  that  these  segregates  are  properly  considered  species. 
The  original  species  was  Gunnera  petalo'idea  Gaudichaud  (Voy.  "Uranie," 
Bot.  512,  1830).  The  plant  was  not  illustrated  and  the  16  word  description 
includes  only  the  major  features,  but  none  of  the  ones  now  found  serviceable 
in  distinguishing  the  local  species.  Neither  Rock  nor  Krajina  gave  any  dis- 
cussion as  to  the  nature  of  G.  petalo'idea  Gaud,  or  its  type  locality.  That  pub- 
lished by  Gaudichaud  was  "In  insulis  Sandwicensibus  (Alt.  450-500  hex.)." 
No  island  is  specified,  and  the  statement  of  altitude  in  fathoms  is  unusual,  but 
was  perhaps  natural  after  months  at  sea  on  a  sailing  vessel !  Search  has  been 
made  for  any  detailed  itinerary  of  Gaudichaud.  He  published  none,  and  the 
narrative  by  the  commander,  Capt.  Louis  de  Freycinet,  and  those  by  other 
members  of  the  expeditions  give  much  on  the  physique  and  social  habits  of  the 
native  inhabitants,  but  nothing  on  the  route  of  the  collecting  botanist.  On  the 
"Uranie"  voyage  they  spent  20  days  in  the  Hawaiian  Islands  and  Gaudichaud 
was  on  shore  7  days.  The  writer  has  long  marveled  at  the  large  number  of 
the  endemic  species  and  genera  that  he  collected  and  described  on  this  first 
voyage.  Starting  from  the  harbor,  with  no  good  roads,  no  cut  mountain  trails, 
to  get  up  into  the  dense,  native  forests  and  through  them  to  the  cloud  zone 
was  a  herculean  task.  Then,  over  half  of  his  collections  were  lost  when  the 
homeward  bound  ship  was  w^recked  on  the  Falkland  Islands.  All  were  soaked 
in  sea  water,  but  he  rescued  many  by  drying  them  before  a  bonfire  on  the 
beach.  These  experiences  are  mentioned  in  a  few  brief  words,  but  what  stark 
tragedy  they  describe !  What  a  grand  naturalist  he  must  have  been,  yet  no 
portrait  of  him  is  known !  He  did  not  specify  where  his  shore  trips  were  made, 
but  the  ship  made  landings  on  Aug.  8,  1819  at  Kayakoua  [Kailua,  Hawaii] ; 
Aug.  12  at  Bay  of  Kohaihai'  [Kawaihae,  Hawaii]  ;  Aug.  17-25  at  Raheina 
[Lahaina,  Maui]  ;  Aug.  26-30  at  d'Onorourou  [Honolulu,  Oahu] .  So  the  seven 
days  on  shore  were  divided  among  these  localities,  but  just  how  is  not  known. 
In  one  day  from  the^arid  regions  near  Kailua  and  Kawaihae,  it  would  not  have 
been  possible  to  ascend  to  the  habitat  of  Gunnera  on  the  far  side  of  the  Kohala 
Mts.  Hence,  the  island  of  Hawaii  can  be  eliminated.  Back  of  Lahaina,  Maui, 
the  mountains  rise  abruptly  from  near  the  shore,  and  on  the  steep  upper  slopes 
of  the  valleys  Gunnera  is  abundant.  From  Honolulu  it  would  have  been  possi- 
ble to  find  Gunnera  in  either  mountain  range.  However,  the  east  cliffs  of  Mt. 
Kaala  were  distant  and  difiicult  of  access.  It  is  unlikely  that  he  could  have 
gotten  there.  Puu  Konahuanui  is  only  6  miles  in  an  air  line  from  the  harbor 
of  Honolulu.  It  is  known  that  he  climbed  to  the  cloud  zone  at  the  crest  of  the 
Koolau  Range,  because  he  collected  Lobelia  Gaudichaudii  which  is  restricted 
to  that  area.  Hence,  it  can  be  deduced  that  Gaudichaud  probably  collected  the 


386  CALIFORNIA  ACADEMY  OF  SCIENCES  [Proc.  4th  Ser. 

Gunnera  in  the  mountains  of  west  Maui  or  in  the  Koolau  Range  near  Hono- 
lulu, Oahu.  In  1935  the  writer  photographed  in  Paris  Gaudichaud's  type  speci- 
men of  0.  petalo'idea,  reproduced  here  as  PI.  38.  The  data  on  the  sheet  read 
"90.  Uranie,  Ins.  Sandwich.  C.  Gaudiehaud."  Stipules  are  lacking  and  the 
pieces  of  the  inflorescence  do  not  permit  one  to  detect  the  minute  floral  differ- 
ences from  the  photograph.  However,  the  margin  of  the  leaf  is  well  shown. 
The  distinct,  rounded  lobes  and  the  low,  rounded  teeth  or  crenations  of  the 
leaf  margin  match  exactly  those  of  the  species  of  west  Maui,  so  that  it  is  to 
be  taken  as  the  type  locality. 

The  Hawaiian  species  all  belong  to  the  subgenus  Panke  (Molina)  Schindler 
(Engler,  Pflanzenreich  IV,  fam.  225:117,  1905).  This  subgenus  has  a  re- 
markable distribution,  occurring  also  in  the  Cordilleras  from  Mexico  to  Chili, 
and  on  Juan  Fernandez  (Skottsberg,  C,  Proc.  6th  Pacif.  Sci.  Congr.  Berkeley 
4:  696,  map  14,  1940). 

In  the  ensuing  systematic  treatment  are  given  key,  diagnoses,  and  descrip- 
tions of  the  species  as  at  present  understood. 

All  the  specimens  examined  and  cited  and  the  new  types  are  in  the  B.  P. 
Bishop  Museum,  Honolulu,  except  the  Gaudiehaud  type  of  G.  petalo'idea 
which  is  in  the  Museum  d'Histoire  Naturelle,  Paris. 

Key  to  the  Hawaiian  Species  op  Gunnera 
Blades  peltate ;  drupes  red  to  purplish 7.  G.  Tcauaiensis  Eock 

Blades  basifixed;  drupes  yellow,  orange  (or  reddish  in  G.  mauiensis) 
Stipules  scurfy  on  back  or  finally  glabrate, 

Petals  entire,  obtuse,  glabrous ;  sepals  0.6—0.8  mm  long,  rounded,  lacerate ;  anthers 
1.2-1.3  mm.  long ;  peduncle  bractless  except  at  base ;  blade  margin  lack- 
ing mucros 5.G.  Tcaalaensis  (Krajina)  St.  John. 

Petals  lacerate-fimbriate,  often  acuminate;  sepals  1-1.2  mm.  long,  broadly  deltoid, 
deeply  lacerate;  anthers  1.6-2.2  mm.  long;  peduncle  bracted;  (stipules 
apparently  scurfy)  ;  teeth  of  blade  margin  with  revolute  mucros 

6.  G.  maTcahaensis  St.  John. 
Stipules  not  scurfy  on  the  back, 
Stipules  glabrous, 

Sepals  1.5-2  mm.  long,  ovate,  deeply  lacerate,  the  lacerations  fimbriate;  petals 
entire 1.  G.  petaloidea  Gaud. 

Sepals  0.6-0.8  mm.  long,  broadly  deltoid  to  rounded,  lacerate ;  petals  minutely 
lacerate 2.  G.  moloTcaiensis  St.  John. 

Stipules  pubescent. 

Petals  1.5  mm.  long,  entire  or  somewhat  erose;  sepals  0.4—0.6  mm.  long,  semi- 
oval,  the  obtuse  tip  minutely  ciliate;  anthers  1.9-2  mm.  long,  elliptic 
oblong 3.  G.  mauiensis  (Krajina)  St.  John. 

Petals  0.8-1  mm.  long,  sparsely  lacerate;  sepals  1-1.2  mm  long,  oblong  lan- 
ceolate, the  tip  lacerate;  anthers  1.5-1.7  mm.  long,  oval 

4.  G.  Eastwoodae  St.  John. 


Vol.  XXV]       ST.  JOHN:  ENDEMISM  IN  HAWAIIAN  FLORA.  GUNNERA         387 

1.  Gunnera  petaloidea  Gaudichaud 

(Plate37,  fig.  d;  pis.  38,  39) 
Gunnera  petaloidea  Gaudichaud,  Voy.  "Uranie,"  Bot.  512,  1826  (=:  1830). 

Giant  herb,  1-6  m.  tall;  stem  2-5  m.  long,  15-22  cm.  in  diameter,  fleshy; 
often  long  decumbent,  the  surface  with  prominent  leaf  scars  and  many  oval 
pits  from  which  prop  roots  develop  on  older,  mostly  decumbent  parts  of  the 
stem ;  roots  reddish,  slimy  at  tip ;  stipides  5-9  cm.  long,  oblanceolate,  pale, 
strongly  nerved,  glabrous;  petioles  2-4  dm.  long,  hirsute,  swollen  at  the  base ; 
blades  27-53  cm.  long,  cordate-reniform,  shallowly  palmately  9-  or  11-lobed, 
firm  chartaceous,  dark  green,  glabrous  or  nearly  so,  impressed  reticulate 
nerved,  the  veins  sparsely  hirsutulous,  below  the  surface  paler  green,  the  veins 
even  to  the  tertiary  ones  densely  hirsute,  the  lobes  1.5-6  cm.  long,  broadly 
deltoid,  the  midrib  and  prominent  lateral  veins  each  ending  in  a  prominent 
deltoid  dentation,  the  margin  crenate  or  low  rounded  dentate,  the  sinus 
broadly  V-shaped ;  inflorescence  probably  axillary ;  bracts  at  base  of  inflor- 
escence 3-7  cm.  long,  lanceolate  to  oblanceolate,  like  the  stipules;  peduncle 
5-16  cm.  long,  hinsute,  bractless;  inflorescence  3-5.5  dm.  long,  12-25  cm.  in 
diameter,  lanceoloid,  hirsute  towards  the  base,  more  or  less  glabrous  towards 
the  tip,  the  principal  lower  bracts  20-28  mm.  long,  broadly  linear,  acute,  the 
margin  membranous,  the  lateral  branches  ascending  or  diverging,  spiciform ; 
flowers  perfect,  sessile;  ovary  barrel-shaped;  calyx  lobes  1.5-2  mm.  long, 
ovate,  deeply  lacerate,  then  the  lacerations  fimbriate;  petals  1.5  mm.  long, 
obovate,  cucullate,  entire,  glabrous ;  the  2  anthers  opposite  the  petals,  anthers 
1.6-1.9  mm.  long,  narrowly  elliptic-oblong;  the  2  stigmas  2  mm.  long,  terete, 
densely  puberulent  throughout;  drupes  2.5-3  mm.  in  diameter,  subglobose, 
greenish-yellow;  stone  1.5-1.8  mm.  long,  0.9-1.3  mm.  wide,  0.9-1  mm.  thick, 
oval  in  outline,  4-angled,  4-sided,  two  of  the  angles  prominent,  the  third  less 
so,  and  the  fourth  still  weaker  and  unequal,  the  stone  laterally  compressed, 
the  surface  dull,  pale  yellowish  browaiish. 

Type:  "In  insulis  Sandwicensibus  (alt.  450-500  hex.)."  The  type  specimen 
in  Paris  is  labeled  "90,  lies  SandAvich,  C.  Gaudichaud."  This  is  illustrated  in 
our  Plate  38. 

Collections  studied,  all  from  western  Maui:  Eeka  [of  Hillebrand  =  Puu  Kukui],  ex  herb. 
Lydgate,  W.  Hillebrand  and  J.  M.  Lydgate;  gulch  sides  by  Mt.  Kukui,  Sept.  24,  1916, 
G.  C.  Munro  426  and  594;  Hanakaoo,  shingles  the  valley  sides  in  places,  May  1910,  C.  N. 
Forbes  69.M;  Olowalu  Valley,  central  ridge,  far  up  on  the  peak,  large  patches,  May  21, 
1920,  C.  N.  Forbes  2,433.M;  without  locality,  J.  F.  RocTc  and  fi'[osme)-]  8215. 

A  discussion  of  the  type  locality  is  given  in  our  introduction.  Gaudichaud 
could  have  discovered  Gunnera  either  on  Oahu  or  on  western  Maui.  Neither 
his  publication  nor  the  type  specimen  in  Paris  gives  any  hint  as  to  the  island 
concerned.  Luckily  there  are  some  differences  in  leaf  shape,  and  by  careful 
examination  of  the  lobing  and  dentation  of  the  leaf  margin  it  has  been  possible 
to  match  the  type  with  recent  collections  from  near  Lahaina,  Maui.  Hence, 


388  CALIFORNIA  ACADEMY  OF  SCIENCES  [Proc.  4th  Ser. 

it  is  deduced  that  Gaudichaud  collected  the  type  in  the  mountains  back  of 
Lahaina,  Maui. 

Most  previous  authors  have  combined  under  G.  petalo'idea  all  of  the  Ha- 
waiian specimens.  This  was  due,  doubtless  in  large  part,  to  the  fact  that  few 
if  any  of  them  had  sufficient  material  to  allow  the  recognition  of  any  diag- 
nostic characters  between  the  specimens  from  the  different  islands.  Conse- 
quently, their  descriptions  are  mostly  composite  ones,  so  that  one  cannot  now 
rely  upon  them.  This  also  applies  to  the  monographic  treatment  by  A.  K. 
Schindler  (Engler's  Pflanzenreich  IV,  fam.  225:117,  fig.  34,  1905).  The  de- 
tails that  he  shows  in  his  figure  34  of  the  blade,  petal,  and  anther  agree  with 
the  characters  of  the  common  Oahu  plant.  The  entire,  broadly  triangular 
calyx  lobes  are  different  from  any  known  Hawaiian  plant,  and  it  seems  that 
they  are  incorrectly  delineated.  He  cited  one  by  Gaudichaud  and  a  total  of 
six  collections,  originating  from  three  islands,  so  it  is  quite  possible  that  his 
drawing  was  a  composite  one  made  from  several  of  these  collections.  Hence, 
this  Schindler  drawing  is  considered  to  be  in  large  part  one  of  G.  kaalaensis, 
but  not  fully  accurate. 

Gaudichaud  in  1830,  when  he  described  G.  petalo'idea,  in  a  footnote  depre- 
cated his  choice  of  a  specific  name  which  alluded  to  the  presence  of  petals.  He 
had  learned  meanwhile  that  some  other  species  of  Gunnera  had  petals,  but 
nevertheless  he  retained  the  name  of  G.  petalo'idea  since  he  had  mentioned  it 
already  in  his  introduction  (p.  98)  printed  in  1827. 

Most  recent  botanists  have  misspelled  the  named  as  G.  petaloidea,  but 
Gaudichaud  clearly  published  it  as  G.  petalo'idea,  spelling  it  thus  with  a 
diaeresis  over  the  i  to  indicate  a  separation  of  the  syllables,  and  spelling  it 
consistently  on  two  remote  pages.  This  was  not  an  error,  but  rather  his  delib- 
erate choice,  so  we  must  follow  his  spelling  despite  its  strange  appearance  in 
a  latinized  name. 


4 


4 


I 


2.  Gunnera  molokaiensis  St.  John,  sp.  no  v. 

(Plate  38,  fig.  e) 

Herba  gigantea,  laminis  43  cm.  longis  cordato-reniformibus  9-lobatis  chartaceis  infra 
pallidioribus  nervis  reticulatis  hirsutulisque,  lobis  5-8  cm.  longis  sinuato-rotundatis  denti- 
culatis,  inflorescentiis  15-40  cm.  longis  laneeoloideis,  ramulis  spicatis,  sepalis  0.6-0.8  mm. 
longis  late  deltoideis  vel  rotundatis  laceratis,  petalis  1.5  mm.  longis  cueullatis  spatulatis 
minute  laceratis  vel  subintegris,  antheris  1.5-1.9  mm.  longis  ovalibus,  fructibus  ignotis. 

Giant  herb  ;  stipules  2  cm.  long,  lanceolate,  glabrous  ( ?)  ;  petiole  (one  seen) 
47  cm.  long,  with  scattered  dark  pustules  especially  at  base,  densely  hirsute  at 
base,  sparsely  so  to  glabrate  elsewhere;  blade  43  cm.  long,  apparently  cordate- 
reniform,  shallowly  palmately  9-lobed,  firm,  chartaceous,  above  dark  green,  \ 
the  surface  and  veins  remotely  hirsutulous  to  glabrate,  below  pale  green,  the 
prominent  reticulate  veins  all  scabrous  hirsutulous,  the  lobes  5-8  cm.  long 
sinuous  rounded,  the  outline  ovate,  the  sinus  U-shaped,  the  margin  irregularly 
rounded  denticulate ;  inflorescence  probably  axillary ;  peduncle  7  cm.  long 
(when  young) ,  bractless,  tomentose  towards  the  base;  inflorescence  15—40  cm. 


Vol.  XXV]       -ST.  JOHN:  ENDEMISM  IN  HAWAIIAN  FLORA.  GUNNEBA         389 

long,  4-15  cm.  in  diameter,  cylindric  to  lanceoloid,  tomentose  on  the  axis  and 
towards  the  base,  glabrate  above,  the  principal  lower  bracts  8-12  mm.  long, 
linear-spatulate,  the  margin  membranous,  the  lateral  branches  ascending, 
spicif orm ;  flowers  perfect,  sessile ;  ovary  barrel-shaped,  glabrous ;  calyx  lobes 
0.6-0.8  mm.  long,  broadly  deltoid  to  rounded,  lacerate ;  petals  1.5  mm,  long, 
cucullate,  spatulate,  minutely  lacerate  or  subentire ;  the  2  anthers  opposite  the 
petals,  anthers  1.5-1.9  mm.  long,  oval;  stigmas  immature;  drupes  immature. 

Tijpe:  Molokai,  Pelekunu  Pali,  July  12,  1912,  C.  N.  Forhes  305. Mo.  B.  P. 
Bishop  Museum,  Honolulu. 

Forbes  was  the  only  one  to  bring  back  Gunnera  from  the  remote  precipi- 
tous, windw^ard  brink  of  the  Molokai  mountains.  His  locality  was  on  the  old 
trail  which  started  from  Kamalo,  ascended  the  broad  ridge  to  the  cabin  near 
Puu  Kolekole,  then  northeastward  across  the  swampy  forest  and  bogs  to  the 
divide  near  Kaunohua  at  about  4,535  feet  altitude,  then  descended  in  switch- 
backs down  the  steep  head  wall  on  the  north  side.  This  deeply  eroded  Pele- 
kunu Valley  is  a  spectacular  bit  of  mountain  scenery.  The  Gunnera  was  prob- 
ably found  just  below  the  windward  or  northern  crest  of  the  divide. 

The  writer  has  a  record  in  his  field  notebook  of  observing  Gunnera  in 
Waikolu  Valley,  the  next  one  to  the  west.  More  exactly  this  was  at  Hanalilolilo, 
near  the  head  of  the  deeply  eroded  part  of  the  valley.  He  did  not  get  close 
enough  to  collect  the  specimens. 

G.  molohaiensis  is  most  closely  related  to  the  species  of  western  Maui,  G. 
petalo'idea  Gaudichaud,  which  has  the  blades  shallowly  lobed,  with  lobes  1.5-6 
cm.  long,  the  margin  crenate  or  low  rounded  dentate ;  the  main  veins  beneath 
densely  hirsute ;  the  sepals  1.5-2  mm.  long,  ovate,  deeply  lacerate,  the  lacera- 
tions fimbriate;  and  the  petals  entire.  G.  molohaiensis  has  the  blades  with 
marked,  well  rounded  lobes,  the  lobes  5-8  cm.  long,  the  margin  irregularly 
rounded  denticulate;  the  main  veins  beneath  sparsely  tomentose  to  early 
glabrate;  the  sepals  0.6-0.8  mm.  long,  broadly  deltoid  to  rounded,  lacerate; 
and  the  petals  minutely  lacerate. 

3.  Gunnera  mauiensis  (Krajina)  St.  John,  comb.  nov. 

(Plate37,fig.  g;pl.  40.) 

Gunnera  petaloidea  Gaudichaud  var.  mauiensis  Krajina,  Acta  Botanica  Bohemica  9 :  50, 

figs.  B2,  F2,  Ij,  1  pi.  1930. 
Illustrations :  by  Bonine,  in  Eock,  J.  P.,  Monograph  of  the  Hawaiian  Lobelioideae,  Mem. 
Bishop  Mus.  7(2) :  pi.  34,  1919;  by  Krajina,  see  above;  by  Degener,  O.,  PI.  Hawaii  Natl. 
Park,  pi.  66, 1930. 

Giant  herb  about  2-3  m.  tall ;  stem  about  1-2  m.  tall,  the  tip  assurgent,  the 
stem  up  to  6  m.  in  length  and  15-20  cm.  in  diameter;  bark  near  apex  hirsute, 
with  numerous  glabrous,  oval  pits,  where  prop  roots  may  later  develop ; 
terminal  bud  completely  covered  by  the  numerous  imbricate  stipules ;  stipules 
of  seedlings  1.5-2  cm.  long,  ovate,  membranous,  strongly  nerved,  slender 
hirsute  on  the  back,  those  of  mature  leaves  7.5-10  cm.  long,  elliptic-lanceolate 
with  a  broad  truncate  base,  pale  greenish,  strongly  nerved,  sparsely  puber- 


390  CALIFORNIA  ACADEMY  OF  SCIENCES  [Proc.  4th  Ser. 

ulous  base  and  tip ;  petioles  3-10  dm.  long,  pustulate,  densely  hirsute  at  base, 
sparsely  so  elsewhere;  blades  30-56  cm.  long,  cordate-reniform,  shallowly  J 
palmately  7-11-lobed,  firm  chartaceous,  above  dark  green,  remotely  hirsutu- 
lous,  soon  glabrate,  below  pale  green,  the  prominent  veins  hirsute,  the  reticu- 
late veinlets  hirsutulous,  the  lobes  2-A  cm.  long,  low  rounded  sinuate,  the 
midrib  and  one  or  more  principal  lateral  veins  ending  in  low,  deltoid  denta- 
tions, the  sinus  broad  U-shaped,  the  margin  with  low  broad  denticulations ; 
inflorescence  supra-axillarj' ;  bracts  at  base  of  inflorescence  5  cm.  long,  ob- 
lanceolate,  membranous,  glabrous ;  peduncle  15-30  cm.  long,  hirsute,  bractless ; 
inflorescence  4.3-10.5  dm.  long,  9-24  cm.  in  diameter,  lanceoloid,  the  axis 
hirsute,  the  principal  low^er  bracts  linear-spatulate,  acute,  with  membranous[ 
margin,  the  lateral  branches  ascending,  spiciform,  hirsute  at  base,  glabrous] 
towards  the  tip;  flowers  perfect,  sessile ;  ovary  depressed  globose ;  calyx  lobes 
0.4—0.6  mm.  long,  semioval,  the  obtuse  tip  minutely  ciliate;  petals  1.5  mm.  long, 
broadly  cucullate-spatulate,  glabrous,  entire  or  somewhat  erose ;  the  2  anthers 
1.9-2  mm.  long,  elliptic-oblong;  the  2  stigmas  2-3  mm.  long,  densely  puberu- 
lous ;  drupes  2.5-3.5  mm.  in  diameter,  globose,  pale  salmon-red ;  stone  1.6-1.9 
mm.  long,  1.4—1.5  mm.  wide,  1-1.2  mm.  thick,  broadly  obovate  in  outline  with 
three  prominent  angles  of  which  one  is  lower  due  to  a  slight  compression  of 
the  stone,  and  a  fourth  angle  more  or  less  developed  or  wanting,  the  surface 
pale  yellowish  brown,  dull.  The  fresh  flowers  are  rose-magenta. 

Type:  "Maui :  Makawao  District  ad  plurimos  rivulos  fluminis  Waikamoi  in 
declivibus  septentrionali-orientalibus  montis  Haleakala,  ca.  1275  m.  s.  m.,  legi 
15.  IV.  1930."  V.  Krajina.  Herb.  Charles  Univ.,  Prag-ue,  not  seen. 

Collections  studied,  all  from  eastern  Maui:  Olinda,  along  pipeline  from  Idlewild  for 
about  7  miles  in  deep  wet  ravine,  July  16,  1927,  0.  Begener  and  H.  Wiebhe  3,404;  the  same, 
July  30,  1927,  D.  L.  Topping,  herb.  Begener  8,185 ;  and  June  19,  1927,  0.  Begener  and  H. 
Wieilce  2,240,  2,240a,  2,240});  Kula  Pipe  Line,  4,000  ft.  alt.,  bank  of  Waikamoi  Stream,  Feb. 
11,  1930,  K.  St.  John  10,302;  Waikamoi,  rain  forest,  4,250  ft.,  Aug.  14,  1933,  M.  Neal  and 
C.  Hartt;  Haiku  Uka,  Waikamoi  Gulch,  steep  side  walls,  4,400  ft.  alt.,  Aug.  4,  1943,  E.  St. 
John  20,358.  The  foregoing  are  all  from  the  same  station  on  the  pipe  line  trail  east  of  Olinda. 
Ukulele,  Sept.  4.  1919,  C.  N.  Forbes  1,255.M;  Haipuaena  Stream,  sides  of  wet  valley,  el. 
4,400  ft.,  Nov.  23,  1927,  G.  C.  Munro  740 ;  below  Koolau  Gap,  Haleakala,  extremely  rainy, 
wet  region,  June  17,  1927,  0.  Begener  8,184 ;  Nahiku,  Kuhiwa  trail,  Koolau  Forest  Eeserve, 
6,000  ft.  alt.,  wet  woods,  Jan.  2,  1937,  E.  St.  John  and  E.  J.  Catto  17,935;  Kipahulu  Valley, 
northerly  side  gulch  near  head  of  Oheo  Stream,  wet  thicket,  5,800  ft.  alt.,  Aug.  22,  1945, 
E.  St.  John  and  A.  L.  Mitchell  21,104. 

Most  of  the  localities  are  in  one  general  region,  within  a  few  miles  of 
each  other.  They  are  on  the  windward  slope  of  Haleakala  between  4,000  and 
6,000  feet  altitude.  Here  in  the  very  wet  forests  are  numerous  narrow  gulches. 
On  their  steep,  often  dripping  side  walls,  the  Ounnera  is  abundant. 

On  the  drier  southeast  side  of  Haleakala,  Gunnera  was  reported  in  Kipa- 
hulu Valley  by  Mr.  David  T.  Fleming.  In  the  summer  of  1945  while  exploring! 
Haleakala  for  the  National  Park  Service,  it  was  possible  for  the  writer  to  enter 
this  area.  The  upper  section  of  Kipahulu  Valley  is  well  watered  and  densely 
forested;  and  the  steep  head  walls  are  nearly  always  blanketed  with  clouds 


Vol.  XXV]       ST.  JOHN :  ENDEMISM  IN  HAWAIIAN  FLORA.  GUNNEEA         391 

and  very  wet.  The  stream  heads  have  cut  narrow  gulches  on  the  precipitous 
head  walls  and  in  these  the  Gunnera  occurs  from  about  4,500  to  6,000  feet 
altitudes.  It  was  found  possible  via  Wai  Anapanapa  to  approach  one  locality 
and  get  specimens.  These  plants  prove  to  be  conspecific  with  those  of  the  wind- 
ward slope  of  Haleakala. 

4.  Gunnera  Eastwoodae  St.  John,  sp.  nov. 

(Plate  3  7,  fig.  f;  plate  42) 

Herba  gigantea  1.5  m.  alta,  stipulis  hispidis,  petiolis  45-60  cm.  longis,  laminis  38-53  em. 
longis  cordato-reniformibus  parve  9-lobatis  chartaceis  infra  pallidioribus  nervis  reticulatis 
hirsutulis  lobis  8-10  cm.  longis  rotundatis  apice  breve  acuminate  marginibus  apiculato- 
denticulatis,  pedunculis  15-20  cm.  longia  hirsutis,  inflorescentiis  37-64  cm.  longis  lance- 
oloideis  axi  primario  tomentoso  ramulis  spicatis,  floribus  rubris,  sepalis  1-1.2  mm.  longis 
oblongo-lanceolatis  ad  apicem  laceratis,  petalis  0.8-1  mm.  longis  obovatis  cucullatis 
laceratis,  antheris  1.5-1.7  mm.  longis  ovalibus,  drupis  3.5-4  mm.  diametro  luteis,  pyrena 
1.6-1.8  mm.  longa  1.4-1.5  mm.  lata  1.1-1.2  mm.  crassa  late  ovali-obovoidea  tetragona 
straminea. 

Giant  herb,  1.5  m.  tall;  stem  stout  and  fleshy;  stipules  of  seedlings  1.5-2.5 
cm.  long,  oval,  erose,  membranous,  strongly  nerved,  sparsely  hispid  on  the 
back,  those  of  mature  leaves  apparently  similar  but  narrower;  petioles  45-60 
cm.  long,  sparsely  hirsute;  blades  38-53  cm.  long,  cordate-reniform,  shallowly 
palmately  9-lobed,  firm  chartaceous,  above  dark  green,  remotely  hirsutulous, 
the  lobes  8-10  cm.  long,  semicircular,  the  end  of  the  midrib  and  one  or  more 
prominent  lateral  veins  ending  in  short  acuminations,  the  margin  irregularly 
apiculate  denticulate,  the  sinus  broad  V-shaped,  below  the  surface  pale  green, 
prominently  reticulate  veined,  subappressed  hirsutulous  on  all  the  veins  and 
veinlets ;  inflorescence  probably  axillary ;  bracts  at  base  of  inflorescence  not 
seen ;  peduncle  15-20  cm.  long,  sparsely  hirsute;  inflorescence  37-64  cm.  long, 
10-15  cm.  in  diameter,  lanceoloid,  the  axis  ferrugineous  tomentose,  bractless, 
the  lateral  branches  ascending,  spiciform,  tomentose  towards  base,  glabrous 
towards  tip ;  the  principal  lower  bracts  15-20  mm.  long,  linear  oblanceolate, 
the  margin  membranous ;  the  flowers  perfect,  bright  reddish,  sessile ;  ovary 
ovoid  truncate;  calyx  lobes  1-1.2  mm.  long,  oblong-lanceolate,  the  tip  lacerate ; 
petals  0.8-1  mm.  long,  obovate,  cucullate,  sparsely  lacerate;  the  2  anthers 
1.5-1.7  mm.  long,  oval;  the  2  stigmas  3  mm.  long,  filiform,  densely  puberulous 
throughout;  drupes  3.5-4  mm.  in  diameter,  globose,  yellow;  seeds  1.6-1.8  mm. 
long,  1.4r-1.5  mm.  wide,  1.1-1.2  mm.  thick,  broadly  oval-obovoid,  the  apex 
emarginate,  4-angled,  but  somewhat  compressed,  two  angles  equal,  the  third 
well  developed  upwards,  the  fourth  weak,  the  surface  dull,  stramineous. 

Type:  Hawaii,  Waipio,  Upper  Hamakua  Ditch  Trail,  3,500  ft.  alt.,  on 
moist  cliff,  Jan.  1,  1932,  H.  St.  John  and  E.  Y.  Hosaka  11,478.  B.  P.  Bishop 
Museum,  Honolulu. 

Collections  studied,  all  from  the  island  of  Hawaii :  Waipio  Valley,  Upper  Hamakua  Ditch 
Trail,  face  of  pali  at  head  of  valley,  alt.  1,300  m.,  Nov.  19,  1926,  L.  H.  MacDaniels  286 ; 
Kohala  Mts.,  Waimea,  Sept.  7,  1911,  C.  N.  Forbes  488.H;  Kohala,  Oct.  1931,  0.  Degener 


392  CALIFORNIA  ACADEMY  OF  SCIENCES  [Proc.  4th  Her. 

S,183 ;  Alakahi,  Kawainui  or  Kawainui  ditch  or  Kawaiiiui  Valley,  3n\y  12,  1909,  J.  F.  Eoclc, 
129,  and  4,374  or  4,375  or  4,376  or  4,382;  Waimea,  Hawaii,  June  1910.  J.  F.  Bock  8,432. 

The  specimens  with  definite  localities  all  came  from  the  windward  slope  of 
the  Kohala  Mountains,  a  region  of  very  wet,  swampy  forests,  at  the  brink  of 
the  precipitous,  deepl^y  entrenched  valleys.  Waimea  is  the  nearest  town,  with 
hotel  and  supply  base.  It  is  in  open  grasslands,  not  the  sort  of  place  where 
Gunnera  grows.  Doubtless  Waimea  was  the  base  camp  of  the  collectors,  not 
the  habitat  and  locality  of  the  plant.  Forest  ranger  Kawai  reports  another 
locality  at  the  head  of  Honopue  Gulch. 

Rock's  collection  in  1909  bears  confusing  data  on  the  four  sheets  examined. 
Alakahi  and  Kawainui  are  distinct  valleys,  both  tributary  to  Waipio.  They 
diverge  sharph%  so  it  is  not  evident  how  the  locality  could  be  in  both  valleys. 
Each  of  his  sheets  bears  two  numbers.  His  number  129  is  perhaps  a  trip  num- 
ber, while  the  others  are  apparently  serial  numbers  given  to  each  sheet  col- 
lected. Though  bearing  different  numbers  they  are  doubtless  duplicate  sheets. 
Degener's  locality  Kohala  is  also  open  to  question.  Kohala  town,  post  office, 
and  mill  are  in  the  lowland,  remote  from  the  area  where  Gunnera  is  known 
to  grow.  Perhaps  this  locality  is  to  be  taken  as  a  general  one,  referring  to  the 
Kohala  District  or  Mountains. 

This  new  species  is  dedicated  to  Miss  Alice  Eastwood,  Curator  of  Botany 
in  the  California  Academy  of  Sciences.  Her  many  decades  of  creative  work 
have  enriched  the  science  of  botany,  and  her  friendliness  and  generous  assist- 
ance have  endeared  her  to  all  botanists  who  have  consulted  her. 

5.  Gunnera  kaalaensis  (Krajina)  St.  John,  comb.  nov. 

(Plate  37,  fig.  c ;  plates  43,  44,  45 ) 
Gunnera  petaloidea  Gaudichaud  var.  Icaalaensis  Krajina,  Acta  Botanica  Bohemica  9:50, 
figs.  F„  B„  Flo,  P„  Ao,  1930. 

Giant  herb ;  stem  1-3  m.  tall,  about  15  cm.  in  diameter,  decumbent  at  base, 
the  surface  marked  with  raised,  pale,  corky  leaf -scars,  and  above  them  numer- 
ous oval,  raised  craters ;  stipules  4-8  cm.  long,  lanceolate,  scurfy  on  the  back, 
membranous,  strongly  nerved ;  petioles  3-6  dm.  long,  at  first  densely  hirsute, 
later  sparsely  so,  with  scattered  dark  tubercles;  blades  20-50  cm.  long,  cordate 
reniform,  shall owly  palmately  7-9-lobed,  firm  chartaceous,  above  dark  green, 
hirsute  on  primary  and  secondary  veins  or  glabrate,  below  paler  green, 
rough  hirsute  on  the  prominent,  reticulate  veins  and  veinlets,  the  lobes  low 
undulate,  shallow,  rounded,  3-7  cm.  long,  the  sinus  broad  U-shaped,  the 
margin  coarsely  salient  dentate;  inflorescence  axillary;  bracts  at  base  of 
peduncle  reduced,  about  2.5  cm.  long,  membranous ;  peduncle  7-12  cm.  long, 
bractless,  hirsute  but  soon  glabrate  ;  inflorescence  15-47  cm.  long,  4-14  cm.  in 
diameter,  narrowly  lanceoloid,  the  axis  hirsute  or  glabrate,  principal  lower 
bracts  10-20  mm.  long,  linear-lanceolate,  the  margin  membranous  and  not  or 
but  partly  fimbriate  ciliate,  the  lateral  branches  ascending  spiciform,  hirsute 
at  base,  glabrous  towards  tip;  flowers  perfect,  sessile,  greenish;  ovary  de- 


Vol.  XXV]       ST.  JOHN:  ENDEMISM  IN  HAWAIIAN  FLORA.  GUNNERA         393 

pressed  globose;  calyx  lobes  0.6-0.8  mm.  long,  rounded,  lacerate;  petals  1.3- 
1.8  mm.  long,  spatulate-obovate,  eucullate,  entire,  obtuse,  glabrous;  the  2 
anthers  1.1-1.3  mm.  long,  ovoid;  the  2  stigmas  about  2  mm.  long,  densely 
puberulous ;  drupes  2.8-3.1  mm.  in  diameter,  globose,  yellow ;  stone  1.8-1.9 
mm.  long,  1.4-1.6  mm.  wide,  1.3-1.5  mm.  thick,  broadly  oval-obovoid,  the  apex 
rounded  or  emarginate,  with  4  prominent  angles,  the  2  lateral  strongest  and 
equal,  the  3rd  nearly  as  strong  but  diminishing  at  base,  the  4th  nearly  as 
strong  as  the  3rd  in  the  upper  half,  in  the  lower  half  nearly  wanting,  the 
surface  yellowish  brown,  dull. 

Type:  "Oahu :  Montes  Waianae,  in  declivibus  orientalibus  sub  summo 
montis  Puu  Kaala,  1150  m.s.  m.,  legi  24.  XI.  1929"  [F.  Krajina  in  Herb. 
Charles  Univ.,  Prague,  not  seen  J . 

Collections  studied,  all  from  Oahu,  Waianae  Mts.,  Mt.  Kaala:  Kaala  Mts.,  H.  Mann  and 
W.  T.  Brigham  626;  Mt.  Kaala,  Jan.  1920,  E.  Bryan;  Mt.  Kaala,  elev.  3,200  ft.,  Feb.  11, 
1928,  H.  F.  Bergman;  Puu  Kaala,  near  summit,  Waianaeuka,  alt.  3,900  ft.,  wet  stream 
bank,  Nov.  24,  1929,  H.  St.  John  10,072 ;  Mt.  Kaala,  on  moist  upper  slope  by  spring,  elev. 
1,100  ±  m.,  April  21,  1929,  E.  H.  Brijan  Jr.  707 ;  Mt.  Kaala,  Waianaeuka,  4,000  ft.  elev., 
Feb.  2,  1930,  E.  Y.  Hosalca  139;  Scliofield  trail  to  Kaala,  steep  wet  slope,  elev.  3,300  ft., 
April  24,  1932,  E.  P.  Hume  547 ;  Puu  Kaala,  3,600-3,800  ft.,  Waianaeuka,  Jan.  8,  1933, 
V.  H.  Krauss;  Kaala,  eastern  slope,  at  summit  near  spring,  Sept.  25,  1938,  0.  Degener,  E. 
Ordonez  and  0.  Selliiig  12,251. 

Koolau  Eauge:  Konahuanui,  near  summit,  Oct.  19,  1930,  J.  R.  Coleman;  Puu  Kona- 
liuanui,  east  face,  n.e.  of  north  peak,  head  of  South  Fork  of  Kahanaiki  Stream,  Kailua, 
2,950  ft.  alt.,  thicket  on  steep  pali,  June  21,  1942,  L.  Euhanic  and  A.  D.  Conger  131.  Puu 
Konahuanui,  precipitous,  small,  hanging  valley,  east  side  of  Koolau  Divide,  head  of  North 
Fork  of  Kahanaiki  Stream,  2300  ft.  alt.,  Kailua,  May  9,  1943,  H.  St.  John  20,335. 

The  localities  for  this  species  are  discussed  in  the  introduction. 

Gunnera  kaalaensis  forms  a  narrow  belt  on  the  eastern  side  of  Mt.  Kaala. 
It  is  conspicuous  and  is  collected  by  most  botanists.  The  older  collections,  as 
that  by  Mann  and  Brigham  in  1864—65  were  probably  made  on  the  old  trail 
that  ascended  from  Waialua.  The  numerous  recent  collections  are  apparently 
all  from  the  station  on  the  trail  from  Schofield  Barracks,  above  the  Firebreak 
Trail  in  Waianaeuka.  The  collectors  have  stated  the  elevations  all  the  way 
from  3,200  to  4,000  feet.  Doubtless  all  of  the  collections  on  this  trail  were 
made  at  the  base  of  the  "pali"  (precipice),  or  by  the  "spring"  or  stream 
above  it  at  elevations  between  3,500  and  3,700  ft.  The  plant  does  not  seem 
to  occur  at  the  actual  summit.  On  the  earlier  U.  S.  Government  topographic 
map  of  1917,  the  mountain  was  given  the  Hawaiian  name,  Puu  Kaala,  and  an 
elevation  of  4,030  ft.  The  recent  one  of  1938  calls  the  peak  Mt.  Kaala,  and 
measures  it  as  4,025  ft.  high. 

The  collection  from  Puu  Kaala,  St.  John  10,072,  shoAvs  on  one  sheet  a  young 
inflorescence  which  has  the  petals  either  entire  or  sparsely  fimbriate.  This 
fringing  of  the  petals  is  not  constant  and  has  not  been  observed  in  any  of  the 
other  numerous  collections. 

Krajina  (p.  50)  in  separating  this  as  var.  kaalaensis  included  among  the 


394  CALIFORNIA  ACADEMY   OF  SCIENCES  [Proc.  4th  Ser. 

characters  that  it  had  the  blade  shining  above,  while  the  blades  of  his  var. 
mauiensis  were  not  shining.  In  this  present  revision,  this  character  is  not  sub- 
stantiated, no  diagnostic  difference  in  the  shiningness  of  the  blade  being  noted 
between  these  two  or  any  others  of  the  Hawaiian  species  of  Gimnera. 

6.  Gunnera  makahaensis  St.  John,  sp.  no  v. 

(Plate37,  fig.  b.) 

Herba  gigantea,  caule  1.5  m.  alta  15  cm.  diametro,  petiolis  ad  7  dm.  longis,  laminis  20-37 
cm.  longis  cordato-reniformibus  arete  7-9-lobatis  chartaceis  molle  hirsutis  in  nerviis  vel 
glabratis  infra  pallidioribus  molle  adpressi-hirsutis  in  nerviis  reticulatis  lobis  sinuatis 
dentieulatis  mucronatis,  pedunculo  bracteoso,  bracteis  15-20  mm.  longis  4-5  mm.  latis 
oblongo-spatulatis  marginibus  membranaceis  bifimbriatis,  pedunculo  5-7  cm.  longo,  in- 
florescentiis  15-60  cm.  longis  anguste  lanceoloideis,  axi  tomentoso,  bracteis  inferioribus 
15-35  mm.  longis,  ramulis  spicatis,  floribus  viridescentibus  sepalis  1-1.2  mm.  longis  late 
deltoideis  profunde  laceratis,  petalis  1.9-2.2  mm.  longis  obovatis  cucullatis  marginibus 
erosis  vel  laceratis  saepe  acuminatis,  antheris  1.6-2.2  mm.  longis  ovalibus,  drupis  ignotis. 

Giant  herb,  2  m.  tall ;  stem  about  1.5  m.  tall  and  15  cm.  in  diameter ;  stipules 
not  seen;  petioles  about  7  dm.  long,  somewhat  hirsute,  soon  glabrate,  sparsely 
tuberculate;  blades  20-37  cm.  long,  cordate-reniform,  shallowly  palmately 
7-9-lobed,  firm  chartaceous,  above  dark  green,  sparsely  soft  hirsute  on  the 
veins  or  glabrate,  below  pale  green,  appressed  soft  hirsute  on  prominent  veins 
and  reticulate  veinlets,  the  lobes  1.5-4  cm.  long,  shallowly  sinuate  semi- 
orbicular,  the  sinus  broad  and  open  U-shaped,  the  margin  minutely  salient 
denticulate  and  the  principal  dentations  revolute  mucronate ;  inflorescence 
probably  axillary;  bracts  at  base  of  peduncle  5-6  cm.  long,  oblanceolate, 
scurfy  towards  the  tip  and  margin ;  peduncle  when  young  clothed  with  numer- 
ous imbricate  bracts  which  are  15-20  mm.  long,  4-5  mm.  wide,  oblong  spatu- 
late,  firm,  glabrous,  with  a  conspicuous  broad  membranous  margin  lacerate 
and  ciliate  fimbriate,  the  peduncle  5-7  cm.  long ;  inflorescence  15-60  cm.  long, 
6-10  cm.  in  diameter,  narrowly  lanceoloid,  the  axis  brown  tomentose,  the  prin- 
cipal lower  bracts  15-35  mm.  long,  oblong-linear,  acute,  firm,  the  conspicuous 
membranous  margin  lacerate  and  ciliate  fimbriate,  the  lateral  branches  ascend- 
ing, densely  spicif orm,  tomentose  except  towards  the  glabrate  tip ;  flowers 
perfect,  sessile,  greenish;  ovary  depressed  globose;  calj^x  lobes  1-1.2  mm. 
long,  broadly  deltoid,  deeply  lacerate;  petals  1.9-2.2  mm.  long,  obovate  cucul- 
late,  the  margin  erose  to  lacerate,  often  acuminate ;  the  2  anthers  1.6-2.2  mm. 
long,  oval;  the  2  stigmas  1.5-2  mm.  long,  filiform,  densely  puberulous ;  drupes 
unknown. 

Type:  Oahu,  Pun  Kawiwi-Mt.  Kaala  ridge,  Makaha-Waianae  Kai,  Wai- 
anae  Mts.,  moist  thicket  in  gulch,  above  pali  [precipice],  3,500  ft.  alt., 
March  31, 1935,  H.  St.  John  17,581. 

Collections  studied:  Oahu:  Kaala  Eange,  Makaha  V[alley],  Feb.  12-19,  1909,  C.  N. 
Forbes;  Mt.  Kaala,  Waianae  Mts.,  on  wet  wooded  slope,  el.  4,000  ft.,  (specimen  brought 
by  other  students  probably  from  the  Makaha  side),  Feb.  2, 1930,  E.  Y.  HosaTca. 


Vol.  XXV]       ST.  JOHN:  ENDEMISM  IN  HAWAIIAN  FLORA.  GUNNEEA         395 

The  specimen  preserved  by  Hosaka  was  given  him  by  other  students  who 
pushed  ahead  and  crossed  the  summit  to  the  Makaha  side.  Hence,  his  general- 
ized data  and  elevation  of  4,000  feet  are  not  to  be  considered  accurate. 

It  is  remarkable  that  there  are  two  species  on  Mt.  Kaala,  G.  kaalaensis  being 
abundant  on  the  wet  windward  slope,  300  to  500  feet  below  the  summit.  G. 
makaJiaensis  is  infrequent  at  about  the  same  altitude  on  the  drier  leeward 
slope.  Actually,  the  broad,  flat  summit  is  well  watered  and  supports  a  swampy 
forest  with  a  few  boggy  openings.  Gunnera  has  not  been  found  on  the  summit. 
The  locality  of  this  new  species  is  along  one  of  the  streams  draining  from  the 
wet  summit  down  towards  the  dry  IMakaha  Valley. 

The  islands  Kauai,  Molokai,  and  Hawaii  each  produce  a  single  species.  Maui 
bears  two  species,  one  on  each  of  the  isolated  mountain  masses  of  east  Maui  and 
w^est  Maui.  Oahu  has  one  species  in  both  mountain  ranges  with  a  considerable 
gap  between  the  localities  on  Puu  Konahuanui  in  the  Koolau  Range  and  Mt. 
Kaala  in  the  Waianae  Mts.  It  is  known  only  on  the  eastern  slopes  of  Mt.  Kaala. 
Yet,  on  the  drier,  westward  slope  we  find  a  second  species.  It  is  amply  distinct 
from  its  close  neighbor  in  characters  of  sepals,  petals,  anthers,  bracts,  leaf 
blade  size,  lobing,  and  toothing.  It  is  the  only  Hawaiian  species  with  the  mar- 
ginal teeth  of  the  blade  bearing  definite  revolute  cartilaginous  mucros.  It  has 
seldom  been  collected  because  of  its  scarcity  and  the  long  difficult  ascent  to 
the  locality. 

7.  Gunnera  kauaiensis  Rock 

(Plate  37,  fig.  a.) 
Gunnera  Tcauaienis  Eock,  Oee.  Pap.  Bishop  Mus.  9(5)  :3-4,  pi.  1,  1930,  Sept. 
G.  Dominii  Krajina,  Acta  Botanica  Bohemica  9 :  50,  52,  figs.  Fj,  Bi,  Ii,  Pli,  Pj,  Ai,  1930. 

Giant  herb,  1-3  m.  tall ;  stem  1-1.3  m.  tall,  10-15  cm.  or  more  in  thickness, 
fleshy,  decumbent  at  base ;  stipules  4-5  cm.  long,  obovate,  erose,  pilose  towards 
the  margin,  membranous,  strongly  nerved;  petioles  3-6  dm.  to  "more  than  1 
meter"  in  length,  at  first  brownish  tomentose  but  soon  glabrate  except  at  base, 
purplish,  with  scattered  pustules;  blades  0.8-2  m.  in  diameter,  suborbicular 
peltate  about  %  way  from  one  side,  firm  chartaceous,  above  dark  green, 
sparsely  hirsute  to  subglabrate,  below  pale  green,  hirsute  on  the  raised  reticu- 
late veins  and  veinlets,  the  margin  palmately  9-11-lobed,  the  lobes  semior- 
bicular,  acute,  5-10  cm.  long,  with  a  broad  U-shaped  sinus,  the  margin  coarsely 
doubly  dentate,  the  teeth  2-4  mm.  long ;  inflorescence  probably'  axillary ;  basal 
bracts  not  seen ;  peduncle  6-10  or  more  cm.  in  length,  brownish  hirsute ;  in- 
florescence 3-6  dm.  long,  ("1  meter  or  more  long,"  fide  Rock),  8-17  cm.  in 
diameter,  dense,  lanceoloid,  the  axis  densely  brownish  hirsute,  at  length  sub- 
glabrate, lower  primary  bracts  15-18  mm.  long,  oblong-lanceolate  to  spatulate, 
glabrous,  the  margin  scarious,  the  lateral  branches  ascending,  spiciform, 
hirsute  at  base,  glabrate  at  tip;  flowers  perfect;  ovary  depressed  globose; 
calyx  lobes  0.3-1  mm.  long,  deltoid,  the  margin  thin,  lacerate;  petals  1.1-1.3 
mm.  long,  obovate  cucullate,  the  margin  ciliate  or  lacerate;  the  2  anthers 
1.9-2.1  mm.  long,  oval ;  the  2  stigmas  1.5-2  mm.  long,  filiform,  densely  puber- 


396  CALIFOBNIA  ACADEMY  OF  SCIENCES  [Proc.  4th  Ser. 

iilous ;  drupes  3^  mm.  in  diameter,  globose,  red  to  purplish ;  stone  1.9  mm. 
long:,  1.6  mm.  wide,  1.4  mm.  thick,  oval,  somewhat  compressed,  the  apex  emar- 
ginate,  conspicuously  3-angled,  the  2  lateral  angles  equal,  acute,  the  3rd  angle 
prominent  and  acute  in  the  upper  half,  wanting  below,  the  4th  angle  unde- 
veloped, the  surface  stramineous,  dull. 

Type:  "Kauai,  along  the  stream  beds  of  Kaluiti  and  Kailiili,  just  below  the 
summit  of  Waialeale,  elevation  4800  to  5000  feet,  September  22,  1909  (fruit- 
ing), J.  F.  Kock  no.  5053.  Type  B.  P.  Bisliop  Mus."  On  one  or  more  of  the 
five  isotypic  sheets  are  found  the  numbers  5,050,  5,053,  and  5,058.  These  are 
apparently  individual  numbers  given  by  the  collector  to  each  sheet.  Later  he 
selected  the  number  5,053  to  stand  for  the  collection.  They  all  bear  also  the 
number  129  which  perhaps  may  be  a  trip  number,  though  it  duplicates  the 
one  for  the  collection  from  Hawaii  in  the  same  year.  Rock's  three  series  of  num- 
bers on  his  collections  are  not  well  understood  by  the  botanists  of  Honolulu. 

Collections  studied,  all  from  Kauai:  Kahili  summit,  Aug.  1909,  C.  N.  Forbes;  Kilohana 
Outlook  above  Wainiha,  Waimea  Drainage  Basin,  West  Side,  July  3  to  Aug.  18,  1917, 
C.  N.  Forbes  903.K;  Kilohana  lookout,  N.  W.  end  Alakai  Swamp,  Na  Pali-Kona  forest 
reserve,  wet  steep  slope,  el.  4,023  ft.,  Dec.  27, 1930,  E.  St.  John  et  al,  10,788;  Avithout  locality, 
Oct.  1916,  J.  F.  Rock. 

Rock  (p.  3)  described  the  "Panicle  terminal."  Neither  his  specimens  nor 
any  others  of  those  available  in  Honolulu  show  an  attached  inflorescence. 
All  of  the  other  Haw^aiian  species  have  their  inflorescences  developing  near 
the  stem  tip,  but  truly  axillary,  and  it  seems  probable  that  this  one  does  too. 
The  inflorescence  is  not  a  true  panicle,  rather  it  has  an  erect  common  axis 
bearing  numerous  ascending,  unbranched  spikes. 

This  is  the  most  distinct  of  the  Hawaiian  species,  differing  markedly  from 
all  the  others.  It  alone  has  orbicular,  peltate  blades  and  drupes  that  are  red  to 
purplish.  It  also  differs  in  the  size  and  structure  of  the  flowers  and  fruits. 

As  indicated  in  the  introduction,  it  is  still  an  open  question  as  to  which  of 
the  two  specific  names  of  this  species  is  the  earlier.  G.  kauaiensis  Rock  was 
published  in  September  1930.  G.  Dominii  Krajina  was  published  in  the  same 
year,  1930  (Acta  Botanica  Bohemica  9  :50,  52,  figs..  1930).  Previous  requests 
for  the  exact  date  of  publication,  sent  to  Prague  have  been  unanswered.  Now 
we  acknowledge  with  thanks  the  assistance  of  the  librarians  of  the  Gray  Her- 
barium, the  British  Museum  of  Natural  History,  and  the  Director  of  the  Kew 
Gardens.  It  is  not  known  whether  or  not  the  several  articles  in  volume  9,  Acta 
Botanica  Bohemica  w^ere  issued  separately  at  various  dates,  or  not.  In  any 
case,  volume  9  was  commonly  distributed  as  a  single  bound  volume,  and  the 
copies  were  received  by  the  three  institutions  listed.  Gray  Herbarium  on  Oct. 
25,  1932,  Kew  on  March  6,  1933,  and  British  Museum  on  April  1,  1936,  but 
these  dates  of  receipt  are  so  late  as  to  be  of  no  help.  Krajina  lists  as  G.  peta- 
loidea  var.  mauiensis  his  own  specimen  collected  on  Maui  on  April  15,  1930. 
His  manuscript  must  have  been  writteu  later  than  April,  and  in  all  proba- 
bility after  he  returued  to  Prague.  From  this  it  can  be  deduced  onlv  that  his 


Vol.  XXV]       ST.  JOHN:  ENDEMISM  IN  HAWAIIAN  FLORA.  GUNNERA         397 

paper  was  not  published  early  in  1930.  The  first  paper  in  vol.  9  was  by  the 
editor,  Karel  Domin,  "A  new  species  of  Psitt acanthus  from  Dominica,"  which 
paper  was  not  listed  in  the  Botanisches  Centralblatt  until  July  6,  1933.  The 
succeeding  volume  10  of  Acta  Botanica  Bohemica  contained  a  chronological 
bibliography  of  Dr.  Karel  Domin's  publications.  Two  of  them  were  published 
in  volume  9,  and  they  are  listed  in  the  bibliography  as  appearing  in  1930.  In 
one  of  these  papers,  page  256,  Domin  cites  a  plant  specimen  collected  on  Sept. 
20,  1930,  which  would  imply  that  his  paper  was  printed  late  in  the  year  1930. 
Volume  12  (1934)  of  the  Prague  journal  Preslia  is  devoted  almost  entirely  to 
bibliographic  notes,  and  it  lists  Krajina's  Gunyiera  paper  as  published  in 
1930,  stating  only  the  year.  It  is  not  known  that  this  vol.  9  was  issued  only  in 
complete  volume  form,  and  the  exact  date  of  Krajina's  publication  is  still 
unknown.  Since  Rock's  publication  was  dated  Sept.  1930,  it  is  tentatively 
accepted  as  the  prior  publication. 


PLATES 


PLATE  37 

Margin  of  terminal  and  one  lateral  lobe  of  blade  of  the  Ha- 
waiian species  of  Gunner  a.  a,  G.  kauaiensis;  b,  G.  malcahaensis ; 
c,  G.  Tcaalaensis;  d,  G.  petaloidea ;  e,  G.  moloTcaiensis ;  f,  G.  East- 
woodae  ;  g,  inauiensis. 


[  4nn 


PROC.  CALIF.  ACAD.  SCI..  4TH  SERIES.  VOL.  XXV  NO.  16  [ST.  JOHN]  PLATE  37 


[  ^01  ] 


PLATE  38 

Gunnera  petalo'idea  Gaudichaud,  type  in  Herb.  Museum  d'His- 
toire  Xaturelle,  Paris. 


[402] 


PROC.   CALIF.  ACAD.   SCI..   4TH   SERIES.   VOL.  XXV.   NO.    16  [ST.  JOHN]  PLATE  38 


Photo  by  H.  St.  John. 


[403] 


PLATE  39 

Gunnera  petaloMea  Gaud.,  at  head  of  Kahana  Valley,  3,800  ft. 
alt.,  west  Maui,  October,  1942. 


[404] 


01 
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0. 


PLATE  40 

Gunnera  mauiensis  (Krajina)   St.  John,  Kula  Pipe  Line,  bank 
of  Waikamoi  Stream,  4,250  ft.  alt.,  east  Maui,  with  figure  of 

Koyal  N.  Chapman. 


[406  J 


PROC.  CALIF.  ACAD.  SCI..  4TH  SERIES.  VOL.  XXV.  NO.  16 


[ST.  JOHN]  PLATE  40 


[407] 


PLATE  41 

Gunnera  mauiensis  (Krajina)  St.  John,  Kula  Pipe  Line,  bank  of 
Waikamoi  Stream,  4,250  ft.  alt.,  east  Maui,  with  figure  of  "Worth 

Aiken,  1925. 


[408  ] 


Ul 

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_l 

Q. 

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o 


I- 


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PLATE  42 

Gunnera  Eastwoodae  St.  John,  tj-pe  specimen.  Upper  Hamakua 
Ditch  Trail,  3,500  ft.  alt.,  Waipio,  HaAvaii,  January  1,  1932. 


[  410  ] 


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a. 


PLATE  43 

Gunnera  kaalaensis   (Krajina)    St.  John,  Mt.  Kaala,  "Waiana- 

euka,  3,500  ft.  alt.,  Oahu,  with  from  left  to  right,  figures   (in 

upper  right)  of  H.  Pope  and  W.  B.  Storey. 


[412] 


PROC.  CALIF.  ACAD.   SCI..   4TH  SERIES.  VOL.  XXV.    NO.    16 


[ST.  JOHN]  PLATE  43 


Photo  by  Willis  T.  Pope. 


[413] 


PLATE  44 

Gunnera  Tcaalaensis    (Krajina)    St.  John,  Mt.  Kaala,  Waiana- 

euka,  3,500  ft.  alt.,  Oahu,  with,  from  left  to  right,  figures   (in 

foreground)  of  H.  Pope  and  W.  B.  Storey. 


[414] 


< 

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I — I 
Z 
I 
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q: 
a. 


PLATE  45 

Gunnera  Tcaalaensis   (Krajina)   St.  John,  Mt.  Kaala,  Waiana- 
euka,  3,500  ft.  alt.,  Oahu, 


[416] 


PROC.  CALIF.  ACAD.  SCI..  4TH  SERIES.  VOL.  XXV.  NO.  16 


[ST.  JOHN]  PLATE  45 


Photo  by  Willis  T.  Pope. 


[417] 


f  PLATE  46 

Distrilnition  of  Gunnera  in  the  Hawaiian  Islands. 
Square:   Gunnera  mololcaiensis ;  Erect  Triangle:   G.  petaloidea; 
Inverted  Triangle:   G.  mauicnsis ;  Shield:   G.  Eastivoodae ;  Cir- 
cle:  G.  kauaiensis ;  J:   G.  makaliaensis ;  L:   G.  kaalacntiis. 


[418] 


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0. 

1— 1 

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a. 


Marine  Biological  Labordtory 

Xj  X  IB  K.  J^  T^  -X" 


JAN  3  1  1947 

WOODS  HOLE,  MASS. 


PROCEEDINGS 

OF  THE 

CALIFORNIA  ACADEMY  OF  SCIENCES 

Fourth  Series 
Vol.  XXV,  No.  17,  pp.  421-468,  ig  text  figs.  November  15,  1946 


ALICE  EASTWOOD 
SEMI-CENTENNIAL  PUBLICATIONS* 

No.  17 

A  REVISION  OF  THE  ARTEMISIA  VULGARIS 

COMPLEX  IN  NORTH  AMERICAf 

BY 

DAVID  D.  KECK 

Carnegie  Institution  of  Washington 

Division  oj  Plant  Biology 

Stanford  University,  California 

THE  NORTH  AMERICAN  ALLIES  of  Artemisia  vulgaris  L,  an  Old  AVorld  species, 
form  a  polyploid  complex  of  great  diversity,  extending  from  Alaska  to 
Guatemala,  and  from  Hudson  Bay  and  the  Mississippi  Valley  to  the  Pacific. 
The  center  of  this  diversity  lies  in  the  western  United  States.  The  extent  of 
the  problem  posed  to  the  systematist  by  this  group  of  plants  is  at  once  appar- 
ent by  referring  to  the  last  two  complete  treatments  of  these  species.  Hall  and 
Clements  (1923)  included  under  Artemisia  vulgaris  L.,  with  fifteen  subspecies, 
all  the  American  forms  which,  but  fifteen  years  previously,  had  been  treated 
as  fifty -four  distinct  species  by  Rydberg  (1916). 

These  authors  were  alike  in  using  comparative  morphology  as  the  principal 
tool  in  their  taxonomic  work,  but  their  species  concepts  were  at  different 
poles  from  each  other.  Rydberg  had  come  to  believe  that  any  striking  morpho- 
logical form  deserved  a  name,  and  that  this  should  be  a  specific  name,  since  he 
could  see  no  advantage  in  using  trinomials.  The  significance  of  geographical 
range  was  rarely  considered.  Rydberg's  species  often  represent  morpho- 
logically well-marked  forms,  but  all  too  frequently  they  are  found  to  be  thor- 
oughly linked  with  others  by  intermediates,  or  to  be  unique  variants,  or  to 
have  no  natural  range  of  their  own.  Often  his  species  are  based  on  one  morpho- 
logical character,  which  is  soon  found  to  occur  sporadically  here  and  there 

*  Printed  from  the  John  W.  Hendrie  Publication  Endowment, 
t  Keceived  for  publication  Feb.  6,  1943. 

[421  ] 


422  CALIFOENIA  ACADEMY  OF  SCIENCES  [Proc.  4th  Ser. 

over  a  large  area  in  which  other  characters  in  the  same  plants  show  trends  of 
variation  in  other  directions.  These  limitations  in  Rydberg's  species  concept 
make  his  entities  artificial  rather  than  natnral. 

In  their  monumental  monograph,  Hall  and  Clements,  at  the  other  extreme, 
offered  the  most  conservative  species  concept  that  could  be  defended  on 
morphological  grounds.  The  result  was  the  telescoping  of  Rydberg's  fift}"- 
four  species  of  this  complex  into  one  species,  as  mentioned  above.  Later 
evaluation  of  the  same  materials  by  other  workers  has  shown  that  the  funda- 
mental units  of  Hall  and  Clements  were  carefully  established.  Other  work- 
ers have  upheld  the  majority  of  their  basic  units  but  have  often  treated 
their  subspecies  as  species  in  order  to  conform  with  more  widely  accepted 
taxonomic  practice.  Hall  and  Clements'  work  is  therefore  sound  from  the 
morphological  viewpoint,  but  the  addition  of  one  significant  new  criterion, 
chromosome  number,  has  now  made  it  possible  to  see  a  little  more  clearly  the 
limits  of  the  species  in  this  group. 

The  present  paper  takes  the  consequences  of  this  discovery  and  presents  a 
study  of  many  new  collections  that  were  not  available  to  Hall  and  Clements. 
It  also  utilizes  genetical,  garden,  and  field  studies  as  background  data  for 
the  herbarium  studies.  Several  members  of  this  complex  were  grown  in  the 
varied  environment  experiments  conducted  by  the  Carnegie  Institution  of 
Washington  along  a  transect  across  central  California,  and  some  cytological 
and  genetical  work  was  done  on  them.  The  results  of  these  experimental 
studies  have  been  reported  by  Clausen,  Keck,  and  Hiesey  (1940a).  Addi- 
tional forms  have  since  become  available  for  cytological  study.  The  herbarium 
investigation  was  made  at  the  time  when  the  results  from  the  transplant 
experiments  were  being  prepared  for  publication,  because  the  cytogenetic 
findings  showed  the  need  for  a  revised  sj'Stematic  treatment. 

The  investigations  of  Clausen,  Keek,  and  Hiesey  (1940a  and  b)  showed 
that  several  ecospecies  must  be  recognized  in  this  polyploid  complex.  On  the 
other  hand,  from  a  limited  amount  of  evidence  these  authors  suggested  the 
possibility  that,  to  use  the  biosystematic  term,  only  one  cenospecies  was  in- 
volved in  the  whole  subsection  Vulgares  of  Artemisia,  i.e.,  that  at  least  a 
limited  amount  of  interbreeding  and  gene  exchange  could  still  take  place 
among  its  members. 

The  morphological  variation  of  the  plants  composing  this  complex  is  very 
great,  making  the  delimitations  of  natural  groups  a  difficult  problem.  The 
prominent  characters  of  taxonomic  use  include  size  of  parts  such  as  leaves  and 
heads,  shape  of  leaves,  and  quality  and  quantity  of  the  pubescence.  IMost , 
of  these  are  subject  to  environmental  modification  and  must  be  interpreted 
with  that  in  mind.  Vegetative  propagation  by  means  of  rhizomes  is  well  de- 
veloped in  most  of  the  species,  and  notable  variations  may  be  perpetuated  for 
long  periods  in  this  manner.  Under  adverse  climatic  conditions,  as  in  the  far 
north  and  at  alpine  heights,  it  may  be  only  the  exceptional  year  in  which  these 
plants  are  able  to  propagate  themselves  sexually,  although  they  are  strongly 


Vol.  XXV]       KECK:  EEVISION  OF  ARTEMISIA  VULGARIS  COMPLEX  423 

established  in  these  localities  (Hulten,  1937,  p.  327;  Clausen,  Keck,  and 
Hiesey,  1940a,  p.  336).  Furthermore,  single  characters,  which  some  authors 
have  utilized  in  the  past  to  differentiate  "species"  in  this  group,  have  been 
found  very  undependable  for  this  purpose.  It  is  only  when  a  group  of  char- 
acters is  found  correlated  to  each  other  within  a  given  territory  that  the 
existence  of  a  natural  S3'stematic  unit  can  be  surmised.  Even  then  the  sys- 
tematist  must  reconcile  himself  to  having  a  residue  of  specimens  that  he  must 
arbitrarily  include  or  exclude  from  a  given  unit  of  lower  order,  because  nat- 
ural crossings  between  such  units  produce  recombinations  that  do  not  fit 
his  key. 

When  historical  precedent  has  entrenched  a  name  in  the  botanical  manuals, 
only  a  thorough  revision  of  the  group  can  properly  appraise  it.  For  example, 
Thomas  Nuttall  in  1818  proposed  four  species  in  this  group.  One,  based  on  a 
specimen  from  near  St.  Louis,  with  leaves  lightly  tomentose  on  the  upper  sur- 
face and  densely  tomentose  beneath,  he  named  A.  ludoviciana.  A  second,  based 
on  specimens  from  the  Wisconsin  area,  with  leaves  white-tomentose  on  both 
sides,  he  named  A.  gnaphalodes.  These  two  species  have  been  maintained  in 
the  majority  of  botanical  works  down  to  the  present,  yet  this  writer  agrees 
with  Asa  Gray  that  there  is  no  warrant  for  separating  these  variations  even 
in  a  subspecific  category.  The  only  characters  that  have  been  proposed  for  dis- 
tinguishing them  prove  to  be  completely  scattered  in  distribution,  wdth  every 
indication  of  complete  interbreeding  between  the  forms. 

This  revision  employs  orthodox  taxonomy  rather  than  experimental  tax- 
onomy (biosystematics),  because  the  experimental  evidence  has  been  very 
incomplete.  Consequently,  it  has  not  been  possible  to  determine  the  exact 
genetic-ecologic  boundaries  of  the  various  units,  and  thus  to  apply  more  than 
superficially  the  concept  of  species  set  down  by  Clausen,  Keek,  and  Hiesey 
(1939a).  Consequently,  most  reliance  has  necessarily  been  placed  upon  com- 
parative morphology  coupled  with  distribution;  and  all  of  the  available 
cytological  information  has  been  used.  In  order  to  recognize  units  as  com- 
parable as  possible  with  those  determined  by  experiment,  the  geographic- 
ecologic  principle  has  been  followed  in  developing  this  species  concept.  That 
is,  an  attempt  has  been  made  to  find  key  morphological  distinctions  that  mark 
groups  limited  by  natural  environmental  barriers,  whether  these  barriers 
be  geographical  or  ecological.  From  the  kind  and  amount  of  intergradation 
observed  between  the  natural  units  thus  set  off,  the  systematic  category  of 
the  units  has  been  determined. 

Under  the  biosystematic  concept  of  species  at  least  partial  genetic  barriers 
exist  between  species.  A  difference  in  chromosome  number  is  a  genetic  barrier 
to  free  intercrossing,  and  so  it  in  itself  may  be  taken  as  evidence  that  distinct 
species  are  involved,  provided  that  morphological  differences  have  had  time 
to  follow  the  genetic  break.  In  the  present  group  of  species  the  differences  in 
chromosome  number  coincide  with  major  morphological  breaks  in  each  in- 
stance examined. 


424  '  CALIFORNIA  ACADEMY  OF  SCIENCES  [Proc.  4th  Ser. 

In  the  vulgaris  complex  the  morphological  breaks  between  species  are  only 
moderately  distinct.  This  prompted  Hall  and  Clements  to  keep  all  its  units  in 
one  species.  As  compared  with  almost  all  the  other  species  of  the  section  Ahro- 
tanum  to  which  this  group  belongs,  the  specific  differences  in  the  subsection 
Vulgares  are  not  morphologically  large.  A  gauge  for  species  is  set,  however, 
by  the  chromosome  data  where  they  are  known.  With  these  as  a  basis,  the  othet 
species  have  been  delimited  as  regional  groups  of  comparable  order,  showing 
relatively  little  intergradation. 

Regional  groups  that  differ  from  each  other  in  morphology  and  distribu- 
tion, but  that  intergrade  completely  where  their  ranges  overlap,  appear  to 
lack  genetic  barriers.  Such  units  are  treated  as  subspecies,  and  all  of  these  are 
geographical.  Because  of  the  extensive  morphological  variation  in  these  plants, 
requiring  broad  subspecies,  it  is  likely  that  the  larger  subspecies  contain  more 
than  one  ecotype. 

Morphological  variants  that  from  herbarium  evidence  do  not  appear  to 
have  either  geographic-ecologic  correlation  or  genetic  barriers  separating 
them  are  not  named.  These  often  striking  variants  are  the  formae  of  many 
taxonomists;  but  such  uncorrelated  forms  cannot  be  considered  of  phylo- 
genetic  significance. 

Cytological  and  Genetical  Evidence.  The  chromosome  numbers  of  37  plants 
from  eight  American  systematic  units,  including  seven  species,  have  thus  far 
been  determined  in  addition  to  the  European  A.  vtdgaris.  These  chromosome 
numbers  have  been  reported  previously  by  Clausen,  Keck,  and  Hiesey  (1939b, 
1940a  and  b) ,  but  several  additional  collections  are  reported  in  this  paper  for 
the  first  time.  The  enumeration  of  the  collections  that  have  been  counted  fol- 
lows the  citation  of  specimens  of  the  unit  concerned. 

The  European  A.  vulgaris  is  a  diploid  species,  with  n  =  8  chromosomes.  In 
contrast,  all  the  American  forms  of  the  Vulgares  that  have  thus  far  been  in- 
vestigated follow  a  9  series  of  chromosome  numbers.  This  cj^tological  difference 
alone  is  sufficient  to  keep  all  the  American  forms  out  of  the  Old  World  vidgaris. 
Artemisia  Carruthii  and  A.  Suksdorfii  are  diploid,  with  n  =  9 ;  A.  serrata,  A. 
longifolia,  and  A.  ludoviciana  sspp.  typica  and  incompta  are  tetraploid,  with 
w  =  18 ;  and  A.  Douglasiana  and  A.  Tilesii  ssp.  unalaschcensis  are  hexaploid, 
with  n  =  27.  Chromosomes  of  these  forms  are  illustrated  in  figures  1-9.  The 
two  American  diploid  species  thus  far  known,  A.  Carruthii  and  A.  S^iksdorfii, 
have  larger  chromosomes  than  any  of  the  other  forms  including  the  diploid 
A.  vtdgaris,  with  the  chromosomes  of  A.  Suksdorfii  the  largest  of  all. 

The  little  breeding  work  that  has  been  done  with  these  plants  has  for  the 
most  part  been  reported  (Clausen,  Keck,  and  Hiesey,  1940a).  This  report 
covered  selfings  of  A.  Douglasiana  from  Santa  Barbara,  and  of  A.  ludovi- 
ciana typica  from  Manitou,  Colorado,  and  also  two  pentaploid  hybrids  that 
were  obtained  when  these  two  races  were  crossed.  An  additional  cross  has 
been  attempted  between  A.  ludoviciana  typica  from  Leevining,  California, 
and  A.  Suksdorfii  from  Trinidad,  California,  in  the  hope  that  A.  Douglasiana 


Vol.  XXV]       KECK:  BEVISION  OF  ARTEMISIA  VULGARIS  COMPLEX 


425 


Figures  1-9.  Heterotypic  metaphases  of  meiosis  in  pollen  mother  cells  and  somatic  meta- 
phases  in  roots.  Fig.  1.  Artemisia  vulgaris,  Skane,  Sweden,  Sn.  Fig.  2.  A.  Carruiliii,  Springer, 
New  Mexico,  2n  =  18.  Fig.  3.  A.  Sulcsdorfii,  White  Horse,  Washington,  On.  Fig.  4.  A.  serrata, 
St.  Paul,  Minnesota,  2n  =  36.  Fig.  5.  A.  longifolia,  Billings  County,  North  Dakota,  2w  =  36. 
Fig.  6.  A.  Douglasiana,  Medford,  Oregon,  2n  =  56.  Fig.  7.  A.  ludoviciana  typica,  Manitou, 
Colorado,  16ii  +  liv  Fig.  8.  A. ludoviciana  incompta, Yosemite  Creek, California,  18„.  Fig.9. 
A.  Tilesii  unalaschcensis,  Unalaska,  Aleutian  Islands,  27„.  Enlargement  for  all  figures:  ca.  x 
2400. 


426  CALIFORNIA  ACADEMY  OF  SCIENCES  [Proc.  4th  Ser. 

might  be  synthesized  through  amphidiploidy,  but  it  did  not  have  a  fair  chance, 
because  the  races  tested  differed  too  much  in  time  of  flowering.  The  supposed 
amphidiploid  origin  of  A.  Douglasiana  is  discussed  in  more  detail  on  page  459. 

The  Subsection  in  Perspective.  The  members  of  the  vulgaris  complex  are 
widely  distributed  across  the  North  American  continent.  There  are  two 
diploid  species  of  radically  different  appearance.  Along  the  Pacific  shores  of 
the  United  States  from  the  Canadian  boundary  to  northern  California  gro^vs 
Artemisia  Suksdorfii,  a  stout,  large-leaved  herb  in  clumps  often  more  than  a 
meter  high  and  half  again  as  broad.  On  the  more  arid  plains  and  plateaus  from 
western  Kansas  and  Colorado  to  Chihuahua  is  A.  Carruthii,  a  low  gray  clump 
with  tiny  fascicled  leaves,  in  marked  contrast  with  the  preceding  diploid. 

Through  the  midlands  of  the  continent  there  is  a  group  of  tetraploids  per- 
plexing in  their  variation.  On  the  periphery  it  is  relatively  simple  to  disjoin 
A.  serrata  and  A.  longifolia  from  the  mass,  the  former  in  the  upper  Mississippi 
Valley,  and  the  latter  spreading  somewhat  westward  from  the  prairies  and 
badlands  of  the  Dakotas.  The  remainder  goes  through  an  impressive  amount 
of  variation  across  the  western  half  of  the  United  States  and  down  the  length 
of  Mexico,  sometimes  in  accord  with  the  climatic  zone  occupied,  and  again  in 
seeming  contradiction  to  the  environment.  From  the  lowlands  and  plains  this 
distribution  may  be  traced  up  through  the  timbered  slopes  to  the  alpine  sum- 
mits, and  from  the  regions  of  abundant  rainfall  out  across  the  arid  desert 
ranges.  No  lines  of  demarcation  of  even  relative  sharpness  are  at  first  apparent, 
for  even  the  regional  facies  of  the  assemblage  show  rather  frequent  intergra- 
dation.  As  a  consequence,  this  tetraploid  diversity  is  gathered  under  one  spe- 
cific name,  A.  ludoviciana.  The  regional  races  of  individual  caste  within  it 
are  demarked  as  subspecies — seven  in  all.  Where  the  sampling  has  been  dense, 
as  in  Colorado,  the  abundance  of  examples  makes  the  broad  gamut  of  variation 
appear  but  the  more  confusing;  however,  from  this  it  becomes  apparent  that 
the  number  of  specific  barriers  to  interbreeding  in  this  assemblage  is  strictly 
limited — that  the  variation  is  intra-,  rather  than  inter-specific.  In  Mexico,  on 
the  other  hand,  additional  sampling  may  well  uncover  regional  subspecies  too 
poorly  discerned  for  recognition  at  present.  The  potential  variation  is  cer- 
tainly there.  The  increased  sampling  available  to  us  now  has  resulted  in  the 
recognition  of  the  validity  of  sulcata  and  albula  as  subspecies  of  ludoviciana 
from  the  arid  Southwest,  while  the  same  abundance  of  material  has  failed  to 
develop  any  substantial  basis  for  separating  gnaphalodes  from  ludoviciana, 
or  Wright a  from  Carruthii,  all  of  these  results  running  at  variance  with 
rather  recent  conclusions  drawn  from  less  material. 

The  two  known  hexaploids  are  A.  Douglasiana,  from  a  Pacific  Coast  region 
between  A.  Suksdorfii  and  A.  ludoviciana,  respectively  its  diploid  and  tetra- 
ploid presumed  progenitors,  its  individuals  becomiug  the  largest  in  all  the 
complex;  and  A.  Tilesii,  a  coarse  herb  with  two  subspecies  distributed  from 
the  Pacific  Northwest  to  the  Arctic  Ocean  and  to  the  western  shore  of  Hudson 
Bay,  but  obviously  coming  to  our  continent  from  Asia,  for  the  trail  is  clear 


Vol.  XXV]       KECK:  EEVISION  OF  ARTEMISIA  VULGARIS  COMPLEX  427 

both  across  Bering"  Strait  and  the  Aleutians  to  Kamchatka,  and  westward  to 
European  Arctic  Russia.  Its  near-relatives  remained  in  the  Old  World. 

The  chromosome  numbers  of  A.  Michauxiana  and  A.  Lindleyana  are  not 
known  as  yet,  and  that  information  is  desirable  for  the  completion  of  the 
picture.  The  alpine  A.  Michauxiana  is  a  quite  well-marked  entity  with  its 
small,  much  dissected  and  finely  cut  leaves,  its  large  heads  and  low  stature, 
and  its  near-restriction  to  the  northern  Rockies.  Artemisia  Lindleyana  is  a 
suffrutescent  plant — -a  distinction  from  the  usual  herbaceous  forms — with  a 
distinctive  subracemose  inflorescence  but  somewhat  variable  leaves.  Its  dis- 
tribution is  a  natural  one  along  the  low  elevation  watercourses  of  the  Columbia 
River  system. 

The  least  well-known  member  of  the  complex  is  A.  Prescottiana,  an  anoma- 
lous species  of  airy  aspect  due  to  its  long  filiform  divided  leaves,  sparse  raceme, 
and  very  slender  branching  stem.  The  root  is  gone  from  the  only  specimen  I 
have  studied,  but  it  might  even  have  been  annual.  David  Douglas  collected  it 
six  score  j^ears  ago  on  the  Quick  Sand  River,  near  The  Dalles,  but  it  seems  to 
have  escaped  recollection,  and  it  appears  to  be  uncertain  today  even  as  to 
along  which  stream  Douglas  found  it. 

Other  conclusions  on  the  derivations  of  the  various  systematic  units  that 
can  be  gathered  from  the  several  lines  of  evidence  available  are  presented  in 
the  discussion  following-  the  systematic  treatment  (pp.  465-467). 

Methods,  Materials,  and  Acknowledgments 

From  the  early  stages  of  the  investigation  it  was  apparent  that  a  study  of 
this  complex  in  any  one  herbarium  would  be  very  inadequate  because  of  the 
great  variability  found  in  nature  and  the  broad  geographic  distribution,  and 
that  it  would  be  essential  to  bring  together  an  abundance  of  herbarium  mate- 
rial for  comparison.  Consequently,  the  material  from  twelve  herbaria  was 
assembled  for  simultaneous  comparison  at  our  central  laboratory  at  Stanford. 
In  the  case  of  the  more  widely  distributed  collections,  this  made  possible  the 
comparison  of  all  the  material  from  one  locality  at  one  time.  On  account  of 
the  variation  within  one  habitat,  this  is  a  much  safer  method  than  comparing 
the  materials  in  one  herbarium  at  a  time. 

So  many  herbarium  specimens  have  been  studied  that  their  citation  in 
entirety  becomes  impractical.  Distribution  maps  therefore  indicate  wherever 
possible  the  approximate  location  of  each  collection,  leaving  for  the  citation 
lists  only  a  limited  number  of  collections.  These  have  been  chosen  because  they 
are  reasonably  representative  and  widely  distributed  to  herbaria,  or  because 
they  mark  the  range  or  otherwise  have  special  interest.  Citation  is  more  com- 
plete for  those  groups  that  are  rare,  or  that  were  previously  understood 
inadequately.  The  complete  documentation  is  to  be  found  in  those  herbaria 
in  which  material  was  studied  and  annotated.  In  so  far  as  possible  citation  is 
by  counties  in  a  north  to  south  and  east  to  west  geographical  sequence ;  the 
county  name  is  followed  by  a  colon. 


428  •  CALIFORNIA  ACADEMY  OF  SCIENCES  [Proc.  4th  Ser. 

The  descriptions  are  cross-comparable  and  made  a.s  brief  as  permissible  if 
they  are  to  have  a  utilitarian  value  in  a  group  marked  by  so  great  diversity. 
The  distribution  and  ecological  data  are  gathered  from  the  herbarium  speci- 
mens and  from  some  personal  field  knowledge  of  the  group. 

During  the  taxonomic  investigation  collections  were  studied  and  annotated 
in  the  following  herbaria.  The  abbreviations  are  those  emploj-ed  in  the  cita- 
tion of  specimens.  In  those  herbaria  marked  with  an  asterisk,  only  selected 
groups  were  examined : 

C — University  of  California,  Berkeley. 
CAS — California  Academy  of  Sciences,  San  Francisco. 
CI — Carnegie  Institution  of  Washington,  Stanford  University,  Calif. 
*Minn — University  of  Minnesota,  Minneapolis. 
NY — New  York  Botanical  Garden,  New  York. 
Ph — Academy  of  Natural  Sciences  of  Philadelphia. 
Po — Pomona  College,  Claremont,  Calif. 

RM — Eocky  Mountain  Herbarium,  University  of  Wyoming,  Laramie. 
SU — Dudley  Herbarium,  Stanford  University,  Calif. 

US — United  States  National  Herbarium,  Smithsonian  Institution,  Washington,  D.  C. 
*UW — University  of  Washington,  Seattle. 

Veg — Vegetation  Type  Map  Herbarium  of  U.  S.  Forest  Service,  University  of  California, 
Berkeley. 

To  the  curators  of  these  herbaria  grateful  acknowledment  is  made  for  the 
privilege  of  study  generously  afforded  and  for  other  help  given.  I  am  indebted 
to  the  late  Sir  Arthur  W.  Hill,  Director  of  the  Royal  Botanic  Gardens  at  Kew, 
for  the  loan  of  a  few  types  and  pertinent  specimens  from  that  institution.  Par- 
ticular thanks  are  due  my  colleague,  Dr.  Jens  Clausen,  for  critical  aid  in 
making  taxonomic  and  cytologic  judgments.  To  him  also,  together  with  my 
colleague.  Dr.  W.  M.  Hiesey,  I  am  grateful  for  a  critical  reading  of  the 
manuscript. 

Systematic  Treatment 

The  great  diversity  of  this  group  of  plants  has  been  mentioned.  One  of  the 
most  variable  organs,  which  at  the  same  time  furnishes  key  characters  of  im- 
portance, is  the  leaf.  In  figure  10  some  of  this  variation  is  shown  in  order  to 
enable  the  reader  to  visualize  the  type  of  leaf-cut  which  words  often  inade- 
quately describe.  One  is  not  to  assume  that  this  illustration  encompasses  by 
any  means  the  entire  range  of  variation  that  is  to  be  encountered  in  these 
systematic  units,  but  these  drawings  are  intended  to  supplement  the  key. 

Key  to  the  Systematic  Units 

A.  Divisions  of  lower  leaves  again  lobed  or  cleft,  5-15  mm.  wide,  directed 
forward,  green  and  essentially  glabrous  above.  Old  World ;  eastern  North 

America 1.  A.  vulgaris 

AA.  Divisions  of  lower  leaves  entire,  or  if  lobed,  the  divisions  less  than  5  mm. 
A\dde,  spreading,  or  leaves  themselves  entire. 
B.  At  least  some  of  the  leaves  bipinnately  parted  with  salient  segments 

again  toothed.  Northern  Rocky  Mts 2.  A.  Michauxiana 


Vol.  XXV]       KECK:  REVISION  OF  ARTEMISIA  VULGARIS  COMPLEX  429 


n  =  9 

SUKSDORFII 


rigure  10.  Typical  leaf  outlines  in  Artemisia  subsection  Vulgares.  x  %. 


430  CALIFORNIA  ACADEMY  OF  SCIEXCES  [Pko^^-  4th  8er. 

BB.  Leaves  entire  to  bipinnatifid  with  entire  lobes. 

C.  Leaves  linear  to  linear-lanceolate,  entire,  or  rarely  divided  into 
few  usually  caudate  lobes,  often  caudate  and  revolute-niargined, 
the  tomentum  on  the  upper  surface  usually  deciduous,  3-12  cm. 
long;  involucre  4-5  mm.  high.  North  Dakota  to  central  Wy- 
oming   3.  A.  longifolia 

CC.  Leaves  not  as  above,  or  if  slender  and  entire  the  involucre  shorter. 

D.  Leaves  finely  and  evenly  serrate,  strongly  discolored;  plant 

1-2.5  m.  tall;  heads  only  2  mm.  wide.  Upper  Mississippi 

Valley 4.  J.  serrata 

DD.  Leaves  not  finely  and  evenly  serrate. 

E.  Leaves  fascicled,  1-3  cm.  long  only,  divided  into  di- 
varicate linear-filiform  lobes  0.5-1  mm.  wide ;  panicle 
narrow,  spike-like;  herbage  mostly  gray-tomentose 
(leaves  sometimes  green  on  upper  surface),  the  heads 
tomentulose  and  ±  shining.  Kansas  and  Arizona  to 

northern  Mexico 5.  A.  Carruthii 

EE.  Leaves  not  fascicled,  or  if  fascicled  their  divisions  more 
more  than  2  mm.  wide  or  l>right  green. 
F.  Involucre  as  high  as  or  higher  than  wide  (except  in 
^  A.  ludoviciana  candicans,  but  in  that  the  leaves  not 

prominently  discolored). 
G.  Principal  leaves  narrow,  1  cm.  or  less  wide 
exclusive  of  lobes  when  present,  tomentose  on 
both  sides  or  green  above;  stems  rarely  more 
than  1  m.  tall. 

H.  Herbaceous  to  base 6.  A.  ludoviciana 

I.  Principal  leaves  entire  or  merely 
lobed. 

J.  Leaves  3-10  cm.  long,  the  mar- 
gin not  revolute. 
K.  Panicle  usually  compact, 
elongated,  not  leafy ;  leaves 
not  thin,  usually  white- 
tomentose  on  both  sides, 
rarely  bright  green  on  up- 
per surface;  involucre  3-4 
mm.  high.  Ontario  and  Mis- 
sissippi Valley  to  Washing- 
ton and  California. 

6a.  .4.  ludoviciana  ssp.  typica 
KK.  Panicle  open,  leafy  (some- 
times contracted)  ;  leaves 
thin,  usually  bright  green 
above,  white-tomentose  be- 
neath ;  involucre  2.5—3  mm. 
high.  Arizona  to  Cliihua- 

hua 6b.  A.  ludoviciana  ssp.  sulcata 

JJ.  Leaves   mostly   1-2   cm.   long, 

lanceolate,    thickish,    usually 

•  white-tomentose  on  both  sides, 

the  margin  often  narrowly  rev- 


Vol.  XXV]       KECK:  REVISION  OF  ABTEMISIA  VULGARIS  COMPLEX  431 

olute;  panicle  open,  the  heads 
pendant  in  slender  zt  leafy  ra- 
cemes. Southwestern  U.  S.  and 
northern  Mexico. 

6c.  A.  ludoviciana  ssp.  albula 
II.  Principal  leaves   ±   parted  or  di- 
vided. 
L.  Involucre  4-5  mm.  high,  4-7  mm. 
vsride;  leaves  white-tomentose 
on  both  sides,  plane.  Western 
Montana  to  northern  Utah  and 
California.  .6d.  A.  ludoviciana  ssp.  candicans 
LL.  Involucre  2.5-3.5  mm.  high,  2- 
3.5  mm.  wide;  leaves  usually 
green  on  upper  surface  (some- 
times tomentose  on  both  sides 
in  incompta). 
M.  Panicle  spike-like  or  race- 
mose;   disk-florets    15-45. 
Montane  ;  Montana  to  Utah 
and  California. 

6e.  A.  ludoviciana  ssp.  incompta 
MM.  Panicle   broader,   branch- 
ing,    leafy;      disk-florets 
6-18. 
N.  Lower  leaves  5-10  cm. 
long,  pinnately  or  ter- 
nately    divided,    the 
lobes  entire,  strongly 
discolored.  Southwest- 
ern U.  S.  to  southern 
Mexico. 

6f.  A.  ludoviciana  ssp.  mexicana 
NN.  Lower  leaves  2—5  cm. 
long,  bipinnately  dis- 
sected, green  and  ± 
glandular,  or  mod- 
erately tomentose  on 
under  surface.  S.W. 
Texas  to  S.E.  Arizona 
and  Durango. 

6g.  A.  ludoviciana  ssp.  reddens 
HH.  Suffrutescent  with  woody  base;  leaves 
narrow,  entire  to  serrate-dentate  apically, 
rarely  more  deeply  lobed;   panicle  nar- 
row,  spike-like   or   racemose.   Banks   of 

Columbia  Eiver  and  tributaries 7.  A.  Lindleyana 

GG.  Principal  leaves  broader,  1-5  cm.  wide  exclu- 
sive of  lobes  when  present,  frequently  entire, 
»  lobes  if  present  few,  short,  entire,  discolored ; 

K  stems  frequently  more  than  1  m.  tall. 


432  CALIFOBNIA  ACADEMY  OF  SCIEXCES  [Proc.  4th  Ser. 

O.  Involucre  terete  or  narrow-ovoid,  less 
than  2  mm.  wide,  yellow-green  and  shin- 
ing; ray-florets  3-7;  disk-florets  2-8; 
leaves  green  above,  silvery  tomentose 
beneath.  Immediate  Pacific  Coast  and 

Puget  Sound  region 8.  ^.  SuTcsdorfii 

00.  Involucre  campanulate,  2-3  mm.  wide,  ± 
tomentose;  ray-florets  6-10;  disk-florets 
10-25;  leaves  less  strikingly  discolored. 
Western  Washington  to  northern  Baja 

California 9.  A.  Douglasiana 

FF.  Involucre  wider  than  high. 

P.  Stem  branching  at  base ;  herbage  very  spar- 
ingly arachnoid-pilose,  somewhat  more  dense- 
ly so  (but  not  tomentose)  on  under  surface 
of  leaves;  leaves  pinnatifid  with  3-7  filiform 
divisions  less  than  1  mm.  wide ;  inflorescence 

racemose.  Columbia  Eiver 10.  A.  Prescotfiana 

PP.  Stem  unbranched  at  base ;  leaves  tomentose 
on  under  surface,  strongly  discolored,  deeply 
incised,  the  divisions  wider;  inflorescence  a 

spike-like  panicle 11.  A.  Tilesii 

Q.  Inflorescence  overtopped  by  the  leaves ; 
involucre  4-5  mm.  high,  6-9  mm.  wide ; 
heads  30-125-flowered.  Siberia  to  Hud- 
son Bay 11a.  A.  Tilesii  ssp.  fi/pica 

QQ.  Inflorescence  well  exceeding  the  leaves ; 
involucre  3-5  mm.  high,  4-8  mm.  wide; 
heads  30-55-flowered.  Northern  Japan, 
Kamchatka,  Alaska  to  Pacific  Northwest. 

lib.  A.  Tilesii  ssp.  unalascJicensis 

1.  Artemisia  vulgaris  L.,  Sp.  PI.  848, 1753. 

Figures  1  and  10 

Type  statement :  "Habitat  in  Buropae  ciiltis,  ruderatis." 
Stems  5  to  15  dm.  tall,  slender  to  moderately  stout,  herbaceous  to  base, 
commonly  unbranched;  lower  leaves  obovate  to  deltoid-ovate  in  outline, 
deeply  incised  into  3-7  lance-ovate  to  narro\vl.y  lanceolate  principal  divisions, 
which  are  for  the  most  part  again  irregularly  and  sharply  toothed  or  lobed, 
the  lobes  and  teeth  all  directed  forward,  5-12  cm.  long,  often  nearly  as  wide, 
green  and  glabrous  or  nearly  so  above,  densely  white-tomentose  beneath,  the 
margin  obscurely  revolute ;  inflorescence  an  ample  leafy  panicle  with  ascend- 
ing branches  crowded  with  subsessile  heads,  3-15  cm.  broad ;  heads  medium- 
sized,  erect  or  nodding;  involucre  campanulate,  3-4  mm.  high.  2.5-3.5  mm. 
wide,  moderately  tomentose,  with  9-13  bracts ;  ray-florets  6-9 ;  disk-florets 
13-20. 

Common  throughout  Europe  and  eastward  to  Siberia ;  probably  adventive 
in  North  America,  but  in  places  appearing  indigenous,  occurring  irregularlj'' 
from  Newfoundland  to  Manitoba  and  Alabama. 


Vol.  XXV]       KECK:  BEFISION  OF  ARTEMISIA  VULGAEIS  COMPLEX  433 

Material  of  this  species  was  examined  in  only  three  herbaria.  In  addition 
to  the  localities  cited  bv  Hall  and  Clements  (1923),  collections  should  be 
reported  from  the  states  of  New  Hampshire,  Vermont,  Massachusetts,  and 
North  Carolina. 

The  chromosomes  were  counted  in  meiosis  of  plants  grown  from  seed  col- 
lected in  Skane,  southern  Sweden,  and  kindly  sent  by  Dr.  Gote  Turesson. 

2.  Artemisia  Michauxiana  Bess,  in  Hook.,  Fl.  Bor.  Amer.  1 :324,  1833. 

Figures  10  and  11 

Type  statement :  "Rocky  Mountains.  Douglas.  Columbia.  Douglas,  in  Herh. 
Lindl."  The  type  is  in  the  Lindley  Herbarium,  according  to  a  note  attached 
to  a  sheet  in  the  Bentham  Herbarium  at  Kew,  which  is  said  to  correspond 
exactly  with  the  type  (fide  R.  A.  Rolfe) .  This  Kew  sheet  has  been  labeled  in 
Douglas'  hand,  "In  fissures  of  dry  rocks  near  the  Kettle  Falls  and  sources  of 
the  Columbia.  1826."  A  piece  of  this  sheet  was  removed  and  sent  to  New  York, 
where  it  has  been  labeled  as  part  of  the  type.  The  first  collection  cited  doubt- 
less came  from  the  Rocky  Mountains  of  Canada;  the  second  may  have  come 
from  the  state  of  Washington. 

Artemisia  discolor  Dougl.  ex  Bess.,  Bull.  Soc.  Nat.  Mosc.  9:46,  1836.  The  type  has  been 
studied  on  loan  from  Kew.  It  was  collected  "on  rocky  situations  near  Spokane  and 
the  Kettle  Falls,"  by  David  Douglas.  It  is  a  somewhat  coarser  plant  than  the  type 
of  A.  Michauxiana,  which  came  from  a  higher  elevation,  but  the  two  can  scarcely  be 
separated  morphologically.  Its  inflorescence  is  considerably  more  open-paniculate 
than  that  of  Michauxiana,  but  this  character  proves  to  be  variable  and  lacking  in 
geographic  significance  in  this  region  as  is  also  the  vesture  of  the  herbage. 

A.  graveolens  Eydb.,  Bull.  Torr.  Club  24:296,  1897.  Leaves  greener  than  in  the  type  of 
Michauxiana,  but  much  dissected  and  to  be  classed  here  ratlier  than  in  incompta. 
Type  locality:  Yogo  Baldy,  Little  Belt  Mts.,  Montana. 

A.  subfflabra  A.  Nels.,  Bull.  Torr.  Club  27:36,  Jan.  24,  1900.  The  same  form  as  A. 
graveolens.  Type  locality :  stony  banks  of  Yellowstone  Eiver,  near  Yancey's,  Yellow- 
stone Park,  "no.  5743." 

A.  tenuis  Eydb.,  Mem.  N.  Y.  Bot.  Gard.  1:431,  Feb.  15,  1900.  A  slender,  small-headed  form 
of  graveolens,  i.e.,  the  green  form  of  the  species.  Type  locality:  Emigrant  Gulch. 
Park  Co.,  Montana. 

A.  vulgaris  ssp.  discolor  Hall  et  Clem.,  Carn.  Inst.  Wash.  Publ.  326 :  74,  1923. 

A.  vulgaris  var.  discolor  Jeps.,  Man.  Fl.  PI.  Calif.  1141,  1925. 

A.  vulgaris  ssp.  Michauxiana  St.  John,  Ees.  Stud.  St.  Coll.  Wash.  1:106,  1929. 

A.  vulgaris  ssp.  Michauxiana  var.  typica  St.  John,  ihid. 

A.  discolor  var.  glandulifera  Hend.,  Ehodora  32:27,  1930.  Leaves  not  tomentose  beneath 
but  glandular.  Type  locality:  eastern  base  of  Steins  Mts.,  Harney  Co.,  Oregon. 

A.  vulgaris  var.  glandulifera  Peck,  Man.  Higher  Pis.  Ore.  766,  1941. 

Stems  2-4 (-7)  dm.  tall;  herbage  white-tomentose  throughout  when  young 
to  nearly  glabrous  and  green ;  leaves  short  and  broad,  rather  crowded,  com- 
monly bipinnately  divided  with  the  secondary  segments  again  toothed,  the 
linear  lobes  widely  spreading,  acute,  rarely  elongated,  even  the  uppermost 
leaves  rarely  entire,  2-5  cm.  long,  typically  bright  green  above  and  white- 


434 


CALIFORNIA  ACADEMY  OF  SCIENCES 


[Proc.  4th  Ser. 


Figure  11.  Distribution  of  Artemisia  Michauxlana. 


tomentose  beneath,  the  margin  plane  or  narrowly  revolute;  inflorescence  a 
slender  racemose  or  spike-like  panicle,  occasionally  more  open,  not  leafy; 
heads  large,  nodding  at  least  at  first ;  involucre  hemispheric,  3.5-4  mm.  high, 
(2.5-) 3.5-5  mm.  wide,  glabrous  or  sparingly  tomentose,  with  9-16  bracts; 
ray-florets  9-12 ;  disk-florets  15-35. 


Vol.  XXV]       KECK:  EEVISION  OF  ARTEMISIA  VULGARIS  COMPLEX  435 

Rocky  Mountains  of  northern  Alberta  and  British  Columbia,  southward 
through  the  mountains  to  northern  Wyoming,  northern  Idaho,  and  central 
Washington;  local  in  Steins  Mountains,  Oregon,  and  in  Washoe  County, 
Nevada.  Occurring  at  elevations  of  500  to  2750  meters,  and  flowering  from 
July  to  August. 

Alberta.  Moberly's  on  Atliabaska,  Scliaffer  1613 ;  Cataract  Creek,  Brown  1471 ;  Poboctan 
Pass;  Lake  Louise;  Vermilion  Mt. ;  Highwood  Eiver  District,  Brln'kDian  3209.  British 
Columbia.  Mt.  Atlin,  Lake  Atlin  region,  Swarth  177 ;  Telegrapli  Creek,  Walker  1208.  Cari- 
boo: Mt.  Sehvyn,  Raup  and  Abbe  3946.  Kootenay:  Mt.  Whaleback,  Yolio  Park,  JJllce  334; 
Mt.  Wapta;  Kicking  Horse  Eiver;  Carbonate  Draw,  Selkirk  Mts.,  Shaw  335 ;  Goldstream; 
Revelstoke.  Yale:  Eraser  Valley;  Lake  House,  Skagit  River,  Macoun  76927.  New  West- 
minster: Yale;  Chilliwack  Valley.  Montana.  Glacier:  Midvale,  Umbaoh  275 ;  Many  Glacier. 
Flathead:  Avalanche  Lake;  Columbia  Falls,  Williams  709 ;  Mission  Creek.  Lincoln:  To- 
bacco Mts.  Judith  Basin:  Yogo  Baldy,  Aug.  24,  1896,  7000  ft.,  J.  E.  Flodman  881  (NY, 
type  of  A.  graveolens)  ;  Long  Baldy,  Little  Belt  Mts.,  Aug.  19,  1896,  7-8000  ft.,  Flodman 
881  (US,  labeled  isotype  of  A.  graveolens).  Deer  Lodge:  Eyans  Lake;  Mt.  Haggin.  Galla- 
tin: Mt.  Bridger.  Park:  Emigrant  Gulch,  6500  ft.,  Rydberg  and  Bessey  5201  (NY,  type 
of  A.  tenuis;  isotype,  US)  ;  Electric  Peak.  Wyoming.  Sheridan:  Little  Tongue  Eiver  Can- 
yon, Williams  3233.  Johnson:  headwaters  of  Clear  Creek  and  Crazy  Woman  Eiver  (NY). 
Big  Horn:  Doyle  Creek,  Goodding  385.  Park:  Meeteetse  Creek,  Griffiths.  Yellowstone  Park: 
Yellowstone  Eiver  near  Junction  Butte,  Nelson  5743  (EM,  type  of  A.  sub  glabra;  isotype, 
NY).  Teton:  Leigh  Lake,  Williams  994.  Utah.  Summit:  Blacks  Fork,  Uinta  Mts.;  Upper 
Henry's  Fork  Basin,  Maguire  et  al.  14412,  14586.  Idaho.  Idaho :  mouth  of  Salmon  Eiver, 
Eastwood  13337.  Washington.  Okanogan:  Oroville;  Mt.  Chapaca,  Elmer  589;  Tonasket; 
Hidden  Lakes;  Angels  Pass,  Thompson  7033.  Whatcom:  Newhalem,  Muenscher  9919. 
Chelan:  Nigger  Creek;  Three  Brothers.  Yakima:  Mt.  Adams,  Suksdorf  (C).  Oregon. 
Harney:  Steins  Mts.,  Cusick  1991;  alpine  rivulet  near  eastern  base  of  Steins  Mts.,  beyond 
Alvord,  Henderson  8489  (CAS,  isotype  of  A.  discolor  var.  glandulifera) .  Nevada.  Washoe: 
head  of  Galena  Creek,  8500  ft.,  Heller  10222. 

The  exact  southern  limits  of  this  species  are  difficult  to  determine  on  mor- 
phological criteria  alone.  Plants  from  the  alpine  zone  in  parts  of  Nevada,  as 
in  the  Ruby,  White,  and  Charleston  mountains,  which  appear  very  similar  to 
the  tenuis  form  of  Michauxiana,  are  referred  in  this  paper  to  A.  ludoviciana 
ssp.  incompta.  This  is  because  intergrades  seem  to  connect  these  forms  with 
very  narrow,  dark  green  leaf -segments  with  forms  having  the  more  usual  type 
of  incompta  leaf,  and  these  Michaiixiana-WkQ  forms  are  fairly  well  isolated 
from  the  other  members  of  Michmixiafia.  Their  disposition  would  be  more 
satisfactorily  made  if  cytological  data  were  available. 


3.  Artemisia  longifolia  Nutt.,  Gen.  2  :  142, 1818. 

Figures  5,  10,  and  12 

Type  statement :  "In  rocky  situations,  on  the  banks  of  the  Missouri,  from 
the  confluence  of  White  River  to  the  Mountains?"  A  single  leaf  preserved 
at  the  Philadelphia  Academy,  labeled  "Artemisia  longifolia  Missouri"  in 
Nuttall's  hand,  is  doubtless  from  the  type. 


436 


CALIFORNIA  ACADEMY  OF  SCIENCES 


[Proc.  4th  Ser. 


A.  ludovieiana  var.  integrifolia  A.  Nels.,  First  Eep.  Fl.  Wyo.  138,  1896.  Based  upon  A. 

integrifolia  of  Pursh,  Fl.  Amer.  Sept.  2:520,  1814.  Not  A.  integrifolia  L.,  1853. 

Pursh  attempted  to  identify  a  specimen  collected  on  bluffs  of  the  Missouri  Eiver, 

October  1,  1804,  by  Lewis,  no.  53,  witli  the  Linnaean  species.  Lewis'  specimen  at 

the  Philadelphia  Academy  has  been  examined. 
A.  naironensis  A.  Nels.,  Bull.  Torr.  Club  26:485,  1899.  A  wide-leaved  form  of  the  species. 

"Type  no.  is  568,  Willow  Creek  [Wyoming],  July  20,  1894,  distributed  as  a  variety 

of  A.  ludovieiana." 
A.  falcata  Eydb.,  N.  Amer.  Fl.  34:271,  1916.  The  form  with  salient  linear  lobes  on  the  very 

narrow  leaves,  but  in  all  other  respects  typical.  Found  on  the  southeastern  border 

of  the  range  of  this  species;  the  type  locality  is  Fort  Pierre,  South  Dakota. 


Figure  12.  Distribution  of  Artemisia  longifolia  and  A.  serrata. 

A.  vulgaris  ssp.  longifolia  Hall  et  Clem.,  Carn.  Inst.  Wash.  Publ.  326:78,  1923. 
A.  vulgaris  var.  longifolia  Peck,  Man.  Higher  PI.  Ore.  767,  1941. 

Stems  2-8  dm.  tall,  usualh-  rather  stout,  erect,  herbaceous  down  to  the 
woody  caudex,  white-tomentose ;  leaves  linear  or  linear-lanceolate,  merely 
acuminate  or,  more  often,  caudate-attenuate,  usually  entire,  sometimes  with 
a  few  slender  teeth  or  lobes,  occasionally  the  lobes  caudate-elongated,  3-12 
cm.  long,  2-5  (-6)  mm.  wide,  those  of  the  inflorescence  very  narrow  and  short, 
aJl  loosely  floccose  and  usually  becoming  glabrate  or  green  above,  densely 
tomentose  beneath,  often  strongly  revolute ;  inflorescence  a  strict  short  panicle, 
sometimes  elongated  or  rather  open;  heads  mostly  erect  and  peduncled;  in- 
volucre campanulate  or  ±  turbinate,  4-5  mm.  high,  3-4  mm.  wide,  tomentu- 
lose  to  densely  tomentose,  with  9-17  bracts ;  ray-florets  3-10;  disk-florets  8-26. 
n  =  18. 


Vol.  XXV]       KECK:  REVISION  OF  ARTEMISIA  VULGARIS  COMPLEX  437 

Prairies  and  badlands  from  Saskatchewan  and  Alberta  to  central  North 
and  South  Dakota,  eastern  Montana,  and  southern  Wyoming,  often  in  heavy 
alkaline  clays.  Flowering  from  July  to  September. 

Saskatchewan.  1858,  Bourgeau.  Alberta.  W.  of  Lacombe,  Moss;  Eosedale,  Moodie 
1097.  North  Dakota.  McLean,  Morton,  McKenzie,  Stark,  Billings,  and  Golden  Valley 
counties.  South  Dakota.  Evarts,  Missouri  Eiver ;  Grand  River.  Stanley :  Fort  Pierre,  Sept. 
29,  1897,  Griffiths  757  (US,  type  of  A.  falcata;  isotype,  US;  phototypes,  C,  NY,  US). 
Washaliaugh :  Pine  Ridge  Reservation.  Washington :  Sheep  Mt.  Harding :  Glaring  Fork ; 
Short  Pine  Hills.  Lawrence  (?):  Crow  Buttes.  Pennington:  Big  Bad  Lands,  Osterlwut 
7529.  Montana.  Milk  River.  Dawson:  Colgate,  Sandberg  et  al.  1011;  Glendive.  Toole: 
Shelby,  Jones.  (Po).  Wyoming.  Natrona:  Willow  Creek,  Nelson  568  (RM,  type  of  A. 
natronensis ;  isotype,  Minn).  Johnson:  Lake  DeSmet,  Nelson  S545.  Carbon:  Como  Ridge 
near  Medicine  Bow,  Osterlwut  6290.  Converse :  Badlands,  T.  38,  R.  68.  Fremont :  Muskrat 
Creek,  Goodding  517.  Sweetwater:  Table  Rock;  Point  of  Rocks,  Nelson  8148. 

Chromosome  counts  were  made  in  root-tips  of  four  plants  from  Billings 
County,  North  Dakota.  The  living  plants  were  supplied  us  through  the  kind- 
ness of  Mr.  E.  C.  Moran. 

4.  Artemisia  serrata  Nutt.,  Gen.  2 :  142, 1818. 

Figures  4,  10,  and  12 
Type  statement :  "Near  the  Prairie  du  Chien,  on  the  banks  of  the  Mississippi, 
also  on  the  banks  of  the  Missouri,  in  open  alluvial  soils."  Upper  Louisiana, 

Nuttall  (Ph,  type). 

Artemisia  ludoviciana  var.  serrata  ToRR.  et  Gray,  F1.  N.  Anier.  2:420,  1843. 
A.  vulgaris  ssp.  serrata  Hall  et  Clem.,  Carn.  Inst.  Wash.  Publ.  326:79,  1923. 

Stems  10-25  dm.  tall,  virgate,  herbaceous;  leaves  lanceolate  to  linear- 
lanceolate,  acuminate,  sharply  and  rather  finely  serrate  from  apex  well  down 
toward  base  (rarely  subentire),  8-12(-15)  cm.  long,  12-20  mm.  wide,  those 
of  the  inflorescence  prominent  but  gradually  reduced  in  size,  all  dark  green 
and  glabrous  above,  white-tomentose  beneath;  inflorescence  a  leafy  panicle 
5-15  cm.  broad;  heads  small,  mostly  erect,  rather  crowded;  involucre  cam- 
panulate,  2.3-3  mm.  high,  ca.  2  mm.  wide,  slightly  floccose,  with  8-12  bracts; 
ray-florets  6-11 ;  disk-florets  3-16.  n  =  18. 

Upper  Mississippi  Valley ;  Minnesota  to  Illinois  and  Iowa ;  introduced  in 
New  York.  Flowering  from  July  to  September. 

Illinois.  Winnebago,  Jo  Daviess,  Rock  Island,  and  Henderson  counties.  Wisconsin. 
Dane:  Wiibe  (US).  Minnesota.  Chisago:  Taylors  Falls,  B.  C.  Taylor.  Isanti:  Sandberg 
(C).  Mille  Lacs:  Milaca,  Sheldon  3278.  Stearns:  Grand  Lake;  Rockville,  Campbell.  Henne- 
pin :  Fort  Snelling ;  Minnesota  River  near  mouth  of  Nine  Mile  Creek.  Wabasha :  Lake  City. 
Winona:  Wiscoy,  Holzinger ;  Winona.  Houston:  Jefferson.  Iowa.  Fayette:  Finlc  406. 
Poweshiek:  Grinnell.  Jasper:  Baxter  (US). 

Chromosome  counts  were  made  in  root-tips  of  two  plants  from  St.  Paul, 
Minnesota.  These  plants  were  supplied  through  the  kindness  of  Dr.  C.  O. 
Rosendahl. 


438  CALIFOBXIA  ACADEMY  OF  SCIENCES  [Proc.  4th  Ser. 

5.  Artemisia  Carruthii  Wood  ex  Cairuth, 
Trans.  Kans.  Acad.  Sci.  5  :  51, 1877. 
Figures  2, 10,  and  13 
Type  locality :  Kansas.  Type  not  seen. 

Artemisia  Wrightii  Gray,  Proc.  Amer.  Acad.  19:48,  1883.  The  type  collection,  Wrlglit 
1279,  made  at  Santa  Rita  del  Cobre,  Grant  Co.,  New  Mexico,  is  scarcely  morphologi- 
cally distinguishable  from  material  from  the  type  region  of  Carruthii.  The  leaves 
are  greener  and  glabrate  on  the  upper  surface  in  Wrightii,  tomentose  in  Carruthii, 
but  this  feature  does  not  appear  in  this  case  to  distinguish  ecotypes  or  geographical 
sulispecies.  While  the  Wrightii  form  does  not  occur  to  the  eastward,  it  grows  together 
with  the  typical  form  at  many  localities  in  New  Mexico.  The  very  broad  range  of 
overlapping  of  the  two  forms  largely  obscures  any  difference  in  ecological  preference 
that  may  exist  betAveen  them,  although  the  incipient  stages  of  ecotype  formation 
may  be  involved. 

A.  kansana  Britt.  in  Britt.  and  Brown,  111.  Fl.  3  :466,  1898.  A  form  identical  with  Carruthii. 
Type  locality:  Lane  County,  Kansas. 

A.  coloradensis  Osterh.,  Bull.  Torr.  Club  27:506,  1900.  Leaf-segments  unusually  wide, 
otherwise  indistinguishable.  In  the  type,  from  Dale  Creek,  Larimer  Co.,  Colorado, 
the  segments  are  less  than  1  mm.  wider  than  those  of  Carruthii  from  Kansas,  but 
other  collections  from  the  same  county  are  broader  and  intergrade  toward  the  Brit- 
tonii  form  of  ^.  ludoviciana  typicd. 

A.  Balceri  Greene,  PI.  Baker.  3:31,  1901.  Allied  to  the  Wrightii  form  in  that  the  leaves 
are  green  on  the  upper  surface,  but  these  are  atypically  scattered  and  wide,  giving 
some  suggestion  of  an  extremely  narrow-leaved  form  of  A.  ludoviciana  mexica.'na. 
This  does  not  appear  to  warrant  separation  from  the  more  usual  forms  of  Carruthii, 
however,  for  plants  of  this  appearance  are  found  in  the  western  portion  of  the  range 
of  the  species  in  scattered  stations  between  which  more  typical  forms  are  found.  In 
other  words,  the  variant  is  of  spasmodic  occurrence  and  by  no  means  of  uniform 
appearance.  The  inflorescence  is  usually  open-paniculate.  Type  locality:  canyon  of 
the  Gunnison  River,  near  Cimmaron,  Colorado. 

A.  Pringlci  Greenm.,  Proc.  Amer.  Acad.  40:50,  1904.  Heads  larger  than  usual,  containing 
tvdce  the  average  number  of  florets  for  the  species  as  a  whole  in  the  type  collection 
(ca.  18  involucral  bracts,  14-18  ray-florets,  20-34  disk-florets),  but  subject  to  con- 
siderable variation  in  the  region  from  which  the  type  came.  Not  readily  separable  by 
any  character  from  Carruthii.  Typo  locality :  plains  near  the  city  of  Chihuahua. 

A.  Wrightii  coloradensis  A.  Nels.  in  Coult.  and  Nels.,  Man.  Rocky  Mt.  568, 1909. 

A.  mexicana  Balceri  A.  Nels.,  ihid.  569. 

A.  vulgaris  ssp.  Wrightii  Hall  et  Clem.,  Carn.  Inst.  "Wash.  Publ.  326 : 80, 1923. 

A.  vulgaris  var.  Wrightii  Palmer  et  Steyerm.,  Ann.  Mo.  Bot.  Gard.  22:677,  1935  ;  Rhodora 
40:134,1938. 

A.  vulgaris  Carruthii  F.  C.  Gates,  Trans.  Kansas  Acad.  Sci.  42 :  138,  1940. 

A.  CarruthiiYar.  Wrightii  Blake,  Jour.  Wash.  Acad.  Sci.  30:472, 1940. 

Stems  3-6  dm.  tall;  herbage  white-tomentose  thronphont  or  the  upper  sur- 
face of  the  leaves  preen ;  leaves  divided  into  divaricate  linear-filiform  lobes, 
these  sometimes  again  toothed,  1-3  cm.  long,  those  of  sterile  shoots  and  of  the 
inflorescence  often  entire,  reduced,  the  margin  closely  revolute;  inflorescence 
a  leafy  narrow  spike-like  panicle,  1-3  cm.  wide,  occasionally  rather  expanded ; 


Vol.  XXV]       KECK:  BKVISION  OF  ABTEMISIA  VULGABIS  COMPLEX  439 


Figure  13.  Distribution  of  A.  Carruthii. 

heads  small,  nodding  when  young;  involucre  eampanulate,  3-3.5  mm.  high, 
1.5-3  mm.  wide,  tomentulose,  often  shining,  with  8-14  bracts;  ray-florets 
6-14 ;  disk-florets  6-20.  n  =  9. 

Western  Kansas  and  northern  Texas  to  central  Utah,  Arizona,  and  northern 
Chihuahua,  at  elevations  from  800  to  3000  m.  Adventive  in  Rhode  Island, 
Indiana,  Missouri,  and  eastern  Kansas.  Flowering  from  late  July  through 
September. 


440  CALIFORNIA  ACADEMY  OF  SCIENCES  [Proc.  4th  Ser. 

Kansas.  Eooks:  Rockport.  Lane:  plains,  Aug.  15,  1895,  HitchcocTc  302  (NY,  type  of  A. 
Icansana;  isotypes,  RM,  US).  Greeley:  Tribune.  Kearney:  Hitchcoclc.  Hamilton:  Syracuse, 
Bose  and  Fitch  17010.  Clark:  White.  Meade:  Meade.  Seward:  Liberal.  Texas.  Potter:  Ama- 
rillo,  Bailey  12S.  Randall:  Paloduro  Canyon.  Colorado.  Larimer:  Dale  Creek,  Sept.  7,  1899, 
Osterhout  2010  (RM,  type  of  A.  coloradensis ;  isotypes,  NY,  Po,  RM).  Eagle:  Red  Cliff. 
Park:  Como.  Douglas:  Pine  Grove.  El  Paso:  Palmer  Lake;  Colorado  Springs.  Teller: 
Florissant.  Fremont:  Canon  City;  Wet  Mountain  Valley.  Saguache:  Marshall  Pass,  10,000 
ft.,  BaTcer  879 ;  San  Luis  Valley.  Montrose:  Bedrock.  Gunnison:  Black  Canyon,  Baker  698 
(C,  NY,  Po,  RM,  SU,  US,  topotypes  of  A.  Bakeri).  Ouray:  5  mi.  N.  of  Ouray.  San  Miquel: 
Norwood  Hill.  Huerfano :  Cucharas  Valley.  Costilla  :  La  Veta  Pass.  Mineral :  Wagon  Wheel 
Gap.  Hinsdale:  Lake  City.  Archuleta:  Piedra.  La  Plata:  W.  of  Hesperus.  New  Mexico. 
Union:  Folsom.  Colfax:  Ute  Park.  Rio  Arriba:  Dulce.  San  Juan:  Tunitcha  Mts.,  Standley 
7622.  Quay:  Nara  Visa;  Llano  Estacado.  San  Miguel;  Las  Vegas.  Santa  Fe:  Canoncito. 
Sandoval:  Jeniez  Springs.  Valencia:  Grants.  Lincoln:  Ruidoso.  Dona  Ana:  Organ  Mts. 
Socorro :  Negrito  Ranger  Station.  Catron :  Patterson.  Grant :  Santa  Rita  del  Cobre,  Wright 
1279  (Gray  Herb.,  type  of  A.  Wrightii;  isotype,  NY;  phototypes,  C,  NY);  N.  of  Cliff. 
Hidalgo:  Cloverdale,  Mearns  456.  Utah.  Sevier:  Mt.  Marvin,  Rydberg  and  Carlton  7567. 
Piute :  Circle  Valley.  Garfield :  Panguitch  Lake.  San  Juan :  Montezuma  Canyon,  E.  of 
Monticello,  Rydberg  and  Garrett  9687 ;  Elk  Mts.;  Bluff.  Arizona.  Apache:  Carrizo  Mts., 
Standley  7510 ;  Greer.  Navajo:  Oraibi,  Voth  76.  Coconino:  Grand  Canyon;  Mt.  Humphreys; 
7  mi.  E.  of  Flagstaff;  Bellemont,  Jones  4074 ;  Williams.  Mohave:  Hualpai  Mts.,  Kearney 
and  Peebles  12727.  Yavapai:  Prescott.  Gila:  Black  River  Bridge.  Greenlee:  Metcalf.  Gra- 
ham: Mt.  Graham.  Cochise:  Chiricahua  Mine,  Blumer  1794.  Chihuahua.  Lake  Santa  Maria; 
plains  near  Chihuahua,  Pringle  625  (Gray  Herb.,  type  of  A.  Pringlei;  isotj'pes,  Ph,  SU, 
US)  ;  Santa  Eulalia  Plains,  Wilkinson.  Chihuahua  or  Coahuila.  Charco  de  Grado,  Oct. 
1852,  Thurber  791.  Sonora.  San  Luis  Mts.,  Mearns  2461;  Badehuachi,  Lloyd  404. 

Chromosome  counts  were  made  in  root-tips  of  three  plants  grown  from  seed 
from  Springer,  Colfax  County,  New  Mexico,  and  kindly  supplied  by  Mrs. 
Gladys  Nisbet. 

In  Colorado,  on  the  one  hand,  occasional  collections  appear  to  be  inter- 
mediate with  the  Brittonii  form  of  A.  lucloviciana  typica,  while  in  New 
Mexico,  on  the  other,  certain  collections  are  found  that  are  intermediate  with 
A.  ludoviciana  mexicana.  More  chromosome  counts  will  help  clarify  this 
situation  and  show  whether,  as  is  now  suspected,  all  forms  of  liidovwiana  are 
tetraploid,  and  of  Carruthii  diploid. 

6.  Artemisia  ludoviciana  Nutt.,  Gen.  2 :  143, 1818. 

6a.  Artemisia  ludoviciana  Nutt.  ssp.  typica  Keck,  Cam.  Inst. 
Wash.Publ.  520:330, 1940. 

Figures  7,  10,  and  14 
Type  statement:  "On  the  banks  of  the  Missisippi.  near  St.  Louis;  also  on 
the  alluvial  plains  of  the  Missouri."  A  specimen  at  Philadelphia  from  St. 
Louis,  collected  and  starred  by  Nuttall,  is  the  type.  It  consists  of  only  four 
leaves,  lightly  tomentose  above  and  densely  tomentose  beneath.  Another 
Nuttall  specimen  at  Philadelphia,  labeled  by  Nuttall  "Artemisia  Ludoviciana. 
Platte  plains,"  is  not  starred,  but  is  a  large  branch  in  flower  with  all  leaves 
entire  and  tomentose  on  both  sides. 


Vol.  XXV]       KECK:  EEVISION  OF  ABTEMISIA  FULGAEIS  COMPLEX 


441 


Arternisia  ludoviciana  Nutt.,  loc.  cit. 

A.  gnaphalodes  Nutt.,  ihid.  Type  statement:  "On  dry  savannahs  about  Green  Bay,  Lake 
Michigan,  and  on  the  banks  of  Fox  river,  and  the  Missouri."  A  starred  specimen  at 
the  Philadelphia  Academy,  collected  by  Nuttall  at  St.  Louis,  was  considered  by  Hall 
and  Clements  to  be  the  type,  but  its  location  is  inconsistent  with  the  type  statement. 
It  consists  merely  of  the  top  of  a  flowering  stem,  in  which  the  leaves  are  white- 
tonientose  on  both  sides.  See  discussion  of  this  synonym  on  p.  423. 

A.  Pursliiana  Bess,  in  Hook.,  Fl.  Bor.  Amer.  1:323,  1833.  Type  locality:  "On  the  plains  of 
the  Saskatchewan.  Dr.  RicJmrdson.  Drummond."  See  the  following. 


Figure  14.  Distribution  of  A.  ludoviciana  ssp.  typica. 


A.  Pursliiana  a  latifolia  Bess.,  ibid.  Specimen  at  Kew  (Herb.  Hooker),  which  has  been 
studied,  taken  in  the  Eocky  Mts.  by  Drummond,  is  doubtless  of  the  type  collection. 
By  his  var.  latifolia,  Besser  meant  the  typical  form  of  Purshianu  (as  both  species 
and  variety  seem  to  rest  on  the  same  type)  as  contrasted  with  his  var.  y3  angustifolia, 
also  described  here  and  based  on  a  plant  from  "Red  Eiver.  Douglas,  in  Herb.  Lindl.," 
which  is  said  to  have  lanceolate  leaves  1%  inches  long  and  3  lines  wide.  From  this 
description  it  appears  probable  that  this  variety,  too,  is  referable  here. 

A.  ludoviciana  var.  gnaphalodes  ToRR.  et  Gray,  Fl.  N.  Amer.  2  :420, 1843. 

A.  ludoviciana  var.  latifolia  ToRR.  et  Gray,  ibid. 

A.  vulgaris  var.  gnaphalodes  O.  Kuntze,  Rev.  1 :309,  1891. 

A.  vulgaris  var.  ludoviciana  O.  Kuntze,  ibid. 

A.  rhizomata  A.  Nels.,  Bull.  Torr.  Club  27:34,  1900.  A  narrow-leaved  form.  Type  locality: 
Sweetwater  River,  Wyoming. 


442  CALIFOBMA  ACADEMY  OF  SCIENCES  IProc.  4th  Ser. 

A.  rhizomaia  var.  pahularis  A.  Nels.,  ihid.  A  small  form  of  the  preceding.  Type  locality: 
Red  Desert,  near  Creston,  Wyoming. 

A.  diversifolia  Rydb.,  Bull.  Torr.  Club  28:20,  1901.  A  form  with  the  principal  leaves  merely 
pinnately  cleft,  but  the  few  lobes  very  narrow  and  acuminate.  It  closely  resembles 
forms  of  ssp.  candicans  from  the  general  region  of  northern  Idaho,  and  should  per- 
haps be  interpreted  as  a  transitional  form.  Type  locality:  Priest  River,  Idaho. 

A.  silvicola  Osterh.,  Bull.  Torr.  Club  28:645,  1901.  A  form  with  lanceolate,  nearly  entire, 
strongly  discolored  leaves  and  large  heads.  Type  locality :  Maclntyre  Creek,  Larimer 
Co.,  Colorado. 

A.  Briitonii  Rydb.,  Bull.  Torr.  Club  32:129,  1905.  Leaves  short,  with  3  to  5  short  rather 
divergent  lobes,  heavily  tomentose  on  both  sides.  Type  locality:  Golden,  Colorado. 

A.  pudica  Rydb.,  ibid.  130.  Inflorescence  loosely  branched,  the  large  heads  nodding,  leaves 
rather  thin  and  elongated.  A  spasmodic  variant  from  widely  scattered  stations  in  the 
Colorado  Rockies.  Perhaps  usually  a  shade  modification.  Type  locality:  Gunnison. 

A.  pabulari,s  Rydb.,  op.  clt.  33 :  157,  190(). 

A.  gnaphalodes  diversifolia  A.  Nels.  in  Coult  and  Nels.,  Man.  Rocky  Mt.  569, 1909. 

A.  mexicana  silvicola  A.  Nels.,  ibid. 

A.  H  err  iota  Rydb.,  Bull.  Torr.  Club  37:455,  1910.  A  large-leaved  form  from  Edmonton, 
Alberta. 

A.  cuneata  Rydb.,  N.  Amer.  Fl.  34:269,  1916.  Leaves  cuneate  obovate,  with  a  few  salient 
teeth  around  the  wide  apex.  Type  locality:  Idaho  Springs,  Colorado. 

A.  argophylla  Rydb.,  ibid.  274.  A  robust  form  with  exceptionally  large  heads.  The  type, 
from  Longs  Peak,  Colorado,  is  well  east  of  the  range  of  ssp.  candicans,  which  form 
it  resembles. 

A.  vulgaris  ssp.  ludoviciana  Hall  et  Clem.,  Carn.  Inst.  Wash.  Publ.  326  :  76, 1923. 

A.  vulgaris  ssp.  gnaphalodes  Hall  et  Clem.,  ibid.  77. 

A.  ludoviciana  var.  pabularis  Fern.,  Rhodora  47:248,  1945. 

A.  ludoviciana  var.  Briitonii  Fern.,  ibid. 

A.  ludoviciana  var.  cuneata  Fern.,  ibid. 

Stems  3-10  dm.  tall,  slender  to  moderately  stout,  herbaceous  to  base;  leaves 
many,  linear  to  lanceolate,  oblanceolate  or  elliptic,  sometimes  cuneate,  entire  or 
few-toothed  or  -lobed  especially  apically,  the  lobes  regularly  entire,  3-10  cm. 
long,  permanently  and  densely  white-tomentose  on  both  sides  or  loosely  floc- 
cose  to  green  and  glabrate  above,  plane  ;  inflorescence  an  elongated  usuallj' 
compact  panicle,  1.5-5  em.  broad  ;  heads  medium-sized,  erect  or  nodding,  often 
in  glomerules;  involucre  ovoid  to  campanulate,  3^  mm.  high,  2—3  mm.  wide, 
usually  densely  tomentose,  occasionally  glabrate,  with  7-13  bracts ;  ray-florets 
5-12  ;  di.sk-florets  6-21.  n  =  18. 

Lowlands  and  mountains  up  to  middle  elevations  (2200  m.,  or  occasionally 
up  to  2900  m.  in  the  Colorado  Rockies)  from  Ontario  to  Alberta,  Washington, 
California,  New  Mexico,  and  Arkansas ;  sporadically  introduced  from  Indiana 
to  Virginia,  New  England,  and  Quebec.  Flowering  from  July  through  Sep- 
tember. 

Ontario.  Thunder  Bay:  near  Fort  William.  Saskatchewan.  Assiniboia,  Indian  Head, 
Macoiin  10084 ;  Ribstone  Creek ;  Moose  Jaw,  Johnson  069.  ?  province,  plains  of  the  Saskatche- 
wan, Dnimmond  (Kew,  type(?)  of  A.  rurshinna  var.  hitifolia).  Alberta.  Stettler  District, 
Donalda  to  Bashaw;  Peace  River;  Grand  Prairie;  Edmonton,  Macovn  and  Ilcrriot  72825 
(NY,  type  of  A.  Ilerriotii)  ;  'RoseAa.\e,Moodic  1183 ;  Banff;  Lethbridge.  Bkitlsh  Columbl^. 
Kootenay:  Beavennonth,  Shaxv  1153;  Natural  Bridge,  TJlke.  Indiana.  Marshall,  Fulton, 


Vol.  XXV]       KECK:  BEFISIOX  OF  AETEMISIA  VULGARIS  COMPLEX  443 

Newton,  and  Sullivan  counties.  Illinois.  Dupage,  Ogle,  and  Eock  Island  counties.  WiscoN- 
.siN.  La  Crosse,  Sauk,  Dane,  and  Rock  counties.  Minnesota.  St.  Louis,  Norman,  Traverse, 
Winona,  Pope,  and  Hennepin  counties.  Iowa.  Emmet,  Fayette,  Linn,  Benton,  Poweschiek, 
Boone,  and  Decatur  counties.  Missouri.  Clark :  Mead.  Jackson :  Sibley.  Arkansas.  Craig- 
head: S.  of  Jonesboro,  Bemaree  7179.  North  Dakota.  Cass,  Eanson,  Stutsman,  Benson, 
Burleigh,  McLean,  and  Golden  Valley  counties.  South  Dakota.  Grant,  Spink,  Kingsbury, 
Harding,  Butte,  Meade,  Lawrence,  Custer,  and  Fall  River  counties.  Nebraska.  Douglas, 
Cass,  Lancaster,  Kearney,  Brown,  and  Cherry  counties.  Kansas.  Riley,  Saline,  McPherson, 
SedgA\dek,  and  Pratt  counties.  Oklahoma.  Woods,  Cimmaron,  Kingfisher,  and  Harmon 
counties.  Texas.  Wood:  Eeverchon  3961.  Montana.  Richland,  Glacier,  (Glacier  National 
Park),  Flathead,  Sanders,  Lewis  and  Clark,  Yellowstone,  Cascade,  Carbon,  Sweetgrass, 
Park,  Gallatin,  Jefferson,  Deer  Lodge,  Silver  Bow,  Ravalli,  and  Beaverhead  counties. 
Wyoming.  Crook,  Albany,  Sheridan,  Johnson,  Natrona,  Park  (Yellowstone  National  Park), 
Teton,  Lincoln,  Carbon,  Sweetwater,  and  Uinta  counties.  Colorado.  Sedwick :  Julesburg. 
Weld:  New  Windsor.  Larimer:  Mclntyre  Creek,  Aug.  24,  1900,  Osterhout  2242  (NY,  type 
of  A.  silvicola;  isotypes,  C,  RM,  US)  ;  Meadows  Inn,  Longs  Peak,  Aug.  14,  1907,  Clements 
(NY,  type  of  A.  argyrophylla).  Boulder:  Foot  of  Flagstaff  Mt.,  Ewan  in  PI.  Exsic.  Gray. 
897.  Routt:  S.  of  Steamboat  Springs.  Jefferson:  Golden,  Oct.  8,  1882,  N.  L.  Britton  (NY, 
type  of  A.  Brittonii).  Grande:  Hot  Sulphur  Springs.  Clear  Creek:  Idaho  Springs,  Shear 
4617  (NY,  type  of  A.  cuneata).  Rio  Blanco:  Buford.  Garfield:  Glenwood  Springs,  Eggle- 
ston  14982.  Otero:  La  Junta.  Fremont:  Canon  City,  Brandegee  501.  Huerfano:  Cucharas 
Valley.  Alamosa:  Alamosa.  Gunnison:  Gunnison,  Baker  573  (NY,  type  of  A.  pudica; 
isotypes,  C,  CAS,  Po,  RM,  US).  Mineral:  Pagosa  Peak,  Balcer  634.  San  Miquel:  Norwood 
Hill.  Montezuma:  Mesa  Verde  National  Park.  New  Mexico.  Colfax:  Raton.  Rio  Arriba: 
Brazos  Canyon ;  Dulce.  San  Juan  :  Tunitcha  Mts.  San  Miguel :  Las  Vegas.  Sandoval :  Sandia 
Mts.,  EUis  125.  Grant:  mts.  near  Condes  Camp,  Wrigld  1275.  Utah.  Rich:  Bear  Lake, 
Hammond  3879.  Box  Elder:  Corinne.  Weber:  Ogden  Canyon.  Salt  Lake:  Parleys  Park. 
Piute:  Marysvale,  Jones  5846.  San  Juan:  Dry  Wash,  Abajo  Mts.,  Ryherg  and  Garrett 
9615.  Iron:  Parowan.  Arizona.  Coconino:  Grand  Canyon,  Eggleston  15665.  Idaho.  Bonner: 
Slough  beach,  forest  between  Priest  Lake  and  East  Fork,  630  m.,  MacDougal  190  (NY, 
type  of  A.  diversifolia ;  Clarks  Fork  Valley  near  Horse  Plains,  Leiberg  1582.  Kootenai: 
Coeur  d'Alene,  Leiberg  1553.  Lemhi:  Gilmore.  Fremont:  Henry's  Lake,  Nelson  6796. 
Bannock:  Lava.  Bingham:  Blackfoot.  Blaine:  Ketchum.  Twin  Falls:  Shoshone  Falls, 
Nelson  and  Macbride  1366.  Gem:  Sweet,  Machride  1626.  Nevada.  Elko,  Eureka,  Lander, 
Humboldt,  White  Pine,  Nye,  Pershing,  Churchill,  Washoe,  Ormsl^y,  Storey,  and  Lyon 
counties.  Washington.  Pend  Oreille:  Kalispell  Lake.  Chelan:  Lake  Chelan.  Klickitat: 
Columbia  River,  Suksdorf  1610.  Oregon.  Wallowa,  Union,  Baker,  Grant,  Deschutes,  Crook, 
Malheur,  Harney,  Lake,  and  Klamath  counties.  California.  Modoc,  Nevada,  Mono,  Inyo, 
San  Bernardino,  and  Riverside  counties. 

Chromosome  counts  were  made  both  in  root-tips  and  in  meiosis  of  one  plant 
from  Manitou  and  of  two  plants  from  Engelmann  Canyon,  all  from  El  Paso 
County,  Colorado,  and  in  root-tips  only  of  one  plant  from  Leevining,  Mono 
County,  California,  and  of  two  plants  from  Upper  Fern  Valley,  San  Jacinto 
Mts.,  Riverside  County,  California. 

An  additional  very  interesting  tetraploid,  whose  chromosomes  were  counted 
in  root -tips,  is  now  referred  here.  It  was  collected  in  Poopenaut  Valley,  Tu- 
olumne River,  Yosemite  National  Park,  at  1065  m.  elevation,  Nohs  1  (CI, 
RM).  This  plant,  from  the  midst  of  the  area  occupied  by  the  hexaploid  A. 
Douglas'iana,  near  the  western  base  of  the  central  Sierra  Nevada,  was  pre- 
sumed to  be  a  peculiar  form  of  that  species  when  first  studied.  It  had  dis- 


444  CALIFORNIA  ACADEMY  OF  SCIENCES  [Proc.  4th  Ser. 

colored  leaves  like  Douglasiana,  but  these  were  extremely  narrow ;  those  on 
the  upper  half  of  the  stem  were  several  centimeters  long  but  less  than  5  mm. 
wide.  The  lower  leaves  were  oblanceolate  and  wider  (up  to  20  mm.  including 
lobes) ,  with  rather  short  forward-projecting  lobes  in  the  outer  half,  precluding 
the  possibility  that  this  could  be  a  form  of  A .  ludoviciana  ssp.  incompta  carried 
down  from  higher  elevations.  This  plant  is  best  explained  as  a  relict  of  ludo- 
viciana  ssp.  typica  stranded  on  the  western  flank  of  the  Sierra  Nevada  after 
most  of  its  fellows  had  been  replaced  by  the  younger  species,  Douglasiana. 
A  similar  plant  from  west  of  Dunlap,  Fresno  County,  Keck  1861  (CI,  SU), 
may  be  the  same.  This  suggests  that  ludoviciana  ssp.  tijpica  once  skirted  the 
southern  end  of  the  Sierra  Nevada  in  a  more  continuous  distribution. 

This  is  probably  the  most  variable  systematic  unit  in  the  whole  complex, 
as  the  long  synonymy  tends  to  show.  Extremes  can  be  pointed  out  that  appear 
very  unlike,  but  intermediates  link  all  together.  This  subspecies  intergrades 
not  uncommonly  with  ssp.  candicans  to  the  northwest,  with  ssp.  incompta  to 
the  west,  and  with  sspp.  alhtda  and  mexicana  to  the  southwest.  It  does  not 
overlap  the  ranges  of  sspp.  sulcata  and  redulens.  Apparently  to  a  much  lesser 
extent  it  intergrades  with  A.  serrata,  longifolia,  and  Carruthii. 

In  the  high  Rocky  Mountains  of  northern  Colorado  there  is  a  form  that 
seems  to  inter  grade  with  the  form  of  A.  Carruthii  known  as  A.  coloradensis 
Osterh.  Because  it  seems  impossible  to  decide  from  herbarium  material  whether 
this  is  best  assigned  to  A.  Carruthii  as  a  subspecies,  or  to  A.  ludoviciana  as  a 
subspecies,  the  question  is  left  open.  Perhaps  cytological  studies  will  be  ulti- 
mately decisive,  but  we  have  no  living  material  of  it.  The  following  specimens 
are  of  this  intergrading  form,  all  with  small  deeply  divided  leaves,  the  divisions 
varying  from  linear  to  lanceolate  :  Colorado.  Weld  :  Windsor,  Ostcrhout  5041 
(RM).  Larimer:  Pole  Hill,  Osterhout  2225  (RM);  Cherokee  Park,  Nelson 
10017  (C,  RM)  ;  Estes  Park,  Thompson  Canyon,  Johnston  6781)  (US). 
Boulder:  Boulder,  Hanson  C26  (US).  Denver:  Denver,  Jones  501c  (Po). 
Clear  Creek:  Empire,  Tweedy  5840  (NY,  RM);  Georgetown,  Shear  4514 
(RM)  ;  above  Silver  Dale  near  Georgetown,  July  10,  1878,  Jones  (Po). 

6b.  Artemisia  ludoviciana  Nutt.  ssp.  sulcata  (Rydb.)  Keck  comb.  nov. 

Figures  10  and  15 

Type  locality  :  Casas  Grandes,  Chihuahua. 
Artemisia  sulcata  Eydb.,  N.  Amer.  Tl.  34:270,  1916. 

Stems  3-12  dm.  tall,  smooth  or  tomentose,  sometimes  rather  prominently 
sulcate ;  leaves  lanceolate,  sharply  acute,  the  lower  tooth  apically  with  short 
salient  lobes,  the  upper  usually  entire,  reduced  in  the  inflorescence,  commonly 
thin  and  rather  flaccid,  mostly  3-10  cm.  long  and  less  than  1  cm.  wide  (exclu- 
sive of  lobes),  usually  bright  green  above,  densely  white-tomentose  beneath, 
.sometimes  ±  tomentose  above,  plane;  inflorescence  a  leafy  spreading  panicle 
or  sometimes  reduced  and  contracted;  heads  usually  distributed  in  individual 
nearly  naked  racemes  and  pendant;  involucre  campaiinlate,  2.5-3  mm.  high. 


Vol.  XXV]       KECK:  REVISION  OF  ABTEMISIA  VULGARIS  COMPLEX  445 


Figure  15.  Distribution  of  A.  ludoviciuna  sspp.  sulcata,  alhula,  mexicana,  and  redolens. 

2-3  mm.  wide,  usually  white-tomentose,  sometimes  quite  glabrate  (in  Chihua- 
hua, as  in  the  type),  with  10-16  bracts;  ray-florets  6-16;  clisk-florets  7-19. 
High  plateaus  and  mountains  from  central  Arizona  to  southern  Chihuahua, 
at  elevations  from  1600  to  2700  m.  Flowering  from  August  to  October. 

Arizona.  Gila :  Upper  Tonto  Creek,  Harrison  and  Kearney  8359.  Graham :  Mt.  Graham, 
9000  ft.,  Kearney  and  Peebles  9874.  Pinal:  Oracle.  Maricopa:  Sierra  Estrella,  Peebles  and 


446  CALIFORNIA  ACADEMY  OF  SCIENCES  [Proc.  4th  Ser. 

Kearney  7756.  Cochise:  between  Portal  and  Paradise;  Barfoot  Park,  Blnmer  1389 ;  War- 
ren; Mule  Mts.;  Bisbee;  Huachuca  Mts.,  Miller  Canyon,  Jones  25073,  Eamsey  Canyon, 
Shreve  5039.  Pima:  Santa  Catalina  Mts.;  Marshall  Gulch,  Shreve  5398;  Ft.  Lowell, 
Thornier  50,  toward  ssp.  mexicana;  Baboquivari  Canyon;  Rincon  Mts.,  Spud  Eanch,  Blumer 
3594.  Santa  Cruz:  Santa  Eita  Mts.,  Kearney  and  Peebles  10569 ;  Patagonia  Mts.  Sonora. 
Cajon  Bonito  Creek,  near  its  head,  Mearns  2474 ;  Cananea,  Donnelly  44 ;  La  Mesa  Colorado, 
Gentry  559.  Chihuahua.  Casas  Grandes,  August  30,  1899,  Townsend  and  Barber  437  (NY, 
type;  isotype,  US;  phototype,  C)  ;  Chuichupa,  Sierra  Madre,  Townsend  and  Barber  423 ; 
Colonia  Garcia,  Sierra  Madre,  Townsend  and  Barber  326;  Norogachi,  E.  Palmer  S.  S.; 
Sierra  de  Papas;  San  Jose  de  Pinal,  Rio  Mayo,  Gentry  2S43. 

In  proposing  Artemisia  sulcata,  R.ydberg-  emphasized  the  shining  glabrate 
involucres  and  the  grooved  stems.  As  pointed  out  by  Hall  and  Clements,  these 
do  not  furnish  good  criteria  for  separating  sulcata  from  ssp.  mexicana.  Other 
characters,  however,  are  more  dependable  for  separating  these  two  units, 
namely,  sulcata  has  entire  to  apieally  toothed  leaves  instead  of  pinnately  di- 
vided ones,  and  these  are  thin  and  plane  instead  of  thickish  and  revolute.  As 
shown  in  the  distribution  map,  figure  15,  the  range  of  ssp.  sulcata  is  adjacent 
to  that  of  ssp.  mexicana  which  it  largely  replaces  in  southern  Arizona. 

Artemisia  ludoviciana  ssp.  sulcata  is  wedged  between  sspp.  mexicana, 
alhula,  and  typica.  Intermediates  occur  between  sulcata  and  sspp.  alhula  and 
mexicana  along  the  periphery  of  its  range  suggesting  intercrossing  and  re- 
combination. Genetic  barriers  of  specific  importance  have  therefore  not  arisen 
between  these  units.  Possibly  the  broader  leaves  of  sulcata  point  to  a  connec- 
tion with  ssp.  typica.  It  appears  that  ssp.  sulcata,  is  a  recognizable  form  Avhich 
may  have  evolved  through  the  recombination  of  characters  from  the  three 
subspecies,  mexicana,  alhula,  and  typica,  which  surround  it. 

6c.  Artemisia  ludoviciana  Nutt.  ssp.  albula  (Woot.)  Keck  comb.  nov. 

Figures  10  and  15 

Type  locality  :  Organ  Mountains,  New  Mexico. 

Artemisia  microcephala  Woot.,  Bull.  Torr.  Club  25:455,  1898;  not  Hillebr.,  1888. 

A.  albula  Woot.,  Contr.  TJ.  S.  Nat.  Herb.  16  :  193, 1913.  New  name  for  A.  microcephala  Woot. 

Stems  3-10  dm.  tall ;  herbage  white-tomentose  throughout,  rarely  the  leaves 
evidently  discolored  and  merely  tomentulose  on  the  upper  surface ;  leaves 
mostly  obovate  to  elliptic,  their  teeth  or  lobes  projecting  forward  and  not 
nearly  cut  to  the  midrib,  the  upper  ones  reduced,  often  lance-linear  and  entire, 
not  fascicled,  thickish,  short  (mostly  1-2  cm.  long),  the  margin  often  nar- 
rowly revolute ;  inflorescence  open-paniculate ;  heads  small,  racemosely  ar- 
ranged along  the  elongated  divergent  remote  leafy  branchlets,  or  sometimes 
dr  glomerate;  involucre  campanulate,  ca.  3  mm.  high,  2  mm.  wide,  densely 
tomentose,  with  11-16  bracts;  ray-florets  8-11;  disk-florets  8-13. 

Western  Texas  and  southern  Colorado  to  southern  Nevada  and  California, 
south  to  northern  Baja  California,  Sonora,  and  Cliihuahua,  at  elevations 
from  150  to  2250  m.  Flowering  from  May  to  October. 


Vol.  XXV]       KECK:  REVISION  OF  ARTEMISIA  TVLGABIS  COMPLEX  447 

Texas.  Oldham:  Canadian  Valley.  Jeff  Davis:  Fort  Davis,  Eggleston  17412 ;  Sawtooth 
Mt.;  Limpia  P.O.  Colorado.  Boulder:  Eldorado  Springs,  Bartholomew  2  (NY).  El  Paso: 
Bear  Creek  Canyon.  Fremont :  Canon  City,  Biltmore  1437a.  Delta :  Paonia,  Eggleston  14954. 
New  Mexico.  Sandoval :  Sandia  Mts.,  Eastwood  15650.  Bernalillo :  Albuquerque.  Valencia : 
Laguna.  Catron:  Mangas.  Lincoln:  Gray,  Earle  457.  Sierra:  Hillsboro,  Metcalfe  1445. 
Grant:  Bear  Mt.,  Metcalfe  709.  Eddy:  Carlsbad  Caverns.  Dona  Ana:  Organ  Mts.,  4500  ft., 
Sept.  1,  1897,  Wooton  504  (US,  type;  phototypes,  C,  NY,  US;  isotypes,  NY,  US).  Luna: 
Florida  Mts.  Hidalgo:  Lordsburg;  Dog  Spring,  Mearns  2370.  Utah.  San  Juan:  Butler 
Wash;  Bluff.  Washington:  Zion  National  Park;  St.  George.  Nevada.  Lincoln:  Panaea. 
Clark:  Charleston  Mts.,  Coville  and  Fvnston  372.  Arizona.  Apache:  Petrified  Forest  N.  of 
Adamana.  Hall  11171.  Navajo:  Shonto.  Coconino:  Grand  Canyon,  Eastwood  and  Hoivell 
7006.  Mohave:  White  Cliffs  E.  of  Kingman.  Yavapai:  Weaver  Mts.,  Gillespie  8498.  Gila: 
Payson ;  Globe.  Maricopa :  Fish  Creek.  Pinal :  Superstition  Mt.,  Gillespie  8585.  Greenlee : 
Blue  Eiver.  Graham:  Ft.  Grant.  Cochise:  Paradise,  Blumer  1798;  Dragoon  Pass,  Jones 
28623;  Bisbee.  Pima:  Tucson;  Baboquivari  Canyon.  California.  Tulare:  Kern  Eiver  at 
Funston  Creek,  6900  ft.,  Sharsmith  3840.  San  Bernardino :  Clark  Mt. ;  5  mi.  S.  of  Barnwell, 
Munz  13838;  Providence  Mts.;  Little  San  Bernardino  Mts.  Eiverside:  Tahquitz  Canyon, 
Palm  Springs,  Wheeler  2296;  San  Jacinto  Eiver  Canyon.  San  Diego:  Borrego  Valley. 
Chihuahua.  Candelaria,  Stearns  276;  Sierra  en  Media,  E.  W.  Nelson  6488 ;  Santa  Eulalia 
Mts.,  Pringle  626.  Sonora.  San  Jose  Mts.,  Mearns  1055.  Baja  California.  Cantillas  Mts., 
1875,  E.  Palmer;  between  Ojos  Negros  and  Neji  Eancho,  Wiggins  and  Gillespie  4146;  La 
Encantada,  Sierra  San  Pedro  Martir,  Wiggins  and  Demaree  4877,  4999  (Po,  SU,  both 
atypical,  toward  ssp.  sulcata  or  mexicana) . 

Hall  and  Clements  included  this  form  under  A.  viilgaris  ssp.  gnaphalodes, 
although  they  accurately  pointed  out  its  outstanding  characteristics.  Blake, 
in  Kearney  and  Peebles'  Flowering  Plants  and  Ferns  of  Arizona,  likewise 
refers  it  to  A.  gnaphalodes.  It  will  be  seen  from  the  distribution  map  that  as 
here  circumscribed  A.  ludoviciana  ssp.  typica  (which  includes  A.  gnapha- 
lodes) and  A.  ludoviciana  ssp.  aUnla  largely  occupy  different  geographical 
areas.  Their  different  morphological  patterns  mark  them  as  excellent  examples 
of  geographical  subspecies. 

6d.  Artemisia  ludoviciana  Nutt.  ssp.  candicans  (Rydb.)  Keck  comb.  nov. 

Figures  10  and  16 
Type  locality  :  Little  Belt  Mountains,  Montana. 

Artemisia  ludoviciana  var.  latiloba  Nutt.,  Trans.  Amer.  Phil.  Soc.  II,  7:400,  1841.  "With 
the  above  [A.  ludoviciana],  in  the  Eocky  Mountains."  According  to  Eydberg,  the 
same  as  his  A.  platyphylla,  but  the  entity  uncertain,  with  no  type  found. 

A.  candicans  Eydb.,  Bull.  Torr.  Club  24:296,  1897. 

A.  floccosa  Eydb.,  ihid.  297.  In  the  type  of  candicans  the  heads  are  sessile  along  the  branch- 
lets  of  the  inflorescence  and  thus  rather  glomerate ;  in  the  type  of  floccosa  the  heads 
are  short-pediceled  and  arranged  in  a  long  spike-like  panicle.  In  other  respects  the 
two  forms  are  closely  similar.  Type  locality:  Lima,  Montana. 

A.  paucicephala  A.  Nels.,  Bull.  Torr.  Club  27:35,  Jan.  24,  1900.  A  form  with  many  of  the 
leaves  entire  and  thus  allied  to  ssp.  typica  ;  however,  some  leaves  are  deeply  parted, 
and  the  very  large  heads  ally  it  more  closely  to  this  than  to  typica.  Type  locality : 
near  Yellowstone  Lake  on  the  banks  of  a  tributary  creek. 

A.  gracilenta  A.  Nels.,  ibid.  A  foliage  form  of  the  preceding,  with  the  leaves  twice  cut  and 
the  lobes  more  slender,  in  this  respect  comparable  with  the  Flodmanii  form  of  ssp. 
incompta.  Type  locality:  sandy  beaches  and  banks  of  Yellowstone  Lake. 


448 


CALIFORNIA  ACADEMY  OF  SCIENCES 


[Proc.  4th  «er. 


Figure  l(i.  Distribution  of  A.  ludoviciana  sspp.  candicans  and  incompta. 

A.  latiloba  Eydb.,  Mem.  N.  Y.  Bot.  Gard.  1:429,  Feb.  15, 1900, 

A.  platyphylla  Rydb.,  N.  Amor.  Fl.  34:275,  1916.  Leaves  of  the  type  large  and  cut  much  like 
those  of  A.  Douglasiana,  that  is,  cuneate-obovate  and  toothed  or  short-Iobed  toward 
apex,  but  the  inflorescence  and  pubescence  are  like  candicans.  Furthermore,  a  large 
series  of  variations  from  the  Spokane  region  in  the  Herbarium  of  the  University  of 
California  make  the  complete  transition  to  candicans.  Ssp.  lypica  is  not  only  very  rare 
in  Washington,  but  has  much  smaller  heads  than  this.  Although  chromosome  counts 


Vol.  XXV]       KECK:  EEVISION  OF  AETEMISIA.  VULGARIS  COMPLEX  449 

are  lacking  which  would  clineli  the  matter,  we  suspect  A.  Douglasiana  may  occur  in 
this  region,  but  in  an  atypically  gray-pubescent  form.  The  type  locality  of  A. 
platyphylla  is  sandy  or  gravelly  banks  of  Spokane  River,  Spokane,  Washington. 

A.  vulgaris  ssp.  candicans  Hall  et  Clem.,  Carn.  Inst.  Wash.  Publ.  326:73,  1923. 

A.  vulgaris  var.  candicans  F^CK,  Man.  Higher  PI.  Ore.  766,  1941. 

Stems  3-10  dm.  tall,  entirely  herbaceous ;  herbage  white-tomentose  through- 
out ;  leaves  oblanceolate  to  broadly  elliptic  in  outline,  deeply  pinnately  cleft 
or  parted  into  forward-projecting  narrow  lobes,  at  least  some  lobes  again 
toothed,  or  sometimes  the  leaves  merely  pinnately  lobed  and  the  upper  ones 
entire.  5-10  cm.  long,  1.5-4  cm.  wide,  sometimes  more  thinly  tomentose  on 
upper  surface,  but  rarely  ever  green,  plane  or  nearly  so ;  inflorescence  a  some- 
what leafy  narrow  panicle,  the  heads  borne  singly  or  in  small  glomerules, 
large,  sometimes  nodding;  involucre  broadly  campanulate  to  hemispheric, 
3.5-4  mm.  high,  3-7  mm.  wide,  tomentose,  with  9-14  bracts;  ray-florets  7-12; 
disk-florets  17-42. 

Lowlands  and  mountains  up  to  middle  elevations  (2400  m.)  from  western 
Montana  and  Wyoming  to  northern  Utah  and  westward  to  central  Washing- 
ton, Oregon,  and  northern  California.  Flowering  from  June  to  September. 

Montana.  Glacier  National  Park:  Baring  Basin,  Standley  17159.  Flathead:  Summit; 
Belton.  Judith  Basin :  Little  Belt  Mts.,  6  mi.  from  Barker,  6000  ft.,  Aug.  18,  1896,  J.  H. 
Flodman  882  (NY,  type  of  A.  candicans;  isotype,  US).  Gallatin:  Sedan.  Beaverhead:  Lima, 
Aug.  6,  1895,  Bydberg  2942  (NY,  type  of  A.  floccosa;  isotype,  US).  Park:  Gardiner, 
Hawkins  72Sa.  Wyoming.  Yellowstone  National  Park:  Yellowstone  Lake,  Aug.  22,  1899, 
A.  and  E.  Nelson  6612  (RM,  type  of  A.  gracilenta ;'isotj])es,  NY,  RM),  ditto,  Aug.  6,  1899, 
A.  and  E.  Nelson  6344  (RM,  type  of  A.  paucicephala ;  isotypes,  NY,  Po,  RM,  SU).  Teton: 
Teton  Forest  Reservation,  Brandegee.  Sweetwater:  6  mi.  N.  of  Rock  Springs.  Uinta:  West 
Fork  of  Bear  River,  Payson  4840.  Utah.  Cache:  Summer  Camp,  Logan  Canyon,  Art  Smith. 
Idaho.  Bonner:  Lake  Pend  d'Oreille.  Kootenai:  Farmingtoh  Landing,  Sandherg  et  al.  569 ; 
Couer  d'Alene,  Leiberg  1552.  Latah:  Comerine  Gulch.  Lemhi:  Gilmore.  Fremont:  St.  An- 
thony. Blaine:  Hailey.  Custer:  Alturas  Inlet.  Washington:  Tamarack,  Clarlc  22?.  Owyhee: 
Twilight  Gulch,  Macbride  486.  Nevada.  Elko:  Little  Lakes  Canyon.  White  Pine:  Duck 
Creek,  Ely,  Mitchcoch  1390.  Humboldt:  Winnemucca,  Griffiths  and  Morris  40.  Washington. 
Spokane:  banks  of  Spokane  River,  Spokane,  June  1897,  Elmer  867  (NY,  type  of  A.  platy- 
phylla; isotypes,  Po,  US),  ditto,  Oct.  1921,  Moore  (C,  a  large  series  of  variations).  Whit- 
man: S.E.  of  Lamont;  Wawawai,  Keclc  382,  Columbia:  Blue  Mts.  Adams:  Sheep  Springs, 
Leiberg  944.  Grant:  Soap  Lake,  Thompson  11795.  Chelan:  Mt.  Stuart.  Kittitas:  Ellensburg. 
Yakima :  Upper  Naches  River.  Oregon.  Wallowa :  Joseph ;  Wallowa  Lake.  Union :  La 
Grande.  Umatilla :  Toll  Gate.  Hood  River :  Hood  River.  Grant :  Izee.  Crook :  Prineville. 
Lake:  Silver  Lake,  Leiberg  758.  Klamath:  Pelican  Bay.  California.  Modoc:  Lily  Lake, 
Pine  Creek.  Nevada :  ridge  S.  of  Donner  Pass,  8500  ft.,  Heller  7153. 

This  subspecies  occurs  through  approximatel,y  the  same  geographical  region 
as  ssp.  incompta,  but  for  the  most  part  at  lower  elevations  and  in  a  different 
life  zone.  Transitional  specimens  are  not  infrequently  found  between  these 
two  subspecies.  Likewise,  transitions  are  sometimes  found  between  ssp.  can- 
dicans and  ssp.  typica.  As  mentioned  under  the  synonym  A.  platyphylla,  forms 
from  Washington  are  sometimes  difficult  to  separate  from  A.  Douglasiana, 
although  presumably  the  two  have  a  genetic  barrier  to  their  interbreeding  in 


450  CALIFORNIA  ACADEMY  OF  SCIENCES  [Proc.  4th  Ser, 

a  difference  in  chromosome  number.  It  is  quite  possible  that  the  Douglasiana- 
like  forms  of  candicans  have  arisen  independently  as  a  parallel  variation  and 
are  not  the  result  of  Douglasiana  genes  picked  up  through  crossing.  This 
possibility  gains  credence  from  the  complete  series  of  intermediates  linking 
these  broad-leaved  forms  with  the  narrow-leaved  forms  of  candicans. 

6e.  Artemisia  ludoviciana  Nutt.  ssp.  incompta  (Nutt.)  Keck,  Carn.  Inst. 

Wash.  Publ.  520  :327, 1940. 
Figures  8,  10,  and  16 

Type  statement :  "In  the  central  chain  of  the  Kocky  Mountains,  in  Thorn- 
berg's  Pass,  near  the  great  passage  to  the  plains  of  the  Oregon."  This  locality 
is  in  Custer  County,  Idaho,  and  the  type  was  collected  by  Nuttall. 

Artemisia  incompta  Nutt.,  Trans.  Amer.  Phil.  Soc.  II,  7:400,  1841. 

A.  discolor  var.  incompta  Gray,  Syn.  PI.  1(2)  :373,  1884, 

A.  atomifera  Piper,  Contr.  U.S.  Nat.  Herb.  11:588,  1906.  A  form  from  Wawawai,  Wash- 
ington, on  the  Snake  Eiver,  ^vith  thick,  strongly  discolored  leaves  that  are  green  on 
the  upper  surface,  and  some  of  which  are  bipinnately  lobed.  An  unusual  variant  from 
within  the  zone  of  candicans,  but  from  its  morphology  best  assigned  to  incompta. 
Wide  variation  is  exhibited  at  the  type  locality,  so  it  is  impossible  to  distinguish  a 
separate  systematic  unit  here. 

A.  ludoviciana  var.  atomifera  M.  E.  Jones,  Bull.  Univ.  Mont.  Biol.  ser.  15  :48,  1910. 

A.  potens  A.  Nels.,  Bot.  Gaz.  54:418,  1912.  A  greenish  form  very  similar  to  the  type  of 
incompta  taken  not  far  from  the  type  locality  of  this  subspecies.  Type  locality: 
Mackay  (Pass  Creek),  Custer  Co.,  Idaho. 

A.  Flodmanii  Rydb.,  N.  Amer.  FI.  34:276,  1916.  A  densely  tomentose  form  with  linear  divi- 
sions to  the  leaves;  heads  medium-sized,  too  small  for  candicans.  An  uncommon 
form,  Init  principally  found  in  western  Montana.  Type  locality:  Little  Belt  Moun- 
tains, Montana,  9  miles  east  of  Barker. 

A.  vulgaris  ssp.  Flodmanii  Hall  et  Clem.,  Carn.  Inst.  Wash.  Publ.  326:75,  1923. 

A.  vulgaris  ssp.  Micltauxiana  var.  incompta  St.  John,  Ees.  Stud.  St.  Coll.  Wash.  1 :  106, 1929. 

A.  vulgaris  var.  Flodmanii  Peck,  Man.  Higher  Pis.  Ore.  767,  1941. 

Stems  3-9  dm.  tall,  usually  entirely  herbaceous ;  herbage  commonl;\'  ±  green, 
with  leaves  glabrate  above  and  white-tomentose  beneath,  but  often  ±  densely 
tomentose  throughout,  and  sometimes  essentially  glabrous  throughout ;  lower 
leaves  mostly  obovate  to  broadly  elliptic  in  outline,  rarely  fascicled,  parted 
or  divided  into  linear  or  lanceolate  often  elongated  forward-projecting  lobes, 
some  of  these  again  toothed  or  lobed,  2-8  cm.  long,  the  upper  leaves  less  cut  or 
even  entire,  the  margin  rarely  revolute ;  inflorescence  a  narrow  spike-like  or 
even  racemose  panicle,  sometimes  much  broader  but  not  open,  not  verj^  leafy ; 
heads  rather  large,  nodding  at  first  or  erect;  involucre  broadly  campanulate, 
3-3.5  mm.  high,  2.5-4  mm.  wide,  sericeous-tomentose  to  glabrate  and  shining, 
with  9-14  bracts ;  ray-florets  6-10;  disk-florets  15-30  (-45).  n  =  18. 

Mountains  of  Montana  and  Idaho,  southward  to  central  Utah,  Nevada  and 
California,  at  elevations  from  1400  to  3000  (to  3500)  m.  Flowering  from  July 
through  September. 


Vol.  XXV]       KECK:  REVISION  OF  ARTEMISIA  VULGARIS  COMPLEX  451 

Montana.  Glacier:  Swiftcurrent  Creek,  Standley  16071.  Cascade:  Belt  Creek.  Judith 
Basin:  Little  Belt  Mts.,  9  mi.  from  Barker,  Aug.  18,  1896,  6,000  ft.,  Flodman  883  (NY, 
type  of  A.  Flodnianii ;  isotype,  US).  Sweet  Grass:  Sweet  Grass  Canyon,  Crazy  Mts.  Lewis 
and  Clark :  Gate  of  the  Mountains.  Jefferson :  Boulder  Creek.  Silver  Bow :  Durant.  Gallatin : 
Bridger  Mts.,  Aug.  21, 1902,  W.  W.  Jones.  Park :  Emigrant  Gulch,  Rydherg  and  Bessey  5206. 
Beaverhead:  Lima,  Shear  5062.  Wyoming.  Yellowstone  National  Park:  Madison  Eiver. 
Teton:  Government  Bridge;  Jackson  Hole,  Payson  2264.  Lincoln:  Salt  Eiver  Mts.  E.  of 
Smoot.  Carbon:  Bridger  Peak.  Colorado.  Routt:  Anita  Peak,  Goodding  1753.  Eagle:  Eed 
Cliff.  Utah.  Cache:  Tony  Lake  basin,  Maguire  16105.  Salt  Lake:  Alta.  Utah:  Mt.  Tim- 
panogas.  Juab:  Mt.  Nebo.  San  Pete:  Eeeder  Canyon.  Sevier:  Fish  Lake  (C)  ;  head  of 
Bullion  Creek,  11,500  ft.,  Jones  5873.  Idaho.  Lemhi:  Gilmore,  Rail  11552.  Custer:  Garden 
Creek,  near  Challis,  Macbride  and  Payson  3350;  Mackay,  Pass  Creek,  6,700  ft.,  July  30, 
1911,  Nelson  and  Machride  1413  (EM,  type  of  A.  poiens;  isotypes,  C,  NY,  Po,  SU,  US)  ; 
Mackay,  Bear  Canyon,  Nelson  and  Macbride  1521;  Thornberg's  Pass,  Nuttall  (Ph,  type  of 
A.  incompta).  Blaine:  Alturas  Lake;  Ketchum.  Owyhee:  Twilight  Gulch,  Macbride  485. 
Nevada.  Elko  :  Coon  Creek,  Jarbidge  Mts.  (SU)  ;  Spruce  Mt. ;  Lamoille  Canyon.  White  Pine. 
Sherman  Peak,  Euby  Mts.,  Hitchcock  and  Martin  5651.  Pershing:  Star  Peak,  Watson  638. 
Lander:  35  mi.  S.W.  of  Austin;  W.  of  Bunkerhill  Mt.  Humboldt:  Winnemucca;  Humboldt 
Canyon,  Heller  10625.  Washoe:  Galena  Creek;  Mt.  Eose.  Mineral:  Walker  Lake,  Tidestrom 
10157.  Clark:  Eainbow  Falls,  Charleston  Mts.,  CloTcey  7388.  Washington.  Whitman: 
Wawawai,  Sept.  1903,  C.  V.  Piper  6466  (US,  type  of  A.  atomifera;  isotypes,  CAS,  NY, 
SU,  US).  Oregon.  Wallowa  Lake;  Bear  Creek,  Sheldon  8809 ;  West  Fork,  Wallowa  Eiver, 
Keclc  366.  Grant :  Strawberry  Mt.  Harney :  base  of  Steins  Mts.  California.  Modoc :  Cedar 
Mt.  Lassen:  Janesville.  Plumas:  Prattville.  Nevada:  Soda  Springs.  Placer:  Summit, 
Heller  12898.  Alpine:  Ebbet  Pass;  Sonora  Pass.  Mono:  Blue  Lake;  Leevining  Grade, 
Keclc  5015;  McAfee  Meadows,  White  Mts.,  11,500  ft.,  Duran  2834  (C,  SU,  with  small 
finely  cut  leaves  like  the  tenuis  form  of  A.  Michanxiana).  Tuolumne:  Upper  Eelief  Valley; 
Tuolumne  Meadows.  Mariposa:  Washburn  Lake.  Madera:  Merced  Pass,  Bolander  6152. 
Inyo:  Piute  Pass-North  Lake  Trail,  Ferris  8937 ;  Mt.  Baxter.  Tulare:  Kaweah  Peaks; 
trail  to  Mt".  Whitney,  Culbertson  in  Baker  4341 ;  Little  Kern ;  Junction  Meadows.  Los 
Angeles:  Swartout  Valley,  Munz  7711;  Pine  Mt.  ridge,  Johnston  1651.  San  Bernardino: 
Vivian  Creek;  Big  Meadows  of  Santa  Ana;  Dry  Lake,  Mt.  San  Gorgonio.  Eiverside:  Straw- 
berry Valley,  San  Jacinto  Mts.,  Wheeler  171. 

Chromosomes  were  counted  in  meiotic  divisions  of  two  plants  from  Yosemite 
Creek,  Tioga  Road,  Yosemite  National  Park,  at  2200  m.  elevation,  and  in 
somatic  divisions  in  root-tips  of  a  plant  from  White  Mountain,  Yosemite  Na- 
tional Park,  at  3200  m.,  and  of  another  from  Slate  Creek  Valley,  Mono  County, 
California,  at  3260  m.  elevation.  An  additional  plant  from  Slate  Creek  Valley 
proved  to  be  hexaploid,  with  2n  =  ca.  54.  (See  Clausen,  Keck,  and  Hiesey, 
1940a,  p.  336,  for  additional  information  on  this  plant.) 

As  is  the  case  with  most  of  the  other  units  in  this  complex,  ssp.  incompta  is 
extremely  variable,  and  many  collections  included  here  neither  fully  fit  the 
description  nor  bear  a  close  resemblance  to  even  neighboring  collections.  The 
closest  resemblance  of  ssp.  incompta  within  the  species  is  to  typica  and  candi- 
cans.  Intergrades  are  not  rare.  As  shown  in  the  map,  figure  16,  ssp.  incompta 
ranges  farther  south  than  ssp.  candicans.  The  exact  boundaries  cannot  be 
automatically  determined.  For  instance,  on  the  higher  summits  of  the  Wasatch 
Range  in  San  Pete  County,  Utah,  far  south  of  the  range  of  typical  candicans, 
four  collections  have  been  studied  all  of  which  bear  a  morphological  resem- 


452  CALIFORNIA  ACADEMY  OF  SCIENCES  [Proc.  4th  Ser. 

blance  to  candicans,  but  which  might  be  classed  as  intermediates.  From  their 
occurrence  well  within  the  region  of  incompta  and  their  transitional  appear- 
ance, these  are  considered  an  example  of  parallel  variation  within  incompta 
rather  than  representatives  of  candicans  of  disjunct  distribution. 

The  relationships  of  ssp.  incompta  outside  the  species  are  with  A.  Michauxi- 
ana  and  Liiidleyana.  This  is  suggested,  also,  in  figure  19,  page  466.  The 
breaks  between  these  units,  however,  seem  to  be  more  clear  cut  than  those 
between  the  subspecies  of  ludoviciana,  and  this  helps  justify  the  maintenance 
of  species  instead  of  subspecies.  A  thorough  cytological  survey  should  precede 
drastic  taxonomic  shifts  in  this  group. 

6f.  Artemisia  ludoviciana  Nutt.  subsp.  mexicana  (Willd.)  Keck  comb.  nov. 

Fig-ures  10  and  15 
Type  locality  :  Mexico.  Type  not  seen. 

Artemisia  mexicana  Willd.  in  Spreng.,  Syst.  3  :490,  1826. 

Oligosporus  mexicanus  Less.,  Syn.  Gen.  Compos.  264,  1832. 

Artemisia  indica  mexicana  Bess.,  Nouv.  Mem.  Soc.  Nat.  Mosc.  3  :56,  1834. 

A.  vulgaris  var.  mexicana  ToRR.  et  Gray,  F1.  N.  Amer.  2 :421, 1843. 

A.  cuneifolia  Scheele,  Linnaea  22:162,  1849.  Type  not  seen,  but  according  to  Hall  and 
Clements  this  is  a  tall,  very  erect  and  rigid,  small-headed  form  of  the  subspecies  here 
considered.  Type  locality :  high  places  on  the  prairie  near  New  Braunf els,  Texas. 

A.  Lindheimsriana  Scheele,  ibid.  163.  Type  not  seen,  but  according  to  Hall  and  Clements 
stems  rigid  and  almost  woody;  heads  only  2  mm.  across.  Type  locality:  in  the  dry 
river  bed  of  the  Cibolo,  fifteen  miles  west  of  New  Braunfels,  Texas. 

A.Underwoodii  Rydb.,  Bull.  Torr.  Club  32:129,  1905.  A  form  with  the  deeply  lobed, 
strongly  discolored  foliage  of  mexicana  and  the  compact  spike-like  panicle  of  typica, 
to  which  it  is  rather  intermediate.  Type  locality:  Ouray,  Colorado. 

A.  revoluta  Rydb.,  N.  Amer.  Fl.  34:272,  1916.  Form  with  very  narrow,  elongated,  revolute 
divisions  of  the  leaves  and  a  narrow  inflorescence.  Type  locality :  city  of  Chihuahua. 

A.  texana  Rydb.,  ihid.  274.  Intermediate  to  typica,  with  broad  leaves,  the  upper  of  which 
are  entire.  Judging  from  its  distribution  and  the  presence  of  intergrades,  a  tomentose 
form  of  mexicana.  Type  locality:  Colorado,  Texas. 

A.  Ghieshreghiii  Rydb.,  ibid.  271.  An  exceptionally  large-headed  form  of  mexicana,  but 
lacking  the  type  of  mexicana,  and  finding  head  size  a  varialile  character  in  Mexico, 
it  seems  best  to  put  this  into  synonymy  here.  Type  locality:  Chiapas. 

A.  Muelleri  Rydb.,  ibid.  270.  A  form  with  rather  wide  mostly  entire  upper  leaves.  Type 
locality:  Orizaba. 

A.  neomexicana  Greene  ex  Rydb.,  ibid.  279.  A  narrow-leaved  form  approaching  A.  Carrutjiii, 
strongly  discolored;  inflorescence  rather  narrow  for  llio  subspecies.  Type  locality: 
in  a  grassy  glade,  Iliilsboro  Peak,  Black  Range,  New  Mexico. 

A.  vulgaris  ssp.  mexicana  Hall  et  Clem.,  Carn.  Inst.  Wash.  Publ.  326:80,  1923. 

A.  ludoviciana  Lindheimeriana  Bush,  Amer.  Midi.  Nat.  11:35,  1928. 

A.  ludoviciana  var.  mexicana  Fern.,  Rhodora  47:248,  1945. 

Stems  3-10  dm.  tall,  densely  leafy,  herbaceous  to  base ;  leaves  variable,  typi- 
cally 5-10  cm.  long,  with  few  linear  pinnate  spreading  entire  divisions  2—1  mm. 
wide,  but  tiie  upper  and  many  of  the  cauline  often  entire  or  nearly  so,  and 
the  lower  cauline  often  merely  ternately  divided,  loosely  floccose  to  glabrous 
and  <liirk  gi-een  above,  densely  white-tomentose  beneath,  often  thickish,  nar- 


i 


Vol.  XXVJ       KECK :  SEVISION  OF  ARTEMISIA  VULGARIS  COMPLEX  453 

rowly  revolute;  inflorescence  a  rather  dense  leafy  branching  panicle  3-10 
cm.  broad ;  heads  rather  small,  erect  or  more  commonly  nodding ;  involucre 
turbinate  to  campanulate  or  hemispheric,  2.5-3.5  mm.  high,  2-3  mm.  wide, 
sparingly  to  densety  tomentose,  with  10-15  bracts;  ray-florets  5-12;  disk- 
florets,  6-18. 

Western  INIissouri  to  Colorado,  Arizona,  and  southward  to  Guatemala,  at 
elevations  from  250  to  3050  m.  Flowering  from  August  through  October.  Type 
locality :  Mexico. 

Missouri.  Jackson :  Courtney,  Bush  6509.  Greene :  Springfield,  Standley  8637.  Arkansas. 
Pulaski:  Little  Bock,  Sept.  1886,  Hasse  (SU).  Kansas.  Cloud:  Aurora,  Eraser  553.  Sedg- 
wick: Wichita.  Oklahoma.  Creek,  Payne,  Cleveland,  Caddo,  and  Kiowa  counties.  Texas. 
Fannin,  Dallas,  Tarrant,  Parker,  Young,  Dickens,  Lubbock,  Garza,  Midland,  Tom  Green, 
Jeff  Davis,  Travis,  Kendall,  Comal,  Bexar,  Valverde,  Maverick,  Webb,  and  Duval  counties. 
Mitchell:  Colorado,  Tracy  8141  (NY,  type  of  A.  texana;  isotypes,  NY,  US).  Colorado. 
Larimer:  Moraine  Park,  OsierJioui  3130  (RM,  atypical).  Summit:  Breckenridge,  MacTcensie 
S37  (NY,  atypical).  Garfield:  Glenwood  Springs,  Osterhout  3309  (EM,  atypical).  Ouray: 
chaparral  covered  hills  S.E.  of  Ouray,  Underwood  and  Selby  74  (NY,  type  of  A.  Under- 
woodii;  phototype,  C).  Montrose:  Naturita,  Payson  590.  San  Miguel:  Telluride,  10,000  ft.. 
Tweedy  332.  New  Mexico.  Colfax,  Rio  Arriba,  San  Juan,  Sandoval,  Lincoln,  Otero,  Catron, 
and  Sierra  counties.  Grant:  Hillsboro  Peak,  Black  Range,  Metcalfe  1248  (NY,  type  of  A. 
neomexicana;  phototype,  C;  isotypes,  CAS,  Po,  RM,  US).  Arizona.  Apache:  Carrizo  Mts.; 
Hannagan  Meadow,  White  Mts.,  1935,  McNeill.  Navajo:  Kayenta;  Holbrook.  Coconino: 
Grand  View,  Grand  Canyon,  Eastwood  3611  (CAS,  toward  albula)  ;  S.  of  Flagstaff.  Sonora. 
Halfway  between  Cumpas  and  Baviacora,  Drouet  et  al.  3654.  Chihuahua.  Boundary,  near 
White  Water,  Mearns  2281;  Soldier  Canyon,  Sierra  Madre;  Colonia  Garcia,  Toivnsend  and 
Barber  296;  by  streams  near  Chihuahua,  Pringle  290  (NY,  type  of  A.  revoluta;  phototype, 
C;  isotypes,  NY,  Ph)  ;  between  Porral  and  San  Julian,  E.  W.  Nelson  4939  (US,  toward 
recJolens).  Coahuila.  Soledad,  E.  Palmer  736  (toward  albida)  ;  Sierra  de  Parras,  Purpus 
4659  (toward  Carruthii) ;  Saltillo,  E.  Palmer  286  (possibly  a  distinct  subspecies,  with  broad, 
venulose,  involute-margined,  tomentose  leaves  like  some  Texan  forms  but  showing  some 
characters  of  typica).  NuEvo  Leon.  Galeana,  Taylor  207.  Durango.  Durango,  E.  Palmer 
600,  601,  602,  907.  San  Luis  Potosi.  Alvarez,  E.  Palmer  59;  San  Luis  Potosi,  Parry  and 
Palmer,  530,  531.  Aguascalientes.  Aguascalientes,  Rose  and  Painter  7760.  Jalisco.  Rio 
Blanco;  Guadalajara,  Pringle  8765.  Hidalgo.  Dublan;  Pachuca;  Tula,  Pringle  9848. 
Mexico.  Contadero;  Eslava.  Federal  District.  Mexico  City.  Michoacan.  Morelia.  Vera 
Cruz.  Orizaba,  Fred  Miiller  235  (NY,  type  and  isotype  of  A.  Muelleri)  Puebla.  Puebla. 
Chiapas.  Without  exact  locality,  Ghiesbrecht  155  (NY,  type  of  A.  Ghiesbreghtii,  the  spelling 
inadvertently  (?)  altered  by  Rydberg).  Guatemalia.  San  Pedro  S.,  Tejada  49  (US).  Ap- 
parently introduced  and  cultivated  in  San  Salvador,  Calderon  2255  (US),  where  the  collector 
reports  it  to  be  "much  used  by  native  people  as  remedy  for  various  affections." 

At  least  as  variable  a  subspecies  as  typica,  but  difficult  to  subdivide  further 
on  the  basis  of  present  collections.  Undoubtedly  distinct  ecotypes  remain  to  be 
worked  out  in  the  Mexican  highlands.  A  few  forms,  now  puzzling,  await  a  fuller 
understanding  before  adequate  treatment.  For  example,  a  collection  from  San 
Pablo,  Baja  California,  collected  April  21, 1889,  by  T.  S.  Brandegee  (C) ,  from 
well  outside  the  range  of  this  otherwise  fall  flowering  subspecies,  is  best  re- 
ferred here  temporarily  on  the  basis  of  form.  Intergrades  link  ssp.  mexicana 
with  ssp.  typica,  albula,  and  reddens. 


454  CALIFORNIA  ACADEMY  OF  SCIENCES    '  [Proc.  4th  Ser. 

6g.  Artemisia  ludoviciana  Niitt.  ssp.  redolens  (Gray)  Keck  comb.  nov. 

Figures  10  and  15 

Type  statement :  "Chihuahua,  on  cool  slopes  under  cliffs,  Pringle,  296." 

Artemisia  redolens  Gray,  Proc.  Amer.  Acad.  21 :  393,  1886. 

A.  vulgaris  ssp.  redolens  Hall  et  Clem.,  Cam.  Inst.  Wash.  Publ.  326:75,  1923. 

Stems  4r-S  dm.  tall ;  herbage  quite  glabrous  throughout  and  glandular- 
atomiferous  to  moderately  tomentose,  especially  on  under  surfaces  of  leaves ; 
leaves  broadly  ovate  in  outline,  the  lower  bipinnately  dissected,  the  salient 
lobes  rather  narrow  and  short  (mostlj-  less  than  1  cm.  long),  2-5  cm.  long, 
nearly  as  wide,  the  upper  toothed  to  entire,  narrowly  linear,  the  margin  some- 
times narrowly  revolute;  inflorescence  a  broad  leafy  panicle  with  ascending 
branches  bearing  small  usually  erect  heads  in  delicate  racemes;  involucre 
broadly  campanulate,  2.5-3  mm.  high,  3  mm.  wide,  sparingly  silky  tomentose, 
with  12-14  bracts;  ray-florets  6-10;  disk -florets  8-14. 

Westernmost  Texas,  southern  New  Mexico  and  southeastern  Arizona  (in 
somewhat  atypical  form)  to  Chihuahua  and  Durango,  at  elevations  from  1800 
to  2500  m. 

Texas.  Brewster :  Chisos  Mts.,  Mueller  8242.  New  Mexico.  Dona  Ana :  Organ  Mts.,  Sept. 
23,  1906,  Wooton  and  Standley.  Grant:  Copper  Mines  (Santa  Eita),  Wright  1278,  in  part 
(Pli).  Arizona.  Graham:  Mt.  Graham,  7500  ft.,  Kearney  and  Peebles  9796  (possibly  a  form 
of  A.  Carruthii).  Chihuahua.  In  shade  of  cliffs,  rocky  hills  near  Chihuahua,  Pringle  296 
(Gray,  type;  isotypes,  NY,  Ph,  US),  1059 ;  Soldier  Canyon,  Sierra  Madre,  Jones.  Durango. 
San  Julian  to  Cerro  Prieto,  E.  W.  Nelson  4956  ;  Sandia  Station,  Pringle  13535. 

This  unit  is  somewhat  variable  and  its  distribution  is  within  the  region  occu- 
pied by  ssp.  mexicana.  However,  the  distinctive  foliage  and  sparse  pubescence 
serve  to  distinguish  it  from  mexicana,  so  that  it  appears  to  be  quite  apart.  A 
very  few  intermediates  are  found.  The  lower  leaves  of  spp.  redolens  are  bipin- 
nately dissected  like  those  in  A.  Michauxiana,  a  form  very  different  otherwise 
and  from  an  entirely  different  habitat  in  the  northern  Rockies,  but  the  cor- 
respondence in  leaves  is  suggestive.  Possibly  each  of  these  units  will  prove  to 
have  a  different  chromosome  number  from  A.  ludovicuina.  and  each  may  have 
had  a  long  independent  history  behind  it. 

7.  Artemisia  Lindleyana  Bess,  in  Hook.,  Fl.  Bor.  Amer.  1 :322,  1833. 

Figures  10  and  17 

Type  statement:  "North-West  coast  of  America.  Douglas,  in  Herh.  Lindl. 
n.  16."  The  following  three  collections  in  the  Kew  Herbarium,  collected  by 
David  Douglas,  may  include  an  isotype  of  the  species :  abundantly  on  the 
sands  of  the  seashore  near  the  straits  of  Juan  de  Fuca,  1825,  Douglas  (Hb. 
Benth.,  I  doubt  the  locality  on  this  label )  ;  in  the  channels  of  mountain  streams 
near  the  sources  of  the  Columbia,  common,  Douglas  (possibly  a  part  of  the 
previous  collection,  in  which  case  these  data  as  to  the  source  are  probably 
correct)  ;  Columbia  River,  1830,  Douglas. 


Vol.  XXV]       KECK :  EEVISION  OF  ARTEMISIA  VULGARIS  COMPLEX  455 


;■■*•■  A.  LINDLEYANA 
-  A.  SUKS  DORFII 
•     A    DOUGLASIANA 


Figure  17.  Distribution  of  Artemisia  Lindleyana,  A.  Sulcsdorfii,  and  A.  Douglasiana. 


456  CALIFORNIA  ACADEMY  OF  SCIENCES  [Proc.  4th  Ser. 

Artemisia  Lindleyana  a  legitima  Bess.,  ihid.  "Folia  intcgcrrinia  1-1%  longa,  vix  i;'.,  lata." 
By  inference,  Douglas  no.  16. 

A.  Lindleyana  /3  brevifolia  Bess.,  ibid.  "Fors  status  serotinus:  nam  adsunt  tantum  folia 
fasciculorum  unguiciihnia."  By  inference,  Douglas  no.  14. 

A.  Lindleyana  y  subdcntata  Bess.,  ibid.  "Folis  versus  apicem  uno  alterne  1-%"  [half  inches] 
longa,  1—2"'  [lines]  lata."  By  inference,  Douglas  no.  18. 

A.  Lindleyana  5  Coronoims  Bess.,  ibid.  "Folia  Flantaginis  Coronopi."  These  varieties  are 
leaf  variations  that  appear  to  have  no  eeotypical  value. 

A.  pumila  Nutt.,  Trans.  Amer.  Phil.  Soc.  II.  7:399,  1841:  not  Link,  1822.  A  starred  speci- 
men labeled  in  Nuttall's  hand  "Artemisia  *pumila.  R.  Mts.  U.  Calif."  seen  in  Pliila- 
delphia  Academy. 

A.  tenuis  var.  integerrima  Eydb.,  Mem.  N.  Y.  Bot.  Gard.  1 :432,  1900.  Lower  leaves  missing 
but  upper  ones  entire,  green,  very  lightly  villous  on  under  surface;  herbage 
glandular ;  stems  glabrate,  sulcate.  Assigned  to  this  species  somewhat  doubtfully. 
Type  locality:  Emigrant  Gulch,  Park  Co.,  Montana. 

A.  arachnoidea  Sheu).,  Bull.  Torr.  Club  30:310,  1903.  Leaves  linear-lanceolate,  tapering  to 
base  and  apex,  very  green,  but  somewhat  tomentose  beneath.  A  wide-leaved  form  of 
this,  rather  than  of  A.  Douglasiana  to  which  it  was  assigned  by  Hall  and  Clements 
(under  the  name  A.  vulgaris  ssp.  heterophyUa).  Type  locality:  sandy  banks  of  the 
Columbia  River,  1  mile  west  of  Vancouver,  Washington. 

A.  Leibergii  Eydb.,  N.  Amer.  Fl.  34:267,  1916.  Leaves  narrow  in  type  specimen,  remotely 
but  sharply  toothed  apieally,  nearly  glabrous.  Type  locality:  Fish  Hook  Ferry, 
Adams  Co.,  "Washington. 

A.  vulgaris  ssp.  Lindleyana  Hall  et  Clem.,  Carn.  Inst.  Wash.  Publ.  326 :  79,  1923. 

A.  vulgaris  var.  Lindleyana  Jeps.,  Man.  Fl.  PL  Calif.  1142,  1925. 

Chamaephj^te ;  stems  2-4  dm.  tall,  several  from  a  woody  base ;  basal  leaves 
linear-oblanceolate  (rarely  spatulate  or  wider),  entire  to  serrate-dentate  api- 
eally or  more  deeply  lobed,  often  crowded,  with  fascicles  developing-  in  their 
axils,  the  npper  mostly  entire,  2—5  cm.  long,  mostly  less  than  1  cm.  wide, 
tomentiilose  tlironghont,  or  more  commonly  green  and  glabrate  above,  white- 
tomentose  beneath ;  inflorescence  a  racemose  or  spike-like  panicle,  usually  nar- 
row and  short,  1-2  cm.  wide ;  involucre  campanulate,  3  mm.  high,  2-3  mm. 
wide,  usually  lightly  tomentulose,  with  9-13  bracts;  ray-florets  5-9;  disk- 
florets  10-30. 

Drainage  of  the  Columbia  River,  mostly  along  the  banks  of  this  stream  and 
its  tributaries,  from  western  Montana  and  southern  British  Columbia  to 
Washington  and  Oregon  ;  mostly  from  low  elevations,  but  ascending  to  1980  m. 
in  IMontaiia.  Flowering  from  July  through  September. 

British  Columbia.  Yale:  Sicamous,  July  3,  1889,  Ilacoun  (NY,  doubtfully  referred 
here).  New  Westminster:  Fort  Hope,  Howell  7766.  Montana.  Flathead:  Flathead  Lake. 
Sanders:  Weeksville,  Leiberg  1567;  Plains,  Kirlcwood  2510.  Lake:  St.  Ignatius,  Mission 
Mts.,  Kirlcwood  1347.  Missoula:  Monture  Trail.  Park:  Emigrant  Gulch,  Aug.  23,  1897, 
6500  ft.,  Fydberg  and  Bessey  5201a  (NY,  type  of  A.  tenuis  var.  integerrima;  isotype,  US). 
Idaho.  Kootenai:  near  Thompson  Mt.,  Leiberg  1610.  Washington.  Stevens:  Old  Ft.  Col- 
ville.  Whitman:  Wawawai.  Franklin:  Fish  Hook  Ferry,  Leiberg  935  (NY,  type  of  A.  Lei- 
bergii; phototype,  C;  isotype,  US),  934,  936,  937.  Chelan:  Entiat;  Wenatchee.  Yakima: 
Priest  Rapids.  Klickitat:  Grand  Dalles,  Keck  341.  Clark:  Vancouver,  Aug.  31, 1902,  Sheldon 
S.  11284  (NY,  type  of  A.  arachnoidea ;  isotypes,  Po,  US).  Oregon.  Malheur:  3  mi.  N.  of 


Vol.  XXV]       KECK:  EEVISION  OF  ARTEMISIA  VULGABIS  COMPLEX  457 

Adrian,  Peck  16070.  Sherman:  Biggs,  Ball  11638.  "Wasco:  mouth  of  Deschutes  River.  Hood 
River :  Hood  River.  Multnomah :  Hayden  Island. 

This  species  has  not  been  well  understood.  Attention  has  been  focused  only 
on  traditional  taxonomic  characters,  and  little  attention  has  been  paid  to  its 
distribution  and  ecologic  connections.  As  a  resvilt  an  unnatural  assemblage 
of  plants,  distributed  very  disruptedly  over  most  of  the  western  states  and 
having-  superficial  similarities  in  leaves  and  inflorescence,  has  been  assembled 
under  this  name.  For  these  reasons  it  was  not  discovered  that  A.  Lindleyana 
is  a  suffrutescent,  narrow-leaved  species,  that  follows  the  banks  of  the  Colum- 
bia River  and  its  tributaries. 

The  nearest  connections  of  Lindleya^ia  appear  to  be  with  sspp.  incompta 
and  typica  of  tetraploid  ludoviciana,  but  possibly  a  close  relationship  can  be 
demonstrated  between  it  and  hexaploid  Douglasiana. 

8.  Artemisia  Suksdorfii  Piper,  Bull.  Torr.  Club  28  :42,  1901. 

Figures  3,  10,  and  17 

Type  locality  :  Stony  sea-beaches  near  Fairhaven,  Washington. 

Artemisia  heterophylla  Nutt.,  Trans.  Amer.  Phil.  Soc.  II,  7:400,  1841,  in  part;  not  Bess., 
1834. 

A.  vulgaris  var.  litoralis  Suksd.,  Deutsch.  Bot.  Monatschr.  18:98,  1900.  This  and  A.  Suks- 
dorfii are  based  upon  the  same  type  collection. 

A.  vulgaris  ssp.  litoralis  Hall  et  Clem.,  Carn.  Inst.  Wash.  Publ.  326 :  76, 1923. 

Stems  6-12  (-15)  dm.  tall,  stout,  herbaceous,  or  somewhat  suffrutescent  at 
base ;  leaves  all  broadly  lanceolate  to  elliptic,  tapering  to  base  and  apex,  or 
some  broadly  oblanceolate  and  merely  obtuse  at  apex,  entire  or  with  a  few 
coarse  teeth  or  short  lobes,  gradually  reduced  within  inflorescence,  8-15  cm. 
long,  1.5-3  cm.  wide,  all  strongly  discolored,  bright  deep  green  and  glabrous 
or  sparsely  tomentulose  above,  silvery  tomentose  beneath,  usually  plane ;  in- 
florescence a  very  dense  terete  or  narrowly  ovoid  leafy  panicle,  4—10  cm. 
broad;  heads  small,  commonly  erect;  involucre  terete  or  narrowly  ovoid,  3-^ 
mm.  high,  less  than  2  mm.  wide,  slighth^  tomentulose  or  glabrate,  yellow-green 
and  shining,  with  6-9  bracts;  ray-florets  3-7 ;  disk-florets  2-8.  w  =  9. 

Vancouver  Island  southward  to  Sonoma  County,  California,  in  the  imme- 
diate vicinity  of  the  coast,  following  a  few  of  the  rivers  a  short  distance  inland, 
apparently  not  ascending  more  than  100  m.  above  sea-level. 

British  Columbia.  Westminster :  Chilliv?ack  Valley,  Macoun  26352.  Nanaimo  :  Qualicum ; 
Alberni  Canal;  Ucluelet.  Victoria:  Sidney;  Victoria.  Washington.  Whatcom:  Chuckanut 
Bay,  Sul-sdorf  980  (C,  NY,  US,  isotypes).  San  Juan:  Kanaka  Bay,  Zeller  909.  Snohomish: 
Dean  Creek,  S.  of  White  Horse,  KecTc  5205.  Clallam:  Sequim;  Olympic  Mts.,  Elmer  2603; 
Beaver;  Mora.  Jefferson:  Duckabush.  King:  Seattle;  E.  of  Scenic.  Mason:  Hoodsport. 
Pierce:  Steilacoom.  Grays  Harbor;  Montesano,  Heller  3976.  Cowlitz:  4  mi.  N.  of  Kelso, 
Keck  5213 ;  Woodland.  Oregon.  Clatsop:  Seaside.  Tillamook:  Garibaldi.  Marion:  Chemawa. 
Lincoln:  Taft,  Keck  5217.  Douglas:  mouth  of  Umpqua  River,  T.  Howell.  Coos:  McKinley- 
ville.  Curry :  Langlois ;  Gold  Beach ;  mouth  of  Chetco  River.  California.  Del  Norte :  S.  of 
Klamath,  Keck  5225.  Humboldt:  Scotia;  Cape  Mendocino;  Shelter  Cove.  Mendocino: 
Westport ;  Fort  Bragg.  Sonoma  :  Duncan  Mills. 


458  CALIFORNIA  ACADEMY  OF  SCIENCES  [Proc.  4th  Ser. 

Chromosome  counts  were  made  in  meiotie  divisions  of  plants  from  White 
Horse,  Snohomish  Co.,  Washington,  and  Trinidad  and  Van  Duzen  River  near 
Carlotta,  Humboldt  Co.,  California,  and  in  somatic  plates  in  root-tips  of 
plants  from  north  of  Fort  Bragg  and  Jenner,  California. 

A  collection  from  the  Columbia  plains,  Nuttall  (NY,  Ph)  is  labeled  and 
starred  by  Nuttall  as  his  A.  heterophylla;  but  Hall  and  Clements  chose  as 
the  type  of  this  entity  the  sheet  in  Gray  Herbarium  which  has  locality  data 
corresponding  to  that  in  the  original  description,  and  this  sheet  is  A.  Doug- 
lasiana  Bess.  Nuttall  lumped  the  two  collections  (and  species)  without  detect- 
ing the  importance  of  the  differences  between  them. 

Despite  an  exact  duplication  of  habit,  the  more  strongly  discolored  leaves, 
very  narrowly  cylindric  and  yellowish  glabrate  heads,  and  compact  inflor- 
escence of  Suksdorfii  serve  to.  distinguish  this  species  at  a  glance  from  Doug- 
lasiana  in  the  field. 

9.  Artemisia  Douglasiana  Bess,  in  Hook.,  Fl.  Bor.  Amer.  1 :323,  1833. 

Figures  6,  10,  and  17 

Type  statement :  "North-West  America.  Douglas,  in  Herb.  Lindl." 

Artemisia  vulgaris  var. calif  ornica'BESS.,'Linjiaea  15:91, 1841.  Type  locality:  San  Francisco. 

A.  heterophylla  Nutt.,  Trans.  Amer.  Phil.  Soc.  II,  7:400,  1841,  in  part  (as  to  the  specimen 
at  Gray  Herbarium  chosen  by  Hall  and  Clements  as  the  type,  not  as  to  starred  speci- 
men at  Philadelphia)  ;  not  Bess.,  1834. 

A.  ludoviciana  var.  Douglasiana  D.  C.  Eat.  in  S.  Wats.,  Bot.  King's  Expl.  183,  1871. 

A.  Kennedyi  A.  Nels.,  Proc.  Biol.  Soc.  Wash.  18:175,  1905.  The  interior  form  of  the  species 
that  has  migrated  eastward  through  the  low  northern  Sierran  passes.  Type  locality: 
Verdi,  Washoe  Co.,  Nevada. 

A.  vulgaris  ssp.  heterophylla  Hall  et  Clem.,  Cam.  Inst.  Wash.  Publ.  326:76, 1923. 

A.  vulgaris  var.  heterophylla  Jeps.,  Man.  Fl.  PL  Calif.  1142,  1925. 

A.  vulgaris  ssp.  Douglasiana  St.  John,  Fl.  S.  E.  Wash.  422, 1937. 

Stems  5-15 (-30)  dm.  tall,  stout  but  herbaceous  to  base,  simple  or  some- 
times ±  branched,  the  branches  erect  to  divergent;  leaves  all  lanceolate  to 
elliptic  and  entire,  or  oblanceolate  to  obovate  in  outline  and  coarsely  fcAV- 
lobed  or  toothed  toward  apex,  with  the  margin  extremely  variable  in  outline, 
the  entire  lobes  mostly  lanceolate,  7-15  cm.  long,  1-3  (-5)  cm.  wide  exclu- 
sive of  salient  lobes,  those  of  the  inflorescence  gradually  reduced  in  size,  all 
strongly  discolored,  sparsely  tomentulose  to  glabrous  and  green  above,  densely 
gray-tomentose  beneath,  usually  plane;  inflorescence  a  leafy  open  or  dense 
elongated  panicle,  3-15  cm.  broad ;  heads  medium-sized,  erect  or  nodding ; 
involucre  campanulate,  3-4  mm.  high,  2-3  nmi.  wide,  ±  tomentose,  with  8-14 
bracts;  ray-florets  6-10;  disk-florets  10-25.  ?i  =  27. 

Washington  to  northern  Baja  California;  rare  east  of  the  Cascadean- 
Sierran  axis  except  about  Lake  Tahoe  where  it  exteiuls  slightly  into  Nevada, 
and  at  elevations  rarely  up  to  2000  m. ;  local  in  northern  Idaho.  Flowering 
from  June  to  October. 


: 


Vol.  XXV]       KECK:  REriSlON  OF  ARTEMISIA  VULGARIS  COMPLEX  459 

"Eocky  Mountains  by  streams,"  Nuttall  (Gray,  type  of  A.  JwteropliyUa  Nutt.  as  fixed  by 
Hall  and  Clements;  tracing,  C).  Idaho.  Kootenai:  Lake  Coeur  d'Alene,  1892,  Aiton  (Po). 
Benewah :  St.  Joe  River  below  St.  Maries.  Nevada.  Washoe :  Verdi,  Sept.  29,  1904,  Kennedy 
963  (RM,  type  of  A.  Kennedyi).  Douglas:  Glenbrook.  Washington.  Okanogan:  Oroville, 
June  26,  1911,  Jones.  Yakima:  Yakima  region.  Cotton  876.  Klickitat:  Sulsdorf  871.  Colum- 
bia River,  North  West  America,  1830,  Douglas  (Herb.  Benth.,  Kew,  possibly  of  the  type  col- 
lection). Oregon.  Hood  River:  Lookout  Mt.,  4000  ft.,  1926,  Simon.  Multnomah:  Portland, 
Sheldon  11155.  Marion:  Salem.  Crook:  Prineville,  Leiberg  837.  Klamath:  Lake  Ewana, 
Lawrence  1196.  Douglas:  Riddle.  Coos:  Marshfield,  House  4991.  Jackson:  Trail  Creek;  S.  of 
Ruch ;  Siskiyou  Mts.,  Heller  12635.  Curry :  Brookings.  California.  Without  exact  locality, 
1835,  Douglas  (Herb.  Hook.,  Kew,  possibly  type  collection  of  A.  vulgaris  var.  calif ornica) . 
Modoc:  Alturas,  Hall  11670.  Siskiyou:  Yreka,  Butler  1798.  Lassen:  Horse  Lake.  Shasta: 
Castella.  Trinity:  summit  Scott  Mts.;  Yolo  Bolly.  Humboldt:  Hoopa  Valley;  Van  Duzen 
River  opposite  Buck  Mt.  Plumas:  Red  Clover  Valley,  Heller  and  Kennedy  8876.  Tehama: 
Payne  Creek.  Butte:  Jonesville,  Copeland  in  U.  C.  set  484.  Glenn:  Chico  Landing.  Men- 
docino: Comptche,  Jones  29108.  Nevada:  Donner  Lake;  Grass  Valley.  Yuba:  Dobbins. 
Colusa:  Princeton.  Lake:  Snow  Mt. ;  Kelseyville.  Eldorado:  Fallen  Leaf  Lake;  White  Hall. 
Solano:  Vallejo.  Marin:  Tiburon,  Heller  5724.  San  Joaquin:  Holt,  plants  over  11  ft.  high, 
Howell  5514.  Contra  Costa:  Berkeley.  Mono:  Walker  River,  11  mi.  S.  of  Coleville,  Wolf 
3279.  Tuolumne:  Confidence;  Mather,  KecTc  1336.  Mariposa:  Yosemite  Valley.  Madera: 
7  mi.  from  French  Meadows,  Kennedy  and  Doten  420.  Inyo:  Sobrina  Lake,  Jones.  Fresno: 
Laton,  4  m.  tall,  Hall.  Tulare :  Hockett  Meadows,  9400  ft.,  Purpus  2095;  10  mi.  E.  of  Visalia. 
Kern:  Rosedale ;  Tehachapi  Pass.  Santa  Clara:  Palo  Alto,  Baker  1563;  Arroyo  Bayo. 
Stanislaus:  Adobe  Creek,  Red  Mts.  Monterey:  Pacific  Grove,  Heller  7195.  San  Luis  Obispo: 
Santa  Margarita.  Santa  Barbara:  Santa  Maria;  Santa  Cruz  Island,  Rand  71.  Ventura: 
Sespe  Creek;  Triunfo.  Los  Angeles:  Mesmer,  Ahrams  2955  ;  Claremont ;  Swartout  Valley; 
Santa  Catalina  Island.  San  Bernardino:  Victorville,  Parish  10583;  Mill  Creek  Canyon. 
Riverside :  Keen  Camp ;  Cold  Water  Canyon.  Orange :  Newport  Beach.  San  Diego :  Warner 
Springs.  Baja  California.  Tia  Juana;  Rio  Santo  Domingo,  Wiggins  and  Demaree  4781; 
Rancho  San  Jose,  E.  of  San  Telmo,  Wiggins  and  Demaree  4835. 

Chromosome  counts  were  made  on  the  following  plants,  all  somatic  counts 
on  single  plants  from  California  stations  unless  otherwise  indicated :  Santa 
Barbara  (also  meiotic) ;  Pismo  Hills,  30  m.  elev. ;  Halls  Valley,  Mt.  Hamilton, 
490  m.  (two  plants)  ;  Santa  Cruz  Mts.  between  Los  Gatos  and  Glenwood, 
450  m. ;  Mather,  1400  m.  (two  plants)  ;  Smith  Peak,  Yosemite  National  Park, 
2200  m.  (one  plant  somatic,  one  meiotic)  ;  Medford,  Oregon,  415  m. 

In  the  San  Francisco  Bay  region,  the  leaves  of  Douglasiana  are  usually 
more  lobed  than  elsewhere,  but  this  deviation  from  the  usual  pattern  is  in- 
sufficient to  warrant  nomenclatorial  recognition. 

The  hypothesis  has  been  proposed  by  Clausen,  Keck,  and  Hiesey  (1940a, 
pp.  342-344),  that  Douglasiana,  a  hexaploid  species  of  inland  valleys  and 
moderate  altitudes  on  the  Pacific  slope,  originated  by  amphidiploidy  (addition 
of  the  chromosomes)  from  the  coastal  diploid  species,  Suksdorfii,  and  the 
interior  montane  tetraploid,  ludoviciana,  whose  essential  characters  it  recom- 
bines.  Transplant  experiments  demonstrated  that  the  tall  Suksdorfii  is  unfitted 
to  occupy  higher  elevations.  It  was  even  a  poor  survivor  at  the  Mather  trans- 
plant station  at  1400  m.  elevation  where  Douglasiana  is  native.  The  smaller 
statured  ludoviciana,  on  the  other  hand,  ranging  widely  toward  the  eastward, 


460  CALIFORNIA  ACADEMY  OF  SCIENCES  [Proc.  4th  Ser. 

has  been  able  to  occupy  even  alpine  habitats.  It  enters  the  range  of  Stiksclorfii 
in  the  Columbia  River  Gap,  and  here  In'bridization  may  have  been  possible. 
Artemisia  Douglasiana  is  found  in  the  geographically  and  climatically  inter- 
mediate zone  between  the  ranges  of  its  presumed  parents. 

10.  Artemisia  Prescottiana  Bess,  in  Hook.,  Fl.  Bor.  Amer.  1 :324,  1833. 

Figure  10 

Type  .statement:  "North-We.st  America.  Douglas  in  Herh.  Lindl."  Known 
only  from  the  type  collection  made  in  1825  by  David  Douglas  on  the  Quick 
Sand  River,  near  the  Grand  Rapids  of  the  Columbia.  (This  is  near  The  Dalles, 
but  whether  from  the  Washington  or  Oregon  side  is  undetermined.)  The  sheet 
in  Bentham's  herbarium  at  Kew  has  been  studied.  A  note  on  it  by  R.  A.  Rolfe 
states  that  it  exactly  corresponds  with  the  type  of  A.  Prescottiana  in  the  Lind- 
ley  Herbarium.  A  drawing  and  fragment  of  this  plant  are  at  New  York. 

Stems  3-4  dm.  tall,  slender,  several  branching  from  the  slender  slightly 
suffrutescent  base,  striate,  smooth  and  shining,  like  especially  the  under 
surface  of  the  leaves  very  sparingly  arachnoid-pilose,  the  very  slender  hairs 
mostly  spreading;  leaves  2-5  cm.  long,  less  than  1  mm.  wide,  pinnatifid  with 
3-7  filiform  caudate  entire  divaricate  remote  divisions  5-18  mm.  long,  or  those 
of  the  inflorescence  entire,  green,  the  margin  closely  revolute ;  inflorescence 
racemose,  sparingly  leafy,  the  few  erect  heads  on  short  filiform  pedicels  well 
spaced;  involucre  hemispheric,  3.5-4  mm.  high,  4-5  mm.  wide,  green  and  shin- 
ing, the  10  bracts  very  sparingly  arachnoid-pilose,  ±:  ciliate ;  ray -florets  6 ; 
disk- florets  23  (only  one  head  counted) . 

I  have  seen  no  other  specimen  than  the  one  in  the  Bentham  herbarium  that 
could  be  referred  to  this  species,  so  it  is  assumed  that  it  is  a  local  endemic  not 
as  yet  rediscovered,  or  that  it  is  by  this  time  extinct.  The  branching  base  is 
unique,  slender  branches  developing  from  each  of  the  lower  nodes ;  the  lowest 
and  strongest  become  flowering  stems,  those  a  little  higher  are  sterile  almost 
capillar}^  leafy  shoots  about  10  cm.  long.  The  filiform  and  filiform-lobed  leaves, 
which  are  not  at  all  fascicled,  and  the  pubescence  are  also  unduplicated,  and 
the  racemose  arrangement  of  the  heads  is  only  approached  in  forms  of  A.  Lirid- 
leyana  and  A.  Michauxiana,  plants  that  otherwise  have  very  little  in  common 
with  this.  In  other  words,  while  the  floral  characters  plainly  show^  that  A. 
Prescottiana  is  a  member  of  this  complex,  it  does  not  seem  to  be  very  closely 
related  to  any  other  species. 

11.  Artemisia  Tilesii  Ledeb.,  Mem.  Acad.  St.  Petersb.  5  :  568, 1814. 

11a.  Artemisia  Tilesii  Ledeb.  ssp.  typica  Keck  nom.  nov. 

Figures  10  and  IS 
Tj^pe  locality :  Kamchatka.  Type  not  seen. 

Artemisia  Tilesii  Ledeb.,  loc.  cit. 

A.  Tilesii  var.  arctica  Bess,  in  Hook.,  Fl.  Bor.  Amer.  1 :324,  18.33.  "Ex  littoribus  arcticis  est 

var.  f  arctica — folioruni  laciniis  intcgerrimis  magis  distantibus  obtusis  brevibus,  pcri- 

cliniis  flosculisque  pallidis." 


Vol.  XXV]       KECK:  EEVISION  OF  ARTEMISIA  VULGARIS  COMPLEX  461 


Figure  18.  Distribution  of  Artemisia  Tilesii  in  North  America. 


462  CALIFOBNIA  ACADEMY  OF  SCIENCES  [PROC.  4th  Ser. 

A.  vulfjaris  var.  kamtschatica  Bess.,  Nouv.  Mem.  Soc.  Nat.  Mosc.  3  -.54,  1834. 

A.  vulgaris  var.  Tilesii  Ledeb.,  F1.  Eossica  2  :586,  1844-46. 

A.  vulgaris  ssp.  Tilesii  Hall  et  Clem.,  Cam.  Inst.  Wash.  Publ.  326:72, 1923. 

Stems  1.5-6  dm.  tall,  floccose  when  young  but  soon  glabrate  ;  leaves  ovate  to 
lanceolate  in  outline,  deeply  incised  into  3-5  lanceolate  attenuate  sparingly 
toothed  or  dentately  lobed  divisions,  the  leaves  of  the  inflorescence  merely 
toothed  or  entire,  all  strongly  discolored,  green  above,  tomentose  beneath;  in- 
florescence a  rather  compact  spike-like  panicle  often  overtopped  by  the  leaves ; 
heads  nodding,  large;  involucre  hemispheric,  4-5  mm.  high,  6-9  mm.  wide,  the 
bracts  12-28,  ±  anthocyanous,  with  broad  scarious  margin,  glabrous  and 
shining,  or  the  outermost  somewhat  tomentose  along  midrib ;  ray-florets  9-25  ; 
disk-florets  20-100. 

Nova  Zembla,  nortliern  fSiberia  and  Kamchatka  to  the  Pribilof  Islands, 
central  and  northern  Alaska,  Yukon  Territory  and  east  to  Hudson  Bay.  Flow- 
ering from  June  to  September. 

North  American  stations:  Alaska.  Bering  Sea,  Macoun  89;  St.  Lawrence  Island;  St.- 
Matthew  Island;  Nunivak  Island;  Punuk  Island;  St.  Paul  Island;  St.  George  Island; 
Kutzeblok ;  Taylor ;  Nome,  G.  N.  Janes  9211 ;  Norton  Sound ;  Postoliak  Elver,  Yukon  Delta ; 
Circle.  Yukon.  Herschel  Island,  Arctic  Ocean,  Aug.  19,  1896,  A.  Seale  (SU)  ;  Dawson, 
Macoun  79012;  Lake  Kluane  to  Don  Jek  Eiver,  1920,  Midler.  British  Columbia.  White 
Pass,  Eastwood  899.  Mackenzie.  Arctic  shore  between  Mackenzie  and  Coppermine  rivers, 
Bichardson  (NY)  ;  Coppermine  Eiver;  Tree  Eiver;  Hood  Eiver,  Bathurst  Inlet.  Manitoba. 
Churchill,  Hudson  Bay,  lat.  58°  50',  Macoun  79263. 

lib.  Artemisia  Tilesii  ssp.  unalaschcensis  (Bess.)  Hulten, 

Fl.  Aleut.  Ids.  327, 1937. 

Figures  9,  10,  andl? 

Type  locality  :  Island  of  Unalaska.  Type  not  seen. 

Artemisia  HooTceriana  Bess,  in  Hook.,  Fl.  Bor.  Amer.  1:322,  1833.  The  type,  from  Eocky 
Mountains,  Canada,  Dntmmond  (Kew),  has  been  studied.  A  sheet  that  matches  the 
fragmentary  type  specimen  so  closely  as  to  seem  to  be  part  of  the  same  plant  is 
labeled,  E.  Mts.,  Saskatchewan,  4-5  ft.,  Drummond  23  (Kew).  Leaves  strongly  dis- 
colored, bright  green  above,  very  white  beneath,  to  17  cm.  long,  many  entire,  linear- 
lanceolate,  tapering  to  base  and  apex,  some  with  1-5  remote  salient  linear:oblong 
usually  entire  lobes  1.5-4.5  cm.  long. 

A.  vulgaris  var.  vulgatissima  Bess,  in  Hook.,  ibid.  Not  seen  but  probably  referable  here. 
Type  statement:  "North-West  coast.  Douglas,  in  Herb.  Lindl." 

A.  vulgaris  var.  americana  Bess.,  Linnaea  15:105,  1841,  in  part,  judging  from  the  distribu- 
tion and  description.  Not  seen. 

A.  Tilesii  var.  unalaschcensis  Bess.,  ihid.  106. 

A.  Tilesii  var.  elatior  Tork.  et  Gray,  Fl.  N.  Amer.  2:422,  1843.  "Subarctic  America, 
Richardson." 

A.  elatior  Eydb.,  Mem.  N.  Y.  Bot.  Gard.  1:430,  1900. 

A.  unalaskensi.s  Eydb.,  N.  Amer.  Fl.  34 :  266,  1916.  Not  based  on  A.  Tilesii  var.  unalaschcensis 
Bess.  1841.  Type  collected  on  the  island  of  Unalaska,  Aug.  22,  1891,  Macoun  20625. 
Leaves  large,  subpalmately  parted  or  divided,  the  lobes  again  cleft. 

A.  Gormanii  Eydb.,  ihid.  267.  "Type  collected  in  the  Lake  Iliamna  Eegion,  Alaska,  Septem- 
ber 1,  1902,  Gorman  281  (U.S.  Nat.  Herb.  no.  420345)."  Leaves  more  finely  cut  and 


Vol.  XXV]       KECK:  BEFISION  OF  ARTEMISIA  VULGARIS  COMPLEX  463 

the  divisions  more  slender  and  attenuate  and  the  inflorescence  less  leafy  than  the 
type  of  A.  wnalasTcensis  Eydb.  This  is  the  form  first  described  as  A.  Tilesii  var.  elaiior 
Torr.  et  Gray, 

A.  obtusa  Eydb.,  ibid.  274.  This  plant,  known  only  from  the  type  specimen,  collected  "on  the 
Columbia"  by  Scouler,  no.  234,  is  very  doubtfully  placed  here.  Scouler's  collections 
were  confined  to  the  year  1825  and  apparently  were  taken  near  the  ocean  along  the 
coast  from  Nootka  Sound,  Vancouver  Island,  southward  to  the  mouth  of  the  Columbia, 
and  at  Fort  Vancouver.  The  fa«t  that  this  form  has  apparently  not  been  recollected 
leaves  its  status  in  much  doubt.  The  plant  has  leaves  thicker  in  texture  and  more 
floccose  on  the  upper  surface,  but  otherwise  cut  about  the  same  as  those  found  in 
A.  Tilesii  ssp.  unalaschcensis. 

A.  Tilesii  var.  elaiior  f.  pubescens  G.  N.  Jones,  Univ.  Wash.  Publ.  Biol.  5  :254, 1936.  A  form 
from  the  Olympic  Mountains  in  which  the  involucre  is  tomentose. 

A.  unalasTcensis  var.  aleutica  Hulten,  F1.  Aleut.  Ids.  327,  1937.  Form  with  even  the  upper 
side  of  the  leaves  subtomentose-lanuginous.  Type  not  given,  but  first  plant  cited  was 
collected  on  Atka  Island  in  1834  by  Wrangell  (Leningrad).  The  variety  is  said  to  be 
limited  to  the  middle  and  western  Aleutians. 

A.  ludoviciana  var.  americana  Febn.,  Ehodora  47:248,  1945.  Both  Fernald  and  the  writer 
have  tried  to  place  Besser's  variety  on  the  basis  of  the  original  description  alone,  and 
the  results  do  not  agree.  Fernald  includes  in  it  an  assemblage  of  forms  having  leaves 
bright  green  above  and  panicles  leafy  and  open,  scattered  from  the  Lower  Athabasca 
Eiver,  northern  All)erta,  to  northern  Mexico.  The  writer  assigns  duplicates  of  the 
collections  Fernald  cites  to  five  different  units :  the  type,  at  least,  to  the  present  sub- 
species ;  most  of  the  remainder  to  four  subspecies  of  A.  ludoviciana,  namely,  sspp. 
mexicana  (Texas,  New  Mexico,  Colorado,  in  part),  incompta  (Idaho),  typica  (Colo- 
rado, in  part),  and  sulcata  (Arizona).  Fernald  himself  cites  two  of  them  again  imder 
his  A.  ludoviciana  var.  mexicana. 

Stems  3-15  dm.  tall,  floccose  when  young  but  soon  glabrate ;  leaves  extremely 
variable,  linear-lanceolate  to  oblanceolate,  ovate,  or  subrotund  in  outline,  the 
narrowest  sometimes  quite  entire,  the  others  bluntly  or  sharply  few-toothed 
or  -lobed  to  deeply  pinnately,  or  commonly  subpalmately,  divided  with  some 
of  the  segments  again  toothed,  lobed,  or  parted,  those  leaves  with  merely  teeth 
or  lobes  often  with  wide  lamina,  those  again  lobed  often  with  narrow  lamina 
and  segments,  floral  leaves  entire  or  nearly  so,  all  strongly  discolored,  green 
above  (rarely  tomentulose),  tomentose  beneath,  plane,  or  the  margin  some- 
what revolute ;  inflorescence  a  ±  elongated  narrow  spike-like  panicle,  some- 
times rather  open,  or  a  broad  dense  spike,  well  exceeding  the  leaves;  heads 
erect  or  nodding,  large;  involucre  hemispheric  to  broadly  campanulate  or 
turbinate,  3-5  mm.  high,  4-8  mm.  wide,  the  bracts  11-13,  often  darkly  antho- 
cyanous.  sometimes  green,  with  broad  hyaline  margin,  glabrate  or  tomentu- 
lose; ray-florets  8-15  ;  disk-florets  20-40.  n  =  27. 

Northern  Japan  (Hokkaido,  etc.)  to  Kamchatka,  the  Aleutian  Islands, 
southern  Alaska,  Yukon  and  Mackenzie,  southward  to  western  Montana  and 
northern  Oregon ;  from  sea-level  in  the  north  up  to  1830  m.  elevation  in  the 
south.  Flowering  from  July  to  September. 

North  American  stations:  Attn  Island  (as  A.  unalasTcensis  Eydb.)  and  Attn,  Agattu, 
Kiska,  and  Atka  islands  (as  A.  unalasTcensis  var.  aleutica  Hult.)  all  reported  by  Hulten, 
loc.  cit.  Unalaska  Id.:  Aug.  22,  1891,  Macoun  20625  (Canada,  type  of  A.  unalasTcensis; 


464  CALIFORNIA  ACADEMY  OF  SCIENCES  [Proc.  4th  Rer. 

phototypes,  C,  CAS,  NY,  US),  Eyerdam  2346;  Dutch  Harbor.  Akutan  Id.,  Norberg  382. 
Unimak  Id. :  False  Pass,  Eyerdam  2250.  Kodiak  Id. :  Old  Harbor.  Alaska.  Porcupine  Eiver, 
Turner ;  Wonder  Lake,  McKinley  National  Park,  Mexia  2252  ;  Lake  Ilianma  region,  sandy 
and  rocky  slopes  above  high  tide  south  of  Iliamna  Point,  Sept.  1,  1902,  M.  W.  Gorman  281 
(US,  type  of  A.  Gormanii;  phototypes,  C,  NY)  ;  Yentna  District,  above  Morgan  Creek; 
Seward  ;  Cordova,  Mexia  2311 ;  Red  Eiver ;  Skagway ;  Chilkat  Valley,  Walker  1077.  Yukon. 
Dawson,  Eastwood  455 ;  Klondyke.  Mackenzie.  Fort  Wrigley,  Preble  and  Cary  86.  Alberta. 
Peace  River  10  mi.  above  Carcajou  Settlement,  Baup  and  Abbe  4359 ;  Boiler  Rapid,  Atha- 
basca B.iYer, Preble  and  Cary  i5J.  British  Columbia.  Cassiar  District,  pass  near  head  Tset-ee- 
yeh  River,  a  branch  of  Klappan  River,  Preble  and  Mixter  640.  New  Westminster:  Chilliwack 
Valley,  Macoun  26350.  Montana.  Flathead:  McDonald  Peak,  Flathead  Lake,  Jones  8775. 
Gallatin:  Bozeman,  mountain  canyons,  6,000  ft.,  Blanlcinship  305.  Idaho.  Bonner:  Lake 
Pend  d'Oreille,  Sandberg  et  al.  754a.  Washington.  Clallam:  Sequim;  Mt.  Angeles,  Thomp- 
son 7502.  Chelan :  Stehekin ;  Mt.  Stuart ;  Tumwater  Canyon,  Thompson  8425.  Pierce :  Mt. 
Ranier;  White  River.  Yakima  (?)  :  Mt.  Adams,  Gardner.  "Columbia"  (River),  Scolder  234 
(NY,  type  of  A.  obtusa;  phototype  and  fragments,  C).  Oregon.  Wallowa:  Wallowa  Mts., 
Piper  2491.  Baker:  Cornucopia,  Jones  29110.  Hood  River:  N.  side  Mt.  Hood,  Thompson 
3530.  Multnomah:  Columbia  River  above  Multnomah  Falls,  Hall  11997. 

The  chromosome  number  was  determined  at  meiosis  in  a  plant  originating 
from  seed  of  Ilulten's  plants  from  Unalaska  grown  at  Stanford. 

Both  ssp.  iypica  and  ssp.  unalaschcensis  are  highly  variable,  but  they  appear 
to  represent  fairly  clear  regional  subdivisions  of  the  species  Tilesii.  Some 
specimens,  however,  must  be  classified  rather  arbitrarily.  The  wide  range  of 
variation  has  prompted  other  workers  to  name  many  forms  as  the  extended 
synonymy  indicates.  But  this  variation  is  extensive  even  in  plants  grown  from 
single  localities  such  as  Unalaska  Island,  and  there  is  no  indication  that  there 
are  any  barriers  to  free  interbreeding  and  recombination. 


Vol.  XXV]       KECK:  BEVISION  OF  ARTEMISIA  VULGABIS  COMPLEX  465 

DISCUSSION  AND  CONCLUSIONS 

111  North  America  eleven  species  are  recognized  in  the  subsection  Vulgares 
of  Ariemisia.  All  but  one  of  these  are  polj-morphic,  several  being  highly  so. 
Two  contain  recognizable  geographic  subspecies  (there  being  seven  such  in 
A.  ludoviciana).  The  conventional  taxonomic  treatment  shows  that  additional 
cytogenetic  investigations  of  this  group  are  needed.  In  fact,  the  abundance 
of  members  of  this  complex  in  the  North  American  flora,  their  great  diversity 
of  form,  and  their  differences  in  chromosome  number  challenge  our  imagina- 
tion and  invite  experimental  studies. 

An  obvious  opportunity  for  further  work  lies  in  a  cytological  survey  of  the 
subsection.  Enough  chromosome  counts  have  been  made  to  show  that  this  is  a 
typical  example  of  the  polyploid  complex.  There  are  special  opportunities  for 
evolution  in  such  complexes,  for  a  great  diversity  of  habitats  is  occupied  by 
forms  closely  enough  related  to  allow  some  mixing  of  hereditary  strains.  This 
in  turn  produces  variation  and  allows  some  of  it  to  become  fixed  through 
genetical  and  ecological  selection.  It  is  to  be  expected  that  in  such  a  complex 
both  mutations  of  various  sorts  and  hybridization  have  played  major  roles 
in  building  up  the  variation. ' 

One  of  the  ways  in  which  sudden  new  steps  in  the  evolutionary  history  may 
arise,  and  the  variation  increase,  is  through  the  crossing  of  two  rather  dis- 
tantly related  species.  The  probable  origin  of  A.  Douglasiana  through  the 
addition  of  all  the  chromosomes  of  two  other  species  is  an  example.  The  strong 
circumstantial  evidence  for  such  an  origin  is  given  elsewhere  (cf.  p.  459),  but 
here  it  should  be  pointed  out  that  one  of  the  supposed  parents,  A.  ludoviciana, 
is  exceedingly  variable  and  occupies  a  wide  range  of  habitats,  while  the  other, 
A.  Suksdorfii,  is  both  morphologically  stable  and  very  limited  in  its  choice  of 
habitats.  The  evolutionary  opportunities  of  Suksdorfii  at  the  present  time 
appear  therefore  limited,  although  this  should  not  lead  us  to  the  conclusion 
that  it  is  either  a  rare  or  decadent  species.  Also,  if  the  theory  of  the  origin  of 
Douglasiana  is  correct,  the  chromosome  set  of  Suksdorfii  combined  with  that 
of  ludoviciana  to  form  the  aggressive  and  variable  species,  Douglasiana.  The 
evolution  of  a  new  species  by  the  addition  of  the  chromosomes  of  two  others 
tends  to  revitalize  the  entire  complex  to  which  they  belong.  The  potential  sig- 
nificance of  Suksdorfii  in  the  evolutionary  course  of  the  complex  is  therefore 
enhanced  through  the  chromosomes  it  probably  contributed  to  Douglasiana. 

The  accompanying  diagram,  figure  19,  attempts  to  show  the  North  Ameri- 
can members  of  subsection  Vtdgares  placed  very  roughly  in  the  relative  posi- 
tions they  would  assume  on  a  map  of  the  continent.  The  geographical  ranges 
are  much  generalized,  and  accurate  limits  are  to  be  found  under  the  taxonomic 
treatment.  Chromosome  numbers  are  given  where  known.  The  arrows  suggest 
possible  courses  of  evolution  as  gathered  from  the  available  evidence  on  mor- 
phological and  cytological  relationships.  The  intangible  impressions  that  come 
from  an  intimate  study  of  a  group  have  also  played  a  part  here. 


466 


CALIFOFXIA  ACADEMY  OF  SCIEXCES 


[Proc.  4th  Ser. 


In  the  case  of  A.  Douglasiana  mentioned  above,  arrows  indicate  its  probable 
origin  from  A.  Suksdorfii  and  A.  ludoviciana.  The  subspecies  of  ludoviciana 
that  seems  to  be  the  most  likely  forebear  of  Douglasiana  is  uncertain,  but 
both  sspp.  candicans  and  typica  are  suggested.  Aside  from  this  case,  in  wliich 

I.VULGARIS 

Old  World 
n  =  8 


typica 
Asia-Alaska 

unalaschcenois 
Alaska-Canada 
V       n  =  27 


n.MICHAUXIANA 

Rocky  Mis. 


m.LONGIFOLIA 

N  Great  Plains 
8 


M.LINDLEYAN 

Columbian  Basi 


SERRATA 

Upper 
Mississippi 
Valley 
n  =  l8 


3Zni.SUKSD0Rr 

Pacific  shores 
n  =  9 


RUTHII 

n  plains 

:9 


H.DOUGLASIANA 

CaliPornia 
n  =  27 


Figure  19.  Chart  of  possible  relationships,  showing  tlie  relative  geographical  position  and 
some  chromosome  numbers  of  the  subsection  Vulgares  of  Artemisia.  See  text. 

the  circumstantial  evidence  Is  strong,  our  present  information  is  too  frag- 
mentary to  indicate  origins ;  therefore,  for  most  of  the  systematic  units  more 
than  one  alternative  must  be  considered. 

In  the  other  instances  in  the  chart  in  which  arrows  from  two  forms  point 
to  a  tliird  one,  am])hidiploidy  or  even  hybrid  origin  is  not  to  be  inferred, 


Vol.  XXV]       KECK:  REVISION  OF  ARTEMISIA  VULGARIS  COMPLEX  467 

although  that  possibility  need  not  be  ruled  out.  Such  arrows  merely  indicate 
that  on  the  present  evidence  it  is  difficult  to  determine  the  exact  relationships 
and  possible  origins.  Furthermore,  it  is  to  be  readily  granted  that  such  variable 
units  as  A.  ludoviciana  sspp.  iypica  and  mexicana  may  have  had  polyphyletic 
origins. 

The  European  source  of  A.  vulgaris  is  indicated  by  an  arrow,  as  is  also  the 
probable  Asiatic  origin  of  A.  Tilesii.  A  broken  line  calls  attention  to  the  mor- 
phological-ecological similarity  between  A.  Tilesii  ssp.  imalaschcensis  and 
A.  ludoviciana  ssp.  candicans,  Possibly  here  is  an  evolutionary  connection 
between  Tilesii  and  the  other  American  species. 

Scarcely  any  two  units  in  the  North  American  assemblage  are  less  alike  than 
the  only  two  known  diploid  species,  A.  Suksdorfii  and  A.  Carruthii.  Both  of 
these  occur  on  the  periphery  of  the  distribution  of  this  complex  and  occupy 
extreme  climates,  the  former,  an  equitable  moist  coastal  zone,  and  the  latter, 
dry  semideserts.  Accordingly,  they  show  extreme  differences  in  size,  habit, 
and  leaf  characters,  although  they  resemble  each  other  in  their  few-flowered 
heads.  These  two  species  inhabit  climates  that  must  have  been  available  from 
mid-Tertiary  time,  at  least.  It  is  of  prime  interest  to  find  such  ecologic  spe- 
cialization in  species  that  have  remained  on  the  diploid  level  and  likely  are 
of  great  age.  Artemisia  Suksdorfii  bears  some  morphological  resemblance  to 
A.  serrata.  The  recurrence  of  this  morphological  type  in  the  central  states 
suggests  that  species  or  ecotypes  of  that  type  in  the  past  may  have  occupied 
those  parts  of  the  North  American  continent  where  now  the  polyploid  species 
are  the  only  representatives. 

The  diversity  of  the  North  American  diploids  and  the  presence  of  another 
anomalous  form  like  A.  Prescottiana  suggest  that  great  gaps  now  exist  in  the 
evolutionary  fabric  of  this  group,  making  the  present-day  picture  incomplete. 
Certainly  the  monophyletic  origin  of  the  subsection  from  any  living  American 
species  appears  quite  impossible. 

The  North  American  diploids  and  the  Eurasiatic  diploid,  A.  vidgaris  L., 
bear  no  close  resemblance  to  one  another.  Other  diploid  progenitors  of  these 
must  therefore  have  existed  which  now  are  lost. 

The  greatest  diversity  in  the  subsection  ViUgares  appears  to  be  found  in 
North  America,  and  the  known  facts  indicate  that  the  American  species  (ex- 
cepting Tilesii)  originated  in  America.  But,  since  the  complex  is  distributed 
both  on  this  continent  and  in  Eurasia,  the  progenitors  of  the  present-day  dip- 
loids may  have  originated  on  either  land  mass  and  migrated  across  the  Bering 
Straits.  Artemisia  Tilesii,  which  now  is  distributed  across  this  migration 
bridge,  appears  to  have  closer  ties  in  Asia  than  in  America,  and  therefore  may 
be  considered  as  having  originated  in  the  west  and  moved  eastward ;  but  the 
Asiatic  flora  is  not  well  known  as  yet,  and  Tilesii  may  prove  to  be  of  American 
origin.  At  all  events,  this  highly  polyploid  species  of  aggressive  distribution 
is  not  presumed  to  be  the  direct  ancestor  of  any  of  the  other  American  forms, 
but  rather  itself  a  derivative  of  the  evolutionary  stream. 


468  CALIFORNIA  ACADEMY  OF  SCIENCES  [Proc.  4th  See. 

LITERATURE  CITED 

Clausen,  J.,  David  D.  Keck,  and  William  M.  Hiesey. 

1939a.  The  concept  of  species  based  on  experiment.  Amer.  Jour.  Bot.  26 :  103-106. 

1939b.  Experimental  taxonomy.  Carnegie  Inst.  Wash.  Year  Book  No.  38:125, 

1940a.  Experimental  studies  on  the  nature  of  species.  I.  Effect  of  varied  environments  on 
western  North  American  plants.  Carnegie  Inst.  Wash.  Publ.  520.  452  pp. 

1940b.  Experimental  taxonomy.  Carnegie  Inst.  Wash.  Year  Book  No.  39:162. 
Hall,  Harvey  M.,  and  Frederic  E.  Clements. 

1923.    The  phylogenetic  method  in  taxonomy.  Carnegie  Inst.  Wash.  Publ.  326.  355  pp. 
HuLT^N,  Eric. 

1937.    Flora  of  the  Aleutian  Islands.  397  pp.  Stockholm,  Aktiebolaget  Thule. 

Rydberg,  Per  Axel. 

1916.    Artemisia;  in  North  American  Flora  34:244-285. 


i 


PROCEEDINGS 

OF  THE 

CALIFORNIA  ACADEMY  OF  SCIENCES 

Fourth  Series 
Vol.  XXV,  No.  18,  pp.  469-478,  pis.  47-48  November  15,  1946 


ALICE  EASTWOOD 


SEMI-CENTENNIAL  PUBLICATICjrWBrine  Biological  LaD'.)fotoiy 

UI  B  K,  A.  H  TT 

JAN  3  1  1947 


No.  18 
THE  GENUS  RUPPIA  L. 


WOODS  HOLE.  MASS. 


BY 


WILLIAM  A.  SETCHELL 

Late  Professor  of  Botany  Emeritus^  University  of  California 

DURING  AN  ATTEMPT  to  find  Suitable  plants  to  demonstrate  the  influence  of 
temperature  on  their  growth,  differentiation,  and  persistence,  attention 
was  attracted  to  those  of  aquatic  habitats,  since  of  the  two  most  essential  cli- 
matic variables,  temperature  and  moisture,  moisture  is  constant.  The  marine 
alg-ae  and  the  marine  spermatophytes  have  lent  themselves  particularly  well 
to  experimentation  and  some  attention  has  been  paid  to  each  group.  Of  sper- 
matophytes which  pass  their  entire  life  history  submerged,  Zostera  and  Ruppia 
are  two  of  those  best  suited  for  study.  Zostera  was  found  to  be  well  adapted 
for  observation  in  the  open  (see  Setchell,  Univ.  Calif.  Pub.  Bot.,  14 :  389—452, 
text  fig.  59, 1929) ,  and  Ruppia  for  observation  both  in  the  open  and  in  culture 
(see  Setchell,  Proc.  Nation.  Acad.  Sci.,  10 :  286-288, 1924) .  Ruppia  is  a  rather 
delicate  aquatic,  without  Avoody  tissue  of  any  appreciable  amount,  with  little 
vegetative  storage  of  starch  or  proteins,  appearing  and  disappearing  accord- 
ing to  variation  in  conditions  of  habitat,  seemingly  likely  to  be  fundamentally 
and  quickly  affected  by  changes  of  temperature  and  salinity,  easy  of  culture, 
and  offering  opportunity  of  being  used  as  a  laboratory  plant  for  the  study  of 
the  influences  of  environmental  factors. 

It  grows  as  a  rule  in  shallow  brackish  water.  The  habitats  vary  from  mari- 
time, and  therefore  coastal,  to  interior  saline  springs,  pools,  and  lakes.  In 
many  ways  it  resembles  in  general  habit  its  near  relative,  ZannichelUa  pahis- 

[469] 


470  CALIFORNIA  ACADEMY  OF  SCIENCES  [Proc.  4th  Ser. 

tris  L.  It  is  called  here  by  the  common  name  of  "Ditch  Grass,"  in  England  as 
"Tassel  Grass,"  but  few  except  botanists  are  aware  of  its  existence. 

Linnaeus  described  the  genus  Ruppia  in  the  Genera  Plantarum  of  1737,  re- 
ferring to  Micheli's  plate  35  of  Buccaferrea  maritima  foliis  minus  acutis  (1729, 
Nova  plantarum  genera),  which  he  quotes,  indicating  by  a  cross  (x)  placed 
after  the  generic  name,  that  he  had  seen,  at  that  time,  only  a  dried  specimen. 
Since  there  are  discrepancies  of  detail  between  the  only  specimen  in  his  her- 
barium, labeled  "I.  maritima"  in  his  own  handwriting  ( !)  (see  Plate  47),  and 
the  plate  of  Micheli,  w^e  may  suspect  that  the  herbarium  specimen  was  acquired 
later.  We  may  conclude  that  the  plate  of  Micheli  clearly  represents  his  con- 
ception of  both  genus  and  the  type  species.  This  specimen  has  a  short,  non- 
spiral  peduncle.  Linnaeus  had  in  his  herbarium  another  specimen  which  he 
labeled  ^'Buppia  spiralis,"  which  has  an  elongated,  spiral  peduncle,  probably 
added  to  his  collection  later  than  the  B.  maritima. 

Linnaeus,  in  1737,  published  the  Hortiis  Cliffortianus.  He  mentions  Buppia 
and  under  it  has  2  series;  first,  among  other  citations  he  lists  Micheli's  Buc- 
caferrea maritima,  foliis  acutissimis ;  and  in  the  second  he  lists  Buccaferrea 
maritima,  foliis  minus  acutis,  pi.  35,  p.  72.  He  also  indicates  that  he  had  col- 
lected a  specimen  near  Leyden  and  had  therefore  become  acquainted  with  the 
living  plant.  Linnaeus  had  not,  at  the  time  of  the  publication  of  the  Hortus 
Cliffortianus,  published  his  idea  of  varieties  under  species,  but  because  of  his 
arrangement  it  may  be  thought  that  he  considered  the  two  "species"  of  Micheli 
to  be  only  varieties  of  one.  Linnaeus'does  not  name  the  second  one. 

In  the  first  edition  of  the  Species  Plantarum,  Linnaeus  simply  refers  to 
"Buccaferrea  maritima,  foliis  acutissimis  Micheli,  Gen.,  72,  t.  35,  and  seem- 
ingly unites  the  two  "species"  of  Micheli.  The  idea  of  possibly  two  species  of 
Buppia,  distinguished  to  some  extent  at  least  by  the  differing  tips  of  the  leaves, 
occurs  several  times  in  the  later  literature  and  perhaps  most  strongly  in 
Hagstrom  (Botan.  Notis.,  1911,  137)  where  his  idea  of  Buppia  maritima  and 
R.  spiralis  is  expressed  thus :  "This  has  obtuse  leaves  and  that  acute."  In  other 
words,  the  Ruppia  maritima  of  Linnaeus  (in  sensu  strict.)  has  acute  leaves, 
corresponding  to  the  ^^ foliis  acutissimis"  of  Micheli,  while  B.  spiralis  has  obtuse 
leaves,  thus  corresponding  to  the  ''foliis  minus  acutis"  of  Micheli.  Micheli's 
figures  on  plate  35,  which  are  really  the  type  of  the  genus  Buppia,  and  there- 
fore presumably  the  Buppia  maritima  of  Linnaeus,  show  short,  noncoiled 
peduncles,  so  that  the  citation  of  "foliis  acutissimis"  by  Linnaeus  in  the 
"Species"  is  misleading.  Linnaeus,  however,  through  the  several  editions  of  his 
"Genera,"  both  in  description  and  also  by  the  specimen  in  his  herbarium, 
suspected  as  having  been  collected  on  the  "Westgota  Resa,"  labeled  (by  him !) 
as  Buppia  maritima  (PL  47,  gives  evidence  that  the  short,  nonspiral  form 
was  the  type  both  of  his  genus  and  of  his  species.  The  occurrence  of  a 
long-spiral,  pedunculate  form  in  his  herbarium,  labeled  (by  him!)  "Buppia 
spiralis"  (PI.  48),  still  further  emphasizes  this  idea.  The  two  specimens  in 
Herb.  Linn,  are  reproduced  in  this  paper.  Linnaeus  then,  in  his  herbarium. 


Vol.  XXV]  SETCHELL:  THE  GENUS  BUPPIA  L.  471 

really  recog-nized  two  species,  Ruppia  maritima  L.  (Spec.  PL,  ed.  1, 127, 1753) 
and  Ruppia  spiralis  L.  (in  herb.),  which  was  later  published  by  Dumortier. 

About  25  species  have  been  proposed  in  Ruppia  by  taxonomists  and  almost 
all  of  them  have  been  reduced  to  varieties  or  forms  of  R.  maritima.  The  chief 
troubles  have  concerned  themselves  in  the  first  place  with  the  lack  of  realiza- 
tion among  both  earlier  and  later  writers  of  the  fact  that  Ruppia  is  dichog- 
amous  (see  Ascherson,  Bot.  Zeit.,  1871,  p,  28,  and  in  Engler  u.  Prantl,  Natl. 
Pfl.  Fam.,  II,  1 :199,  1889),  with  proterandry ;  in  the  second  place,  with  the 
confusion  of  the  types  of  peduncles,  and  nature  of  pollination;  in  the  third 
place,  the  ecophenic  variable  as  to  length  of  both  peduncle  and  podogyne,  even 
in  the  same  plant ;  in  the  fourth  place,  varying  persistence  and  robustness  of 
the  vegetative  parts;  in  the  fifth  place,  the  variability  in  size  and  relative 
obliquity  in  the  maturing  fruits,  the  varying  behavior  of  stigma  and  rostrum; 
and  finally  in  the  relation  of  robust  and  slim  fruits  so  commonly  found  in  the 
same  inflorescence.  The  tendencies  to  recognize  only  one  species,  two  species, 
or  several  species  of  the  genus  have  been  influenced  by  these  considerations. 

After  Linnaeus,  botanists  seemingly  regarded  both  series  of  forms  as  states 
or  stages  of  one  another.  Botanists,  such  as  C.  A.  Agardh  (1823) ,  W.  D.  J.  Koch 
(1823, 1824),  and  others,  began  to  split  the  Linnaean  species,  regarding,  how- 
ever, the  spiralis  form  as  the  type.  Dumortier  (in  1827),  to  avoid  confusion, 
published  the  name  Ruppia  spiralis.  From  then  on  some  authors  have  applied 
this  name  to  the  spiral,  peduncled  form,  but  the  majority  have  insisted,  in 
spite  of  the  evidence  of  the  record,  that  this  is  the  type  of  R.  maritima  L.,  and 
that  the  short,  peduncled  var.  rostrata  C.  Ag.  (R.  rostellata  Koch)  is  a  variety 
of  it.  Fernald  and  "Wiegand  (Khodora,  16,  1914,  p.  121)  adopt  this  interpre- 
tation of  R.  maritima.  Hagstrom  (loc.  cit.,  1911,  pp.  137,  138)  recognizes  the 
true  R.  maritima  as  distinct  from  the  true  R.  spiralis. 

Ascherson  and  Graebner  (Synopsis,  1907,  pp.  142-145)  recognize  one 
species,  Ruppia  maritima  L.,  consisting  of  two  subspecies,  A.  spiralis  L.  Herb., 
Dumort.  and  B.  rostellata  Koch. 

As  a  result  of  our  studies,  it  seems  best  to  recognize  two  species,  Ruppia 
maritima  L.  (1763)  with  short,  nonspiral  peduncles  (the  R.  maritima  var. 
rostrata  of  my  earlier,  1924,  preliminary  account),  and  R.  spiralis  Dumort. 
(1827)  with  elongated,  spiral  peduncles  (the  R.  longipes  Fernald  and  Wie- 
gand, non  Hagstrom). 

These  conclusions  were  arrived  at  by  an  extensive  study  undertaken  by 
myself,  with  the  assistance  of  several  graduate  students,  of  the  representatives 
of  the  genus  as  they  occur  in  nature  in  the  marshes  around  San  Francisco 
Ba3%  and  by  culturing  plants  from  these  and  many  other  localities. 

Our  cultures  were  established  in  beakers  of  3  liter  capacity.  Into  each 
beaker  was  placed  about  100  grams  of  garden  soil  and  2.5  liters  of  water  from 
the  taps  of  the  Berkeley  water  supply.  In  such  cultures  cuttings  of  rhizomes 
or  seeds  were  planted.  The  beaker  was  covered  with  ordinary  glass  to  prevent 
evaporation,  and  the  plants  grew  and  flourished,  some  of  them  for  several 


472  CALIFORNIA  ACADEMY  OF  SCIENCES  [Proc.  4th  Ser. 

years  -without  change  of  water  or  soil.  Fresh  tap  water  had  to  be  added  from 
time  to  time,  in  small  quantities,  to  make  up  for  the  slight  evaporation.  A  few 
cultures  were  made  with  natural  spring  water,  with  distilled  water,  and  with 
potassium  nitrate  or  magnesium  sulphate  added.  In  the  two  latter  eases  the 
plants  grew  without  soil.  Under  favorable  conditions,  cultures  from  seed  pro- 
duced seeds  in  6()-80  days. 

Buppia  plants,  of  both  types  of  peduncle,  were  studied  for  about  15  years, 
both  from  certain  localities  more  or  less  readily  accessible  and  in  cultures. 
The  gross  morpliology  of  both  types  of  Buppia  was  studied  in  all  stages  from 
germination  to  fruit  production,  particular  attention  being  paid  to  such  un- 
certain matters  as  possession  or  absence  of  a  primary  root,  types  of  peduncles, 
variations  in  fruit  shape,  and  varying  lengths  of  peduncles  and  podogynes.  The 
definite  discontinuity  of  the  two  types  of  peduncle  is  maintained.  The  ever- 
varying  characters  of  fruits,  podogynes,  and  peduncles  may  be  expressed 
very  differently  on  the  same  individual. 

From  the  foregoing  studies  it  is  concluded  that  out  of  the  many  described 
species,  there  are  two  valid  species  in  the  genus,  and  that  the  various  species, 
subspecies,  varieties,  proles,  and  forms  which  have  been  described  are  only 
reversible  ecophenic,  probably  quantitative  expressions  of  form  and  size.  Per- 
sistence in  the  habitat  is  due  to  a  continuation  of  conditions  favorable  to 
development.  Buppia  plants  are  delicate  in  response  to  habitat  and  store  little 
starch  or  other  nutrient  materials.  In  culture  they  live  on  indefinitely,  but  in 
their  native  habitats  they  may  be  short  seasonal  plants  or  longer  or  shorter 
lived  perennials. 

My  sincerest  thanks  are  due  the  Linnaean  Society  of  London  for  permis- 
sion to  study  and  photograph  the  specimens  of  Buppia  in  the  Herbarium  of 
Linnaeus. 


PLATES 


PLATE  47 
Specimen  of  Euppia  maritima  L.  Herb.  Linn.,  Genus  175.  Sheet  1. 


[474] 


PROC.  CALIF.  ACAD.  SCI..  4TH  SERIES.  VOL.  XXV.  NO.  18 


[SETCHELL]  PLATE  47 


[475] 


PLATE  48 
Specimen  of  Ruppia  maritima  L.  Herb.  Linn.,  Genus  175.  Sheet  2. 


[476  J 


1« 

i 


PROC.  CALIF.  ACAD.  SCI..  4TH  SERIES.  VOL.  XXV.  NO.    18 


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INDEX  TO  VOLUME  XXV 


FOURTH  SERIES 
New  names  in  bold-face  type 


abrupta,  Fuchsia,  50 
acanthocarpa,  Opuntia,  246 

var,  Thornberi,  Opuntia,  247 
acanthodes,  Echinocactus,  261 
adenophora,  Phacelia,  360 
var.  affinis,  Fuchsia  regia,  14 
var.  aggregata,  Mammillaria  vivipara,  262 
ssp.  albula,  Artemisia  ludoviciana,  446 
var.  alpestris.  Fuchsia  regia,  14 
Andrei,  Fuchsia,  61 
apetala.  Fuchsia,  80 
var.  aprica.  Fuchsia  microphylla,  95 
arborescens.  Fuchsia,  84 

forma  parva,  Fuchsia,  86 

forma  tenuis,  Fuchsia,  86 

forma  typica,  Fuchsia,  85 
Arctostaphylos  glauca,  226 

pungens,  227 
arenarioides,  Drymaria,  201 
Arentsii,  Nicotiana,  297 
var.  arizonica,  Mammillaria  vivipara,  263 
arizonicus,  Bromus,  309 
Artemisia  Carruthii,  438 

Douglasiana,  458 

Lindleyana,  454 

longif  olia,  435 

ludoviciana,  440 

ludoviciana  ssp.  albula,  446 

ludoviciana  ssp.  candicans,  447 

ludoviciana  ssp.  incompta,  450 

ludoviciana  ssp.  mexicana,  452 

ludoviciana  ssp.  redolens,  454 

ludoviciana  ssp.  sulcata,  444 

ludoviciana  ssp.  typica,  440 

Michauxiana,  433 

Prescottiana,  460 

serrata,  437 

Suksdorfii,  457 

Tilesii,  460 

Tilesii  ssp.  typica,  460 

Tilesii  ssp.  unalaschcensis,  462 

vulgaris,  432 

vulgaris  Complex  in  North  America,  421 
Arthu-Schottii,  Astragalus,  158 
asperif  olia,  Fuchsia,  65 
Aspiazui,  Fuchsia,  43 
Asplundii,  Fuchsia,  37 


Astragalorum  Alter,  Pugillus,  147 
Astragalus  Arthu-Schottii,  158 

Bryantii,  156 

chamaemeniscus,  158 

diversif  olius,  148 

ensiformis  var.  gracilior,  158 

ensif  ormis  var.  typicus,  157 

eremiticus  var.  spencianus,  155 

eremiticus  var.  typicus,  155 

lentiginosus  var.  caesariatus,  161 

marianus,  150 

musimonum,  150 

oophorus,  152 

oophorus  var.  caulescens,  154 

oophorus  var.  typicus,  154 

panguicensis,  149 

Pattersonii,  157 

pinonis,  152 

sabinarum,  150 

straturensis,  155 

striatiflorus,  162,  163 

uncialis,  150 

Whitedii  forma  speirocarpoides,  148 

zionis,  149 
Atriplex  lentif  ormis,  226 
austromontana,  Fuchsia,  22 
ayavacensis.  Fuchsia,  32 
Babcock,  Ernest  B.,  Endemism  in  Crepis, 

269 
baeillaris.  Fuchsia,  92 
Barneby,  E.  C,  Pugillus  Astragalorum 

Alter,  147 
Benson,  Lyman,  Kevision  of  some  Arizona 

Cactaceae,  245 
var.  bolivarensis,  Fuchsia  petiolaris,  35 
boliviana  var.  luxurians.  Fuchsia,  54 

forma  puberulenta.  Fuchsia,  54 

var.  tsrpica,  Fuchsia,  53 
var.  Bonkerae,  Echinocereus  Fendleri,  260 
var.  Boyce-Thompsonii,  Echinocereus  Fend- 
leri, 260 
Bracelinae,  Fuchsia,  7 
Bromus  arizonicus,  309 
Bryantii,  Astragalus,  156 
Cactaceae,  Eevision  of  Arizona,  245 
var.  caesariatus,  Astragalus  lentiginosus, 
161 


[479] 


480 


CALIFORNIA  ACADEMY  OF  SCIENCES 


[Proc.  4th  Ser. 


californica,  Juniperus,  225 
Campbell,  Douglas   H.,  Eelations   of  the 
Temperate  Floras  of  North  and  South 
America,  139 
Campos-Portoi,  Fuchsia,  7 
ssp.  candicans,  Artemisia  ludoviciana,  447 
canescens,  Fuchsia,  26 
carinata,  Drymaria,  196 
carinata  var.  perennls,  Drymaria,  197 
Carruthii,  Artemisia,  438 
var.  caulescens,  Astragalus  oophorus,  154 
Ceanothus,  Field  Hybrids  in,  323 
cestroides,  Fuchsia,  83 
chamaemeniscus.  Astragalus,  158 
coccinea.  Fuchsia,  11 
Colensoi,  Fuchsia,  72 
colimae,  Fuchsia,  90 
compressa,  Opuntia,  250 

var.  microsperma,  Opuntia,  250 

var.  macrorhiza,  Opuntia,  251 
conf  ertif  olia.  Fuchsia,  42 
cordif  olia.  Fuchsia,  21 
corymbiflora.  Fuchsia,  49 
crassif  olia,  Drymaria,  195 
Crepis,  Endemism  in,  269 
Cuatrecasasii,  Fuchsia,  51 
cylindracea,  Fuchsia,  99 
cyrtandroides.  Fuchsia,  72 
Cytogenetics  of  Hybrids  in  Bromus,  307 
Datura  and  Matthew's  Hypothesis,  235 
debilis,  Drymaria,  198 
decidua,  Fuchsia,  75 
var.    decumbens,    Echinocereus    Engelman- 

nii,  258 
decussata.  Fuchsia,  58 
denticulata,  Fuchsia,  23 
depressa,  Drymaria,  205 
var.  deserti,  Mammillaria  vivipara,  263 
var.  diffusa,  Drymaria  polystachya,  198 
diversif  olius.  Astragalus,  148 
Douglasiana,  Artemisia,  458 
Drymaria,  192 
Drymaria,  The  Genus,  189 

arenarioides,  201 

carinata,  196 

carinata  var.  perennis,  197 

crassifolia,  195 

debilis,  198 

depressa,  205 

effusa,  204 

Fendleri,  199 

holosteoides,  194 

Johnstonii,  203 


pachyphylla,  195 

peninsularis,  202 

polystachya,  197 

polystachya  var.  diffusa,  198 

sperguloides,  200 

tenella,  204 

tenella  var.  nodosa,  205 

viscosa,  200 
Eastwoodae,  Gunnera,  391 
Echinocactus  acanthodes,  261 

polycephalus  var.  xeranthemoides,  262 
Echinocereus  Engelmannii,  257 

Engelmannii  var.  decumbens,  258 

Engelmannii  var.  Nicholii,  258 

Fendleri,  258 

Fendleri  var.  Bonkerae,  260 

Fendleri  var.  Boyce-Thompsonii,  260 

Fendleri  var.  rectispinus,  259 

Fendleri  var.  robustus,  259 

Ledingii,  261 

pectinatus,  255 

pectinatus  var.  neomexicanus,  256 

pectinatus  var.  rigidissimus,  256 

trigloehidiatus,  252 

triglochidiatus  var.  gonacanthus,  253 

trigloehidiatus  var.  polyacanthus,  253 

triglochidiatus  var.  mojavensis,  255 

triglochidiatus  var.  melanacan thus,  254 
Echinocystis  f  abacea,  227 
effusa,  Drymaria,  204 
Encliandra,  87 
Encliandra,  Fuchsia,  98 
Endemism  in  Crepis,  269 
Endemism  in  the  Hawaiian  Flora,  and  a 
Eevision   of   the   Hawaiian   Species   of 
Gunnera  (Haloragidaceae),  377 
Engelmannii,  Echinocereus,  257 

var.  decumbens,  Echinocereus,  258 

var.  Nicholii,  Echinocereus,  258 
ensif  ormis  var.  gracilior,  Astragalus,  158 

var.  t3rpicus,  Astragalus,  157 
eremiticus  var.  spencianus.  Astragalus,  155 

var.  typicus,  Astragalus,  155 
erinacea,  Opuntia,  258 

var.  hystricina,  Opuntia,  249 

var.  rhodantha,  Opuntia,  249 

var.  ursina,  Opuntia,  249 
Eufuchsia,  15 

Ewan,  Joseph,  Perennial  Datura  and  Mat- 
thew's Hypothesis,  235 
fabacea,  Echinocystis,  227 
excorticata,  Fuchsia,  71 
Fendleri,  Drymaria,  199 


Vol.  XXV] 


INDEX 


481 


Fendleri,  Echinocereus,  258 

var.  Bonkerae,  Echinocereus,  260 

var.  Boyce-Thompsonii,  Echinocereus, 
260 

var.  rectispinus,  Echinocereus,  259 

var.  robustus,  Echinocerus,  259 
Field  Hybrids  in  the  Genus  Ceanothus,  323 
Fischeri,  Fuchsia,  45 
Fuchsia  abrupta,  50 

Andrei,  61 

apetala,  80 

arborescens,  84 

arborescens  forma  parva,  86 

arborescens  forma  tenuis,  86 

arborescens  forma  typica,  85 

asperifolia,  65 

Aspiazui,  43 

Asplundii,  37 

austromontana,  22 

ayavacensis,  32 

bacillaris,  92 

boliviana  var.  luxurians,  54 

boliviana  forma  pubemlenta,  54 

boliviana  var.  typica,  53 

Bracelinae,  7 

Campos-Portoi,  7 

canescens,  26 

cestroides,  83 

coccinea,  11 

Colensoi,  72 

colimae,  90 

conf  ertif  olia,  42 

cordif  olia,  21 

corymbiflora,  49 

Cuatrecasasii,  51 

cylindracea,  99 

cyrtandroides,  72 

decidua,  75 

decussata,  56 

denticulata,  23 

Encliandra,  98 

excorticata,  71 

Fischeri,  45 

f  ulgens,  55 

f urfuracea,  46 

Garleppiana,  82 

Gehrigeri,  41 

glaberrima,  66 

Hartwegii,  63 

Hemsleyana,  95 

hirtella,  47 

hirsuta,  81 

hybrida,  12 


hypoleuca,  57 

Jahnii,  40 

juntasensis,  77 

Killipii,  52 

Kirkii,  74 

Lehmannii,  61 

leptopoda,  24 

Llewelynii,  38 

loxensis,  28 

lycioides,  69 

macrantha,  83 

macrophylla,  66 

macrostigma,  30 

macrostigma  var.  longiflora,  30 

macrostigma  var.  typica,  31 

magdalenae,  25 

magellaniea,  8 

magellanica  var.  macrostema,  10 

magellaniea  var.  Molinae,  11 

magellanica  var.  typica,  9 

Mathewsii,  44 

membranacea,  78 

Mexiae,  101 

michoacanensis,  97 

microphylla,  93 

microphylla  var.  aprica,  95 

microphylla  var.  typica,  94 

minimiflora,  90 

minutiflora,  92 

minutiflora  var.  hidalgensis,  93 

minutiflora  var.  typica,  93 

Munzii,  46 

(Onagraceae)  a  revision  of  the  Genus,  1 

Osgoodii,  60 

ovalis,  64 

pallescens,  29 

perscandens,  73 

petiolaris,  34 

petiolaris  var.  bolivarensis,  35 

petiolaris  var.  typica,  35 

platypetala,  30 

pilosa,  65 

polyantha,  48 

Pringlei,  89 

Pringsheimii,  33 

procumbens,  73 

putumayensis,  62 

regia,  13 

regia  var.  affinis,  14 

regia  var.  alpestris,  14 

regia  var.  radicans,  14 

regia  var.  typica,  13 

Eevision  of  the  Genus,  1 


482 


CALIFORNIA  ACADEMY  OF  SCIENCES 


[Peoc.  4th  Ser. 


rivularis,  27 

salicifolia,  78 

sanctae-rosae,  59 

scabriuscula,  58 

sessilifolia,  67 

simplicicaulis,  41 

Skiitchiana,  91 

Smitbii,  36 

splendens,  20 

Storkii,  45 

striolata,  96 

sylvatica,  68 

tacanensis,  91 

tetradactyla,  100 

thymif  olia,  88 

tincta,  43 

^Townsendii,  29 

triphylla,  33 

tuberosa,  76 

tuberosa  var.  inflata,  77 

tuberosa  var.  typica,  76 

tunariensis,  79 

unduavensis,  81 

venusta,  38 

venusta  var.  huilensis,  39 

venusta  var.  typica,  39 

verrucosa,  58 

Woytkowskii,  25 
fulgens.  Fuchsia,  55 
f  urf  uracea.  Fuchsia,  46 
Ganderi,  Quercus,  178 
Garleppiana,  Fuchsia,  82 
Gehrigeri,  Fuchsia,  41 
Genus  Drymaria  in,  and  Adjacent  to,  the 

Sonoran  Desert,  189 
glaberrima.  Fuchsia,  66 
glaberrima,  Phacelia,  367 
glauca,  Arctostaphylos,  226 
var.    gonacanthus,    Echinocereus    triglochi- 

diatus,  253 
Goodspeed,  T.  H.,  Nicotiana  Arentsii,  291 
var.  gracilior,  Astragalus  ensiformis,  158 
Gunnera  Eastwoodae,  391 

Hawaiian  Species  of,  380 

kaalaensis,  392 

kauaiensis,  395 

makahaensis,  394 

mauiensis,  389 

molokaiensis,  388 

petaloidea,  387 
Harlan,  J.  R.,  et  al..  Cytogenetics  of  Hy- 
brids in  Bromus,  307 
Hartwegii,  Fuchsia,  63 


Hawaiian  Flora,  Endemism  in,  377 

Hemsleyana,  Fuchsia,  95 

Hemsleyella,  74 

Heyderi,  Mammillaria,  264 

var.  MacDougalii,  Mammillaria,  265 

var.  hidalgensis,  Fuchsia  minutiflora,  93 

hirsuta.  Fuchsia,  81 

hirtella.  Fuchsia,  47 

holosteoides,  Drymaria,  194 

Howell,  John  Thomas,  Revision  of  Phacelia 
Sec.  Miltitzia,  357 

var.  hnilensis,  Fuchsia  venusta,  39 

Hybrid  Oak  from  San  Diego  County,  Cali- 
fornia, 177 

hybrida.  Fuchsia,  12 

hypoleuca.  Fuchsia,  57 

var.  hystricina,  Opuntia  erinacea,  249 

ssp.  incompta,  Artemisia  ludoviciana,  450 

var.  inflata.  Fuchsia  tuberosa,  77 

inundata,  Phacelia,  365 

inyoensis,  Phacelia,  372 

Jahnii,  Fuchsia,  40 

Johnstonii,  Drymaria,  203 

Juniperus  calif  ornica,  225 

juntasensis.  Fuchsia,  77 

kaalaensis,  Gunnera,  392 

kauaiensis,  Gunnera,  395 

Keck,  David  D.,  Revision  of  the  Artemisia 
vulgaris  complex  in  North  America,  421 

Kierschlegeria,  69 

Killipii,  Fuchsia,  52 

Kirkii,  Fuchsia,  74 

var.  Kunzei,  Opuntia  Stanlyi,  248 

Ledingii,  Echinocereus,  261 

Lehmannii,  Fuchsia,  61 

lentif  ormis,  Atriplex,  226 

lentiginosus  var.  caesariatus,  Astragalus, 
161 

leptopoda.  Fuchsia,  24 

Lindleyana,  Artemisia,  454 

Llewelynii,  Fuchsia,  38 

var.  longiflora.  Fuchsia  macrostigma,  31 

longif  olia,  Artemisia,  435 

loxensis,  Fuchsia,  28 

ludoviciana,  Artemisia,  440 
ssp.  albula,  Artemisia,  446 
ssp.  candicans,  Artemisia,  447 
ssp.  incompta,  Artemisia,  450 
ssp.  mexicana,  Artemisia,  452 
ssp.  redolens,  Artemisia,  454 
ssp.  sulcata,  Artemisia,  444 
ssp.  typica,  Artemisia,  440 


Vol.  XXV] 


INDEX 


483 


lutea,  Phacelia,  363 

var.  purpurascens,  Phacelia,  365 

var.  typica,  Phacelia,  364 
var.  luxurians,  Fuchsia  boliviana,  54 
lycioides,  Fuchsia,  69 
var.    MacDougalii,    Mammillaria    Heyderi, 

265 
McMinn,  H.  E.,  Field  Hybrids  in  the  Genus 

Ceanothus,  323 
macrantha,  Fuchsia,  83 
macrophylla,  Fuchsia,  66 
var.  macrorhiza,  Opuntia  compressa,  251 
var.  macrostema,  Fuchsia  magellanica,  10 
macrostigma,  Fuschia,  30 

var.  longiflora,  Fuchsia,  31 

var.  typica,  Fuchsia,  31 
magdalenae,  Fuchsia,  25 
magellanica.  Fuchsia,  8 

var.  Molinae,  Fuchsia,  11 

var.  macrostema.  Fuchsia,  10 

var.  tjrpica,  Fuchsia,  9 
makahaensis,  Gunnera,  394 
Mammillaria  Heyderi,  264 

Heyderi  var.  MacDougalii,  265 

vivipara  var.  aggregata,  262 

vivipara  var.  arizonica,  263 

vivipara  var.  deserti,  263 
marianus.  Astragalus,  150 
Marine  Algae  of  the  Monterey  Peninsula, 

171 
Mason,  Herbert  L.,  Pleistocene  Flora  Mc- 

Kittrick  Deposits,  California,  221 
Mathewsii,  Fuchsia,  44 
mauiensis,  Gunnera,  389 
var.  melanacanthus,  Echinocereus  triglo- 

chidiatus,  254 
membranacea.  Fuchsia,  78 
Mexiae,  Fuchsia,  101 

ssp.  mexicana,  Artemisia  ludoviciana,  452 
Michauxiana,  Artemisia,  433 
michoacanensis,  Fuchsia,  97 
microphylla,  Fuchsia,  93 

var.  aprica,  Fuchsia,  95 

var.  typica,  Fuchsia,  94 
var.  microsperma,  Opuntia  compressa,  250 
minimiflora.  Fuchsia,  90 
minutiflora  var.  hidalgensis,  Fuchsia,  93 

var.  tsrpica,  Fuchsia,  93 
var.   mojavensis,   Echinocereus   triglochi- 

diatus,  255 
molokaiensis,  Gunnera,  388 
monophylla,  Pinus,  225 
Munzii,  Fuchsia,  46 


Munz,   Philip   A.,   Eevision   of   the  Genus 

Fuchsia  (Onagraceae),  1 
musimonum,  Astragalus,  150 
var.    neomexicanus,    Echinocereus   pectina- 

tus,  256 
var.  Nicholii,  Echinocereus  Engelmannii, 

258 
Nicotiana  Arentsii,  297 
var.  nodosa,  Drymaria  tenella,  205 
oophorus,  Astragalus,  152 

var.  cauleseens.  Astragalus,  154 

var.  typicus,  Astragalus,  154 
Opuntia  acanthocarpa,  246 

acanthocarpa  var.  Thornberi,  247 

compressa,  250 

compressa  var.  macrorhiza,  251 

compressa  var.  microsperma,  250 

erinaeea,  248 

erinacea  var.  hystricina,  249 

erinaeea  var.  rhodantha,  249 

erinacea  var.  ursina,  249 

Stanlyi,  247 

Stanlyi  var.  Kunzei,  248 

Stanlyi  var.  Wrightiana,  248 

tortispina,  251 
Osgoodii,  Fuchsia,  60 
ovaHs,  Fuchsia,  64 
pachyphylla,  Drymaria,  195 
pallescens.  Fuchsia,  29 
panguicensis.  Astragalus,  149 
forma  parva,  Fuchsia  arborescens,  86 
Pattersonii,  Astragalus,  157 
pectinatus,  Echinocereus,  255 

var.  neomexicanus,  Echinocereus,  256 

var.  rigidissimus,  Echinocereus,  256 
Peirce,  George  J.,  Water  and  Plant  Anat- 
omy, 215 
peninsularis,  Drymaria,  202 
Perennial  Datura  and  Validity  of  Matthew's 

Hypothesis,  235 
var.  perennis,  Drymaria  carinata,  197 
perscandens.  Fuchsia,  73 
petaloidea,  Gunnera,  387 
petiolaris,  Fuchsia,  34 

var.  bolivarensis,  Fuchsia,  35 

var.  typica,  Fuchsia,  35 
Phacelia  adenophora,  360 

glaberrima,  367 

inundata,  365 

inyoensis,  372 

lutea,  363 

lutea  var.  purpurascens,  365 

lutea  var.  typica,  364 


484 


CALIFORNIA  ACADEMY  OF  SCIENCES 


[Proc.  4th  See. 


salina,  373 

scopulina,  368 

Sec.  Miltitzia,  Revision  of,  357 

submutica,  370 

tetramera,  374 
pilosa,  Fuchsia,  65 
pinonis.  Astragalus,  152 
Pinus  monophylla,  225 
platypetala,  Fuchsia,  30 
Pleistocene  Flora  McKittrick  Deposits, 

California,  221 
var.  polyacanthus,  Echinocereus  triglochi- 

diatus,  253 
polyantha,  Fuchsia,  48 
polycephalus  var.  xeranthemoides,  Echino- 

cactus,  262 
polystachya,  Drymaria,  197 

var.  diffusa,  Drymaria,  198 
Prescottiana,  Artemisia,  460 
Pringlei,  Fuchsia,  89 
Pringsheimii,  Fuchsia,  33 
procumbens.  Fuchsia,  73 
forma  puberulenta,  Fuchsia  boliviana,  54 
Pugillus  Astragalorum  Alter,  147 
pungens,  Arctostaphylos,  227 
var.  purpurascens,  Phacelia  lutea,  365 
putumayensis,  Fuchsia,  62 
Quercus  Ganderi,  178 
Quelusia,  6 

var.  radicans.  Fuchsia  regia,  14 
var.  rectispinus,  Echinocereus  Fendleri,  259 
ssp.  redolens,  Artemisia  ludoviciana,  454 
regia,  Fuchsia,  13 

var.  affinis,  Fuchsia,  14 

var.  alpestris.  Fuchsia,  14 

var.  radicans.  Fuchsia,  14 

var.  typica,  Fuchsia,  13 
Relations  of  the  Temperate  Floras  of  North 

and  South  America,  139 
Revision  of  some  Arizona  Cactaceae,  245 
Revision  of  the  Genus  Fuchsia  (Onagra- 

ceae),  1 
var.  rhodantha,  Opuntia  erinacea,  249 
var.  rigidissimus,  Echinocereus  pectinatus, 

256 
rivularis,  Fuchsia,  27 
var.  robustus,  Echinocereus  Fendleri,  259 
Ruppia,  469 

sabinarum.  Astragalus,  150 
St.  John,  Harold,  Endemism  in  the  Hawaiian 
Flora,  and  a  Revision  of  the  Hawaiian 
Species  of  Gunnera(Haloragidaceae),377 
salicif  olia,  Fuchsia,  78 


salina,  Phacelia,  373 

sanctae-rosae,  Fuchsia,  59 

scabriuscula.  Fuchsia,  58 

Schufia,  84 

scopulina,  Phacelia,  368 

serrata,  Artemisia,  437 

sessilifolia.  Fuchsia,  67 

Setchell,  Wm.  A.,  The  Genus  Ruppia,  469 

simplicicaulis.  Fuchsia,  41 

Skinnera,  70 

Skutchiana,  Fuchsia,  91 

Smith,  Gilbert  M.,  Sublittoral  Marine  Algae 
of  the  Monterey  Peninsula,  171 

Smithii,  Fuchsia,  36 

forma  speirocarpoides,  Astragalus  Whitedii, 
148 

var.  spencianus.  Astragalus  eremiticus,  155 

sperguloides,  Drymaria,  200 

splendens,  Fuchsia,  20 

Stanly i,  Opuntia,  247 

var.  Kunzei,  Opuntia,  248 
var.  Wrightiana,  Opuntia,  248 

Stebbins,  G.  L.,  Jr.,  et  al..  Cytogenetics  of 
Hybrids  in  Bromus,  307 

Storkii,  Fuchsia,  45 

striatiflorus.  Astragalus,  162,  163 

straturensis,  Astragalus,  155 

striolata,  Fuchsia,  96 

submutica,  Phacelia,  370 

Suksdorfii,  Artemisia,  457 

ssp.  sulcata,  Artemisia  ludoviciana,  444 

sylvatica.  Fuchsia,  68 

tacanensis.  Fuchsia,  91 

Temperate    Floras    of    North    and    South 
America,  139 

tenella,  Drymaria,  204 

var.  nodosa,  Drymaria,  205 

forma  tenuis,  Fuchsia  arborescens,  86 

tetradactyla,  Fuchsia,  100 

tetramera,  Phacelia,  374 

var.  Thornberi,  Opuntia  acanthocarpa,  247 

thymifolia.  Fuchsia,  88 

Tilesii,  Artemisia,  460 

ssp.  typica,  Artemisia,  460 

ssp.  unalaschcensis,  Artemisia,  462 

tincta.  Fuchsia,  43 

Tobgy,  H.  A.,  et  al..  Cytogenetics  of  Hy- 
brids in  Bromus,  307 

tortispina,  Opuntia,  251 

Townsendii,  Fuchsia,  29 

triglochidiatus,  Echinocereus,  252 

var.  gonacanthus,  Echinocereus,  253 
var.  melanacanthus,  Echinocereus,  254 


Vol.  XXV] 


INDEX 


485 


var.  mojavensis,  Echinocereus,  255 

var.  polycantlius,  Echinocereus,  253 
tripliylla,  Fuchsia,  33 
tuberosa,  Fuchsia,  76 

var.  inflata,  Fuchsia,  77 

var.  typica,  Fuchsia,  76 
tunariensis.  Fuchsia,  79 
ssp.  typica,  Artemisia  ludoviciana,  440 
ssp.  typica,  Artemisia  Tilesii,  460 
var.  typicus,  Astragalus  ensif  ormis,  157 
var.  typicus,  Astragalus  eremiticus,  155 
var,  typicus,  Astragalus  oophorus,  154 
forma  typica,  Fuchsia  arborescens,  85 
var.  typica,  Fuchsia  boliviana,  53 
var.  typica,  Fuchsia  macrostigma,  31 
var.  tjrpica,  Fuchsia  magellanica,  9 
var.  typica,  Fuchsia  microphylla,  94 
var.  typica.  Fuchsia  minutiflora,  93 
var.  typica,  Fuchsia  petiolaris,  35 
var.  typica.  Fuchsia  regia,  13 
var.  typica,  Fuchsia  tuberosa,  76 
var.  typica,  Fuchsia  venusta,  39 
var.  typica,  Phacelia  lutea,  364 
ssp.  unalaschcensis,  Artemisia 
Tilesii,  462 


uncialis,  Astragalus,  150 
unduavensis,  Fuchsia,  81 
var.  ursina,  Opuntia  erinacea,  249 
venusta,  Fuchsia,  38 

var.  huilensis,  Fuchsia,  39 

var.  tjrpica,  Fuchsia,  39 
verrucosa.  Fuchsia,  58 
viscosa,  Drymaria,  200 
vivipara  var.  aggregata,  Mammillaria,  262 

var.  arizonica,  Mammillaria,  263 

var.  deserti,  Mammillaria,  263 
vulgaris,  Artemisia,  432 
Water  and  Plant  Anatomy,  215 
Whitedii,  forma  speirocarpoides,  Astraga- 
lus, 148 
Wiggins,  Ira  L.,  The  Genus  Drymaria  in, 
and  adjacent  to,  the  Sonoran  Desert,  189 
Wolf,  Carl  B.,  The  Gander  Oak,  a  New 
Hybrid  Oak  from   San  Diego   County, 
California,  177 
var.  Wrightiana,  Opuntia  Stanlyi,  248 
Woytkowskii,  Fuchsia,  25 
var.  xeranthemoides,  Echinocactus  poly- 

cephalus,  262 
zionis,  Astragalus,  149 


a      --■ 


PROCEEDINGS 


OF  THE 


California  Academy  of  Sciences 

FOURTH    SERIES 

Vol.  XXV,  No.  i,  pp.  1-138,  pis.  1-16  December  1 1,  1943 


.ALICE  EASTWOOD 
SEMI-CENTENNIAL  PUBLICATIONS 

No.  1 

A  REVISION  OF  THE  GENUS  FUCHSIA 

(ONAGRACEAE) 

Pn 


By  PHILIP  A.  MUNZ     /l^^^l:^^/^ 

wfessor  of  Botany,  Pomona  Colle^i^  /,0  ^  <%  ^  ^^ 


., »i 

^  \  L^. 


( L I  e  2^  A  R  Y  ) 


PRINTED  FROM  THE  JOHN  W.  HENDRIE  PUBLICATION  ENDOWMENT 

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'  PUBLISHED    BY  THE    ACADEMY       :        1943 


TRUSTEES,  OFFICERS,  COUNCIL,  AND  MUSEUM  STAFF 
OF  THE  CALIFORNIA  ACADEMY  OF  SCIENCES 


BOARD  OF  TRUSTEES 

NORMAN  B.  LIVERMORE,  President Term  expires  1945 

BRUCE  CORxNWALL,  Vice-President Term  expires  1946 

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CHARLES  PAGE Term  expires  1947 

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OFFICERS  AND  COUNCIL 

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SCIENTIFIC  STAFF 
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Department  of  Botany Alice  Eastwood,  Curator 

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Department  of  Entomology E.  C.  Van  Dyke,  Honorary  Curator 

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Department  of  Exhibits Frank  Tose,  Chief  ^ 

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and  Mammalogy Robert  T.  Orr,  Assistant  Curator 

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Department  of  Paleontolog)' G.  Dallas  Hanna,  Curator 

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PROCEEDINGS  OF  THE  CALIFORNIA  ACADEMY  OF  SCIENCES 

Fourth  Series 

Volume  XXIII 

1.  Cretaceous  Geology  of  Lower  California,  by  Frank  M.  Anderson  and  G.  Dallas 

Hanna.  Pp.  1-34,  pis.  1-1 1,  2  figs,  in  text.  December  1935 $0.75 

2.  A  List  of  the  Birds  of  the  Atlin  Region,  British  Columbia,  by  Harry  S.  Swarth. 

Pp-  35-58-  June  1936 25 

3.  Origins  of  the  Fauna  of  the  Sitkan  District,  Alaska,  by  Harry  S.  Swarth.  Pp.  59-78, 

1  fig.  in  text.  June  1936. 20 

4.  A  New  Central  American  Snake,  by  Joseph  R.  Slevin.  Pp.  79-81.  August  1936 20 

5.  The  Cranium  of  the  Miocene  Gannet  Moris  vagabundus  Wetmore,  by  Lawrence 

V.  Compton.  Pp.  83-84,  1  fig.  in  text.  August  1936 10 

6.  A  New  Member  of  the  Blenny  Family,  by  Alvin  Scale.  Pp.  85-86.  August  1936 10 

7.  The  Fishes  of  the  Atlantic  and  Pacific  Slopes  near  Cajamarca,  Peru,  by  Nathan  E. 

Pearson.  Pp.  87-98,  pis.  12-13,  1  fig.  in  text.  April  1937 .20 

8.  The  Fishes  of  the  Beni-Mamord  and  Paraguay  Basins,  and  a  Discussion  of  the  Origin 

of  the  Paraguayan  Fauna,  by  Nathan  E.  Pearson.  Pp.  99-114.  May  1937 .20 

9.  Mammals  of  Death  Valley,  by  Joseph  Grinnell.  Pp.  1 15-169.  August  1937 60 

10.  A  New  Marine  Turtle  from  the  Miocene  of  California,  by  Charles  W.  Gilmore. 

Pp.  171-174,  pi.  14.  December  1937 15 

11.  Contributions  to  Oriental  Herpetology.  V.  Honshu  or  Hondo,  the  Neighboring 

Islands  of  Sado  and  Awaji,  and  the  Seven  Islands  of  Idzu,  by  Joseph  R.  Slevin. 

Pp.  175-190.  December  1937 .10 

12.  Marine  MoUusca  of  San  Martin  Island,  Mexico,  by  A.  M.  Strong.  Pp.  191-194. 

December  1937 10 

13.  A  Record  of  the  Fossil  Grebe,  Colymbus  parvus,  from  the  Pliocene  of  California, 

with  Remarks  on  other  American  Fossils  of  this  Family,  by  Alexander  Wetmore. 

Pp.  195-201,  15  figs,  in  text.  December  1937 .15 

14.  New  Species  of  West  American  Shells,  by  A,  M.  Strong.  Pp.  203-216,  pis.  15-16. 

May  1938 35 

15.  Some  MoUusca  of  the  Families  Cerithiopsidae,  Cerithiidae  and  Cyclostrematidae 

from  the  Gulf  of  California  and  Adjacent  Waters,  by  Fred  Baker,  G.  D.  Hanna 

and  A.  M.  Strong.  Pp.  217-244,  pis.  17-23.  May  1938 70 

16.  Columbellidae  from  Western  Mexico,  by  Fred  Baker,  G.  D.  Hanna  and  A.  M.  Strong. 

Pp.  245-254,  pi.  24.  May  1938 25 

17.  On  Some  Birds  Rare  in,  or  Hitherto  Unrecorded  from  Chiriqui  Province,  Panama, 

by  M.  E.  McLellan  Davidson.  Pp.  255-261.  September  1938 15 

18.  Studies  in  the  Andrenidae  of  North  America^ — I,  by  E.  Gorton  Linsley.  Pp.  263-282. 

September  1938 25 

19.  Notes  on  the  Breeding  Seasons  of  the  Rocky  Beach  Fauna  of  Monterey  Bay,  Cali- 

fornia, by  Willis  G.  Hewatt.  Pp.  283-288.  September  1938 15 

20.  Review  of  the  Genus  Gastrodes,  by  Robert  L.  Usinger.  Pp.  289-301,  2  figs,  in  text. 

September  1938 20 

21.  A  New  Rodent  of  the  Genus  Nesoryzomys  from  the  Galapagos  Islands,  by  Robert  T. 

Orr.  Pp.  303-306,  pi.  25.  September  1938 15 

22.  Mammals  from  Sikang,  China,  by  Robert  T  Orr.  Pp.  307-310.  September  1938 10 

23.  Studies  on  the  Cladocera  of  Monterey  Bay,  by  Harriet  Marguerite  Baker.  Pp.  311- 

365,  pis.  26-31.  November  1938 70 

24.  Marine  Pleistocene  Mollusks  from  the  Galapagos  Islands,  by  Leo  George  Hertlein 

and  A.  M.  Strong.  Pp.  367-380.  July  1939 25 

25.  Notes  on  Some  Forms  of  Oreohelix  strigosa  (Gould),  by  G.  Dallas  Hanna  and  Allyn 

G.  Smith.  Pp.  381-392,  pis.  33-36.  December  1939 35 

26.  Notes  on  a  Collection  of  Reptiles  and  Amphibians  from  Guatemala.  I.  Snakes,  by 

Joseph  R.  Slevin.  Pp.  393-414,  pis.  37-38.  December  1939 35 

27.  A  New  Genus  and  Species  of  Marine  Ostracods  from  South  Georgia,  by  Tage  Skogs- 

berg.  Pp.  415-425,  13  figs,  in  text.  December  1939 25 

28.  The  Bees  of  the  Southern  California  Islands,  by  T.  D.  A.  Cockerell.  Pp.  427-436. 

December  1939 15 

29.  The  Neotropical  Anchovies  of  the  Genus  Amplova,  by  George  Sprague  Myers.  Pp. 

437-442.  December  1940 .15 

30.  Notes  on  a  California  Earthworm,  Plutellus  papillifer  (Eisen,  1893),  by  Gordon  E. 

Gates.  Pp.  443-452,  4  figs,  in  text.  September  1941 20 

31.  Notes  on  a  collection  of  Reptiles  and  Amphibians  from  Guatemala.  II.  Lizards, 

by  Joseph  R.  Slevin.  Pp.  453-462.  June  1942 20 


PROCEEDINGS  OF  THE  CALIFORNIA  ACADEMY  OF  SCIENCES 
Fourth  Series,  Volume  XXIII  (Continued) 

32.  Notes  on  a  Collection  of  Reptiles  from  Boquete,  Panama,  with  the  Description  of 

a  new  Species  of  Hydromorphys,  by  Joseph  R.  Slevin.  Pp.  463-480,  pis.  39-42. 

June  1942  50 

33.  Redescription  of  three  Species  of  the  Polychaetous  Family  Polynoidae  from  Cali- 

fornia, by  Tage  Skogsberg.  Pp.  481-532,  pi.  43,  text  figs.  A-D.  July  1942 50 

34.  Reef  Corals  from  the  California  Middle  Eocene,  by  J.  Wyatt  Durham.  Pp.  503-510, 

pi.  44.  October  1942 30 

35.  Mammals  of  the  Clearwater  Mountains,  Idaho,  by  Robert  T.  Orr.  Pp.  511-536, 

pis.  45-47.  August  18,  1943 50 

36.  Mollusks  of  the  Clearwater  Mountains,  Idaho,  by  Allyn  G.  Smith.  Pp.  537-554,  pi. 

48.  August  18,  1943 35 

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PROCEEDINGS  OF  THE  CALIFORNIA  ACADEMY  OF  SCIENCES 

Fourth  Series 
Volume  XXIV 

Contributions  Toward  a  Knowledge  of  the  Insect  Fauna  of  Lower  California. 

1.  Introductory  Account,  by  A.  E.  Michelbacher  and  E.  S.  Ross.  Pp.  1-20,  pis.  1-3. 

February  1942  $0.25 

2.  Coleoptera:  Cerambycidae,  by  E.  Gorton  Linsley.  Pp.  21-96,  pis.  4-5.  February  1942        .75 

3.  Coleoptera:  Buprestidae,  by  Edwin  C.  Van  Dyke.  Pp.  97-132,  pis.  6-7.  March  1942        .35 

4.  Neuroptera:  Myrmeleonidae,  by  Nathan  Banks.  Pp.  133-152,  pi.  8.  March  1942 20 

5.  Symphyla,  by  A.  E.  Michelbacher.  Pp.  153-160,  pi.  9.  March  1942 15 

6.  Diptera:  Culicidae,  by  Thomas  H.  G.  Aitken.  Pp.  161-170.  June  1942 20 

7.  Coleoptera:  Tenebrionidae,  by  Frank  E.  Blaisdell,  Sr.  Pp.  171-288,  pis.  10,  1 1 1.50 

Volume  XXV 
Alice  Eastwood  Semi-Centennial  Publications 
i.A  Revision  of  the  Genus  Fuchsia  (Onagraceae),  by  Philip  A.  Munz.  Pp.  1-138, 

pis.  1-16.  December  11,  1943 $i-75 

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10.  The  Reptiles  of  Western  North  America;  an  Account  of  the  Species  known  to  in- 

habit California  and  Oregon,  Washington,  Idaho,  Utah,  Nevada,  Arizona,  British 
Columbia,  Sonora  and  Lower  California,  by  John  Van  Denburgh.  2  vols.,  1028  pp., 
128  pis.  November  1922 $10.50 

11.  Fauna  and  Stratigraphic  Relations  of  the  Tejon  Eocene  at  the  Type  Locality  in 

Kern  County,  California,  by  Frank  M.  Anderson  and  G.  Dallas  Hanna.  249  pp., 

16  pis.  March  1925 2.00 

12.  A  Review  of  the  Giant  Mackerel -like  Fishes,  Tunnies,  Spearfishes  and  Swordfishes, 

by  David  Starr  Jordan  and  Barton  Warren  Evermann.  1 13  pp.,  20  pis.  September 

1926 1.25 

13.  Cretaceous  Diatoms  from  California,  by  G.  Dallas  Hanna.  48  pp.,  5  pis.  September 

•927  •  •, 75 

14.  The  Rudistids  of  Southern  Mexico,  by  Robert  H.  Palmer.  132  pp.,  18  pis.  February 

1928 1.75 

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text.  August  1928 2.00 

16.  The  Amphibians  of  Western  North  America,  by  Joseph  R.  Slevin.  152  pp.,  23  pis. 

September  1928 3.00 

17.  Log  of  the  Schooner  Academy  on  a  Voyage  of  Scientific  Research  to  the  Galapagos 

Islands,  by  Joseph  R.  Slevin.  162  pp.;  17  pis.  February  1931 3.00 

18.  The  Avifauna  of  the  Galapagos  Islands,  by  Harry  S.  Swarth.  299  pp.,  1  map,  57  figs. 

in  text.  June  1931 3.00 

19.  The  Rabbits  of  California,  by  Robert  T.  Orr.  227  pp.,  10  pis.,  30  figs,  in  text.  May    < 

1940 3-50 

20.  Early  Naturalists  in  the  Far  West,  by  Roland  H.  Alden  and  John  D.  Ifft.  60  pp., 

1  map,  6  figs,  in  text.  April,  1943 1.00 

8 


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