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I       CiUntni  af  ^ 

^^^^^^ 

l^kn\}nsth  from 


MEMOIRS 

ToRREY  Botanical  Club- 
volume  VII 


THE 

HEPATICAE  AND  ANTHOCEROTES 

OF 

CALIFORNIA 


BY 

MARSHALL  AVERY  HOW^E 


NEW  YORK 
1899 


THE 


HEPATICAE  AND  ANTHOCEROTES 

OF 

CALIEORNIA 


BY 

MARSHALL  AVERY  HOWE 


Issued  August  5,  1899 


PRESS  OF 

The  new  Era  Printing  Company 
LANCASTER,  Pa. 


INTRODUCTION. 


The  aim  of  the  subjoined  work  is  to  present  a  descriptive  hst 
of  the  Hepaticae  and  Anthocerotes  of  Cahfornia,-  together  with 
delineations  of  such  species  as  have  been  hitherto  either  not  at  all 
or  only  quite  imperfectly  figured.  Most  of  the  material  upon 
which  the  paper  has  been  based  was  collected  by  the  author  dur- 
ing a  residence  of  about  five  years  in  California.  Of  the  Anthoc- 
erotes and  the  thalloid  Hepaticae,  specimens  obtained  in  various 
stages  of  development  were  fixed  with  chromic  acid  and  preserved 
in  alcohol  in  order  to  retain  better  the  natural  outlines  and  also  to 
permit  a  more  accurate  study  of  the  internal  structure  through  the 
use  of  microtome  sections.  Additional  material  has  been  contributed 
from  time  to  time  by  friends  and  associates,  among  whom  thanks 
are  due  especially  to  Miss  Edith  S.  Byxbee,  Dr.  Willis  L.  Jepson, 
Professor  L.  M.  Underwood,  Professor  Douglas  H.  Campbell, 
Professor  A.  J.  McClatchie,  Mr.  S.  B.  Parish,  Professor  Edward 
L.  Greene,  Mr.  M.  S.  Baker,  Mr.  F.  P.  Nutting,  Mr.  J.  Burtt 
Davy,  Mr.  W.  C.  Blasdale,  and  Professor  William  A.  Setchell. 
Specimens  secured  by  the  earlier  collectors,  notably  the  very  im- 
portant collections  of  Dr.  Henry  N.  Bolander,  have  likewise  been 
of  great  service.  Through  the  kindness  of  the  owners  or  custo- 
dians of  various  herbaria  in  Europe  and  America  we  have  enjoyed 
the  privilege  of  seeing,  with  two  or  three  exceptions,  the  original 
materials  of  all  the  distinctively  W^est  American  species  which  have 
been  recorded  as  occurring  within  our  limits. 

It  is  possible  that  the  first  botanist  to  collect  any  Hepaticae  in 
California  was  Archibald  Menzies,  who  visited  the  region  during  the 

Memoirs  of  the  Torrey  Botanical  Club,  Vol.  VII. 

(1) 


2  Hepaticae  and  Anthocerotes  of  California 


last  decade  of  the  eighteenth  century,  though  we  are  not  in  a  position 
to  afifirm  with  certainty  that  he  actually  gathered  any  of  these  plants 
within  the  limits  of  what  is  now  California,  inasmuch  as  the  only 
hepatic  that  we  have  seen,  definitely  known  to  be  of  Menzies'  col- 
lecting (the  original  of  Porella  navicularis),  is  referred  simply  to  the 
West  Coast  of  North  America,  and  the  species  is  one  that  he  may 
have  found  equally  well,  or  better,  farther  to  the  northward.  Dr. 
Thomas  Coulter,  about  four  decades  later,  seems  to  have  been  the 
discoverer  of  Asterella  vlolacea,  though  it  was  a  plant  of  Bolan- 
der's  which,  after  the  lapse  of  another  forty  years,  served  as  the 
basis  of  Mr.  Austin's  diagnosis.  Occasional  Hepaticae  were  col- 
lected within  the  state  from  1850  to  1880  by  Dr.  Albert  Kellogg, 
Dr.  J.  M.  Bigelow,  Professor  William  H.  Brewer,  Professor  John 
Torrey,  Dr.  C.  C.  Parry,  and  Dr.  Edward  Palmer.  It  was,  how- 
ever. Dr.  Henry  N.  Bolander,  who  did  much  more  than  all  the 
others  to  pave  the  way  to  a  knowledge  of  the  Californian  hepatics. 
Dr.  Bolander's  services  to  Californian  botany  are  too  well  known 
to  need  discussion  here.*  Suffice  it  to  say  that  he  was  a  resident 
of  Cahfornia  from  1861  to  1878,  and  that  while  giving  no  more 
attention  to  cryptogamous  plants  than  to  the  spermatophytes,  his 
eye  for  mosses,  liverworts,  and  lichens  was  astonishingly  keen. 
He  not  only  explored  with  much  thoroughness  the  San  Francisco 
Bay  region,  but  as  botanist  of  the  State  Geological  Survey,  and  later 
as  State  Superintendent  of  Schools,  was  enabled  to  visit  more  remote 
portions  of  the  state.  Fully  one  half  the  species  in  the  California 
hepatic  list  of  today  were  known  to  him.  Of  these,  at  least  twelve 
were  new  to  hepaticology  ;  seven  now  bear  his  name.  The  Hepati- 
cae of  Bolander's  collecting  were  studied  by  Mr.  Austin  and  by  Dr. 
Gottsche.  Professor  W.  G.  Farlow  visited  the  Pacific  Coast  in  1885 
and  collected  a  few  plants  of  this  group  at  various  points  in  Califor- 
nia. Professor  L.  M.  Underwood  also  visited  California  in  the  sum- 
mer of  1 888  and  made  collections  in  Alameda,  San  Francisco,  Marin, 
and  Santa  Cruz  counties.  Twelve  specimens  of  Hepaticae  were 
secured  by  Messrs.  Coville  and  Funston  on  the  Death  Valley  Ex- 
pedition in  the  summer  of  1891.  In  the  summer  of  1894,  Mr.  M. 
S.  Baker  and  Mr.  F.  P.  Nutting  made  a  botanical  journey  through 

*  An  interesting  paper  on  "  Dr.  Henry  N.  Bolander,  Botanical  Explorer,"  by  Dr. 
Willis  L.  Jepson,  has  appeared  in  Erythea,  6:  100-107.  1898. 


Introduction 


3 


portions  of  Shasta,  Lassen,  and  Modoc  counties,  including  in  their 
collections  several  liverworts.  Mr.  Samuel  B.  Parish  has  found 
several  species  of  Hepaticae  and  Anthocerotes  in  the  San  Bernar- 
dino Mts.,  and  Professor  A.  J.  McClatchie  lists  seventeen  species 
in  the  Flora  of  Pasadena  and  Vicinity."*  Professor  D.  H.  Camp- 
bell has  made  interesting  discoveries  in  the  neighborhood  of  Stan- 
ford University,  two  of  which  are  the  novelties  SpJiaerocarpiis  cris- 
tatus  and  Riccia  Campbelliana,  first  described  in  the  present  paper. 
Mrs.  Katharine  Brandegee  is  thus  far  the  only  collector  of  Pro- 
fessor Campbell's  new  genus  Geothalhis  and,  so  far  as  we  know,  is 
the  only  one  to  have  found  Reboulia  Jiemisphacrica  in  California. 
Our  own  collections  have  been  made  in  the  region  of  the  Bay  of  San 
Francisco,  and  in  Sonoma,  Mendocino,  Humboldt,  Trinity,  Shasta, 
and  Siskiyou  counties.  Although  the  hepatic  flora  of  the  coast 
counties  may  now  be  considered  to  be  fairly  well  known,  it  is  per- 
haps not  too  much  to  expect  that  explorations  of  the  future  will 
nearly  double  the  number  of  species  now  known  to  occur  in  the 
state  as  a  whole.  That  the  present  number — 86,  including  one 
well-marked  variety — is  a  very  respectable  one  for  the  area  involved 
may  be  readily  seen  by  a  comparison  with  the  hepatic  flora  of  the 
northern  and  eastern  United  States.  In  the  sixth  edition  of  Gray's 
Manual  of  the  Botany  of  the  Northern  United  States,  141  species 
are  recognized  by  Professor  Underwood  under  the  Hepaticae. 
Subsequent  additions  and  revisions  leave  the  total  about  1 50. 
When  it  is  borne  in  mind  that  the  area  covered  by  the  sixth  edi- 
tion of  Gray's  Manual  is  five  times  that  of  California,  the  86 
species  now  attributed  to  California  make  a  fair  showing  in  point 
of  numbers  as  against  the  150  of  the  northern  and  eastern  states. 
But  California  has  mountains  of  more  than  twice  the  height  of  any 
in  the  Gray's  Manual  region  and  in  spite  of  the  fact  that  the  an- 
nual rainfall  in  many  parts  of  the  state  is  less  than  is  requisite 
for  the  best  development  of  bryophytic  vegetation,  the  diversities 
of  altitude,  temperature,  and  humidity,  afforded  by  the  state  as  a 
whole,  combine  to  give  it  a  rich  and  varied  hepatic  flora,  and  one 
which  in  respect  to  the  numbers  of  its  species  may  be  expected  to 
compare  favorably  with  any  region  of  equal  area  in  the  United  States. 
The  whole  great  range  of  the  Sierra  Nevada  is  still  almost  a  terra 


*  Reid,  History  of  Pasadena,  624.  1895. 


4 


Hepaticae  and  Axthocerotes  of  California 


incognita  so  far  as  its  bryology  is  concerned.  The  mountains  of  this 
range  in  order  to  yield  the  best  results  should  be  explored  during 
the  spring  months,  while  the  snows  are  leaving  and  while  the 
ground  is  still  moist.  During  the  dry  season,  the  terrestrial  Hep- 
aticae become  shriveled  and  more  or  less  covered  with  dust,  and 
are  usually  difficult  to  detect.  Several  species  known  to  occur  in 
British  Columbia,  Washington,  and  Oregon  are  likely  to  be  found 
also  in  northern  California,  especially,  perhaps,  near  the  well- 
watered  coast.  The  more  arid  portions  of  the  state  will  doubt- 
less still  yield  Riccias  and  quite  possibly  new  hepatic  forms  with 
special  provisions  for  withstanding  a  season  of  drought,  such  as  are 
exhibited  by  GeotJiallns  tubcrosiis  and  AntJioccros  pJiymatodcs.  Ad- 
ditions to  the  state  list  are  to  be  expected  from  any  quarter,  but 
more  particularly,  we  think,  from  the  regions  indicated. 

On  the  following  pages  is  a  table  designed  to  give  some  idea 
of  the  geographical  range  of  the  Hepaticae  and  Anthocerotes  of 
California,  taking  into  consideration  only  the  hepaticologically  bet- 
ter known  portions  of  the  Northern  Hemisphere,  the  facts  of 
distribution  being  compiled  from  the  most  recent  and  trustworthy 
sources.  It  will  be  seen  that  of  our  86  species  (representing  36 
genera),  37  (43-1-^6)  occur  also  in  the  Gray's  Manual  Region  (/.  Cy 
the  northern,  eastern,  and  central  United  States,  extending  beyond 
the  Mississippi  River  to  the  looth  meridian  and  southward  to 
North  Carolina  and  Tennessee);  40  (46 -j- J/o)  in  the  British  Isles  ; 
46  (53^ — northern  and  central  Europe;  46(531^ — fo)  in 
the  Mediterranean  Region  of  Europe  ;  and  36(42  —  ^)  in  northern 
Asia  (including  Siberia,  China,  and  Japan);  37  (43  +  fo)  are  pecu- 
liar to  Pacific  Slope  of  America,  several  of  them  having  not  yet 
been  found  outside  of  California.  It  is  of  some  interest  to  remark 
that  the  number  of  species  peculiar  to  the  Pacific  Slope  is  propor- 
tionally greater  than  of  those  peculiar  to  the  Gray's  Manual  Re- 
gion, even  though  we  may  not  stop  to  prove  this  by  figures.  The 
difference  between  43^  and  53j^I/o  is  sufficient  to  justify  the  asser- 
tion that  the  hepatic  flora  of  California  has  more  in  common  with 
Europe  than  with  the  eastern  and  central  United  States.  But  facts 
involving  the  consideration  of  species  do  not  always  lend  them- 
selves with  grace  and  accuracy  to  mathematical  modes  of  expression. 
The  European  species  Porclla  laevigata,  Riccia  ciliata,2ir\d  AntJioccros 


Introduction 


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6  Hepaticae  and  Anthocerotes  of  California 


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Introduction 


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14 


8-  Hepaticae  and  Axthocerotes  of  California' 


dichotomiis  are  represented  on  the  Californian  coast  by  the  close 
analogues,  Porella  Rocllii,  Riccia  tricliocarpa  and  AntJwceros  phy- 
matodes,  their  nearest  relatives  in  the  eastern  United  States  being 
much  farther  removed  ;  again,  a  species  may  be  very  common  in 
Europe  and  very  rare  in  California  or,  vice  versa ;  and  facts  like 
these  are  wholly  lost  sight  of  in  merely  numerical  comparisons  of 
what  we  choose  to  regard  as  species,  A  few  special  considerations 
serv^e  to  emphasize  the  affinity  of  the  California  hepatic  flora  with 
that  of  Europe,  particularly  with  that  of  the  Mediterranean  region. 
We  have,  to  begin  with,  five  species  of  the  genus  Asterdla  in  Cal- 
ifornia and  the  same  number  in  Italy.  In  the  eastern  United  States 
there  is  but  one  representative  of  this  genus.  Ten  species  of  Riccia 
— a  genus  which  seems  to  find  its  best  development  in  southern 
Europe  and  northern  Africa — are  known  to  occur  in  California. 
Of  these,  one  cannot  be  distinguished  from  a  species  otherwise 
known  only  in  southern  France  and  Italy,  and  one  or  two  others 
are  closely  allied  to  species  peculiar  to  the  sam.e  regions.  Targi- 
onia  hypophylla,  too,  is  another  link  in  the  chain  that  binds  Cali- 
fornia to  the  warmer  parts  of  Europe.  This  genus  is  absent  in  the 
eastern  states,  but  what  seems  to  be  the  very  same  species  is  found 
sparingly  in  the  British  Isles  and  central  Europe,  is  more  common 
in  northern  Italy,  and  is  said  to  become  very  abundant  in  the  south- 
ern portion  of  the  peninsula.  One  rather  common  Californian 
hepatic  seems  to  be  identical  with  CepJialozia  Tiinicri — one  of  the 
rarest  of  the  Old  World  species,  occurring  in  limited  quantity  at 
a  few  localities  in  the  British  Isles,  France,  the  Canaries,  Corsica 
and  Algeria.  No  stations  for  this  plant  intermediate  between  Cal- 
ifornia and  the  Old  World  are  known,  but  it  has  been  collected  near 
San  Diego  and  there  is  a  probability  that  this  species,  like  Targi- 
onia  hypophylla,  will  be  found  to  extend  into  Mexico.  Targionia 
is  represented  in  Mexico  by  two  forms  which  were  once  considered 
distinct  species  but  which  have  recently  been  reduced  by  Herr 
Stephani*  to  T.  JiypopJiylla.  The  presence  of  this  peculiar  genus 
Targiojiia'm  Mexico,  the  Canary  Islands  and  Africa,  audits  absence, 
so  far  as  is  known,  in  eastern  Asia  suggest  the  possibility  of  some  pre- 
historic land  communication  across  what  is  now  the  Atlantic  Ocean. 
The  possibility  of  spores  or  entire  plants  finding  their  way  across 

*  Bull.  Herb.  Boiss.  6:  764.  1898. 


Introduction 


9 


any  extended  portion  of  the  sea  and  retaining  their  vitaHty  seems 
very  remote. 

A  departure  from  the  customary  classification  is  made  in  the 
treatment  of  the  Anthocerotales.  The  commonly  accepted  arrange- 
ment recognizes  two  classes — the  Hepaticae  and  the  Miisci — in  the 
pJiylinii  or  siibkingdoin  Bryophyta,  though  by  some  writers  the 
SpJiagnaccae  are  taken  out  from  the  Miisci  and  made  a  separate 
class.  Under  the  class  Hepaticae  are  ordinarily  recognized  the 
three  orders  Marchautiales,  Jitngennainiiales,  and  AntJiocerotales. 
The  last-named  group  differs  from  the  Hepaticae  proper  in  very 
important  characters  and  we  think  these  differences  are  best  em- 
phasized by  raising  the  Aiithocerotales  to  the  rank  of  a  class  (under 
the  name  Anthocerotes),  coordinate  with  the  Hepaticae,  restrict- 
ing this  last  term  to  the.  Marchajitiales  and  Jiingernianiiiales.  This 
idea  is  not  wholly  a  new  one.  Students  of  plant  morphology, 
among  whom  Professor  D.  H.  Campbell  may  be  especially 
mentioned,  have  been  approaching  this  view  for  several  years, 
and  this  position  was  definitely  taken  by  Mons.  L.  A.  Gayef^ 
in  1897  as  a  result  of  investigations  on  the  archegonium 
alone,  though  the  class  was  not  named  and  defined  by  him  in  the 
lormal  systematic  way.  The  peculiarities  of  the  Anthocerotes, 
briefly  enumerated,  are:  (i)  A  single  large  chloroplast  to  each 
cell,  instead  of  several  smaller  chloroplasts  as  in  the  assimilative 
tissues  of  the  Hepaticae  proper.  Chlorophyll  bodies  like  these — 
solitary  in  the  cell — are  known  to  occur  elsewhere  only  in  certain 
green  algae  and  in  certain  tissues  of  Selaginella.\  (2)  The  an- 
theridia  arise  within  the  thallus — are  endogenous  in  origin — and 
the  walls  of  the  immersed  archegonia  are  confluent  with  the 
adjacent  tissues.  (3)  The  presence  of  meristematic  tissue  in  the 
capsule  near  its  base,  by  the  activity  of  which  the  capsule  has  a 
long  continued  growth,  ripening  spores  toward  its  apex  while  form- 
ing new  spore-mother-cells  below.  (4)  The  presence,  in  probably 
all  cases,  of  a  columella  around  which  the  archesporium  is  formed. 
(5)  The  presence,  in  many  cases,  of  stomata  on  the  capsule,  with 
accompanying  assimilative  tissue.  These  five  characters  are  pecu- 
liar to  the  Anthocerotes  as  distinguished  from  the  Hepaticae  proper 

^  Ann.  Sci.  Nat.  Bot.  VIII.  3  :  243,  244.  1897. 

I  Campbell,  D.  H.  The  Structure  and  Development  of  Mosses  and  Ferns,  115.  1895. 


10         Hepaticae  and  Anthocerotes  of  California 

and  they  are  characters  of  much  significance.  For  analogies  we 
must  look  chiefly  in  two  directions — toward  the  Pteridophyta  and 
toward  the  Musci. 

The  principal  list  of  Californian  Hepaticae  and  Anthocerotes 
hitherto  published  is  that  of  Dr.  Bolander,  who  included  thirty 
species  belonging  to  these  two  groups  in  his  Catalogue  of  the 
Plants  Growing  in  the  Vicinity  of  San  Francisco."*  Dr.  Sereno 
Watson  and  Mr.  Coe  F.  Austin  prepared  a  manuscript  descriptive 
of  the  Hepaticae  and  Anthocerotes  of  California  for  incorporation 
in  the  volumes  devoted  to  botany  in  the  Report  of  the  Geological 
Survey  of  California,  but  for  some  reason  it  never  saw  the  light. 
This  manuscript,  in  which  forty  species  are  diagnosed,  is  now  owned 
by  Professor  Underwood  and  has  been  accessible.  At  the  end  of 
the  present  paper  will  be  found  a  list  of  the  existing  literature  bear- 
ing directly  upon  the  hepaticology  of  California  and  also  a  list  of 
other  hepaticological  works  which  have  been  of  the  most  service 
in  the  preparation  of  this  paper. 

With  the  hope  of  making  the  paper  more  useful  to  beginners  in 
the  study  of  the  Pacific  Coast  Hepaticae,  keys  to  the  genera  and 
species  have  been  supplied.  In  the  matter  of  synonymy  and 
bibliography,  no  attempt  is  made  to  give  anything  beyond  the  most 
important  references.  Exsiccatae  are  cited  only  when  Californian 
specimens  are  involved.  In  regard  to  nomenclature,  the  Rochester 
Code  is  applied  in  general,  though  perhaps  not  always  rigidly  and 
consistently ;  apparent  exceptions  are  discussed  under  Grivialdiay 
Lepidozia^  and  Asterella  Calif ornica. 

It  has  been  our  aim  to  distinguish  as  clearly  as  possible 
between  statements  based  upon  our  own  observations  and  those 
based  upon  the  recorded  observations  of  others,  yet  borrowings 
doubtless  exist  on  the  following  pages,  particularly  in  the  diagnoses 
of  genera,  which  are  not  expressly  acknowledged  to  be  such  at 
the  place  of  their  occurrence.  In  framing  definitions  of  genera,  we 
have  been  assisted  especially  by  the  writings  of  Spruce,  Schiffner, 
Lindberg,  and  Limpricht. 

Whatever  of  value  the  present  work  may  possess  is  largely  due 
to  the  encouragement  received  from  Professor  L.  M.  Underwood, 
whose  library,  herbarium,  and  mature  experience  have  been  re- 

*  California  Medical  Gazette,  May,  1870. 


Introduction 


11 


sources  ever  most  generously  placed  at  our  command.  In  addi- 
tion to  the  friends  already  mentioned,  who  have  communicated 
Californian  specimens,  our  thanks  are  due  to  numerous  others  who 
have  kindly  loaned  types  or  sent  specimens  for  comparison.  We 
are  especially  indebted  in  this  way  to  Mr.  W.  H.  Pearson,  Professor 
F.  E.  Weiss,  Herr  F.  Stephani,  Dr.  B.  L.  Robinson,  Dr.  E.  Levier, 
Dr.  H.  W.  Arnell,  Sir  W.  T.  Thiselton-Dyer,  Professor  C.  G.  Limp- 
richt.  Dr.  Harald  Lindberg,  Herr  M.  Heeg,  Mr.  Matthew  B.  Slater, 
Dr.  Alexander  W.  Evans,  Professor  Romualdo  Pirotta,  Professor 
Gunther  Ritter  von  Beck,  Mr.  O.  F.  Cook  and  Mons.  P.  Hariot. 


12 


Hepatic  AE 


Class  HEPATICAE. 

Protonema  very  small,  often  rudimentary.  Gametophore  thal- 
lose  or  foliose  or,  in  a  few  genera,  of  a  character  intermediate 
between  these  two  types,  mostly  dorsi ventral.  Leaves,  when 
present,  composed  of  a  single  layer  of  cells,  never  with  a  true 
midvein,  a.rranged  upon  the  stem  in  two  lateral  ranks,*  with  some- 
times" a  more  or  less  well-developed  ventral  (third)  rank.  Root- 
hairs  unicellular.  Chloroplasts  small,  several  or  many  in  a  cell 
in  the  assimilative  parts.  Antheridia  and  archegonia  arising  each 
from  the  subdivisions  of  a  surface  cell  ;  archegonium  individual- 
ized, its  wall  (except,  rarely,  the  basal  portion  of  the  venter)  al- 
ways free  from  the  adjacent  tissues. 

Sporogonium  permanently  enclosed  within  the  modified  arche- 
gonium wall  (calyptra)  or  more  frequently  included  until  the 
capsule  is  nearly  mature,  when  the  calyptra  is  ruptured  near  the 
top  by  the  growth  of  the  capsule  or  by  the  elongation  of  the  seta. 
Capsule  without  columella  and  without  stomata,  irregularly  dehis- 
cent, operculate,  or  more  often  opening  by  (usually)  four  valves. 
Spores  ripening  simultaneously,  accompanied  (except  m  Ricciaceac) 
by  elongated,  thin-walled  sterile  cells  which  (with  rare  exceptions) 
are  provided  with-  spiral  thickenings  (when  each  sterile  cell  is 
known  as  an  elater). 

Order  I.    MARCHANTIALES.  f 

Gametophore  a  thallus,  consisting  of  several  distinct  layers  of 
tissue,  the  usually  clearly  defined  dorsal  epidermis  %  containing 
pores  ("stomata"  §)  which  communicate  with  narrow  vertical 

*  Except  in  a  very  few  genera,  none  of  which  are  known  to  occur  in  California, 
■f  For  diagnosis  of  Order  II,  JungermannialeSy  see  page  62. 

\  Dumorliera,  not  yet  found  in  California,  has  a  very  exceptional  thallus,  being 
without  perceptibly  differentiated  epidermis,  pores,  or  air-chambers. 

^  The  so-called  "  stomata  "  of  the  gametophore  of  the  ISIarchantiales  are  quite  dif- 
ferent in  structure  from  the  stomata  of  the  sporophyte  in  the  Anthoccrotes,  Musn\ 
Pteridophyia,  and  Sperniatophyta^  and  are,  perhaps,  only  in  part  analogous  in  function. 
But  '*  stoma  "  as  applied  to  the  "  breathing  pore,"  together  with  its  more  or  less  modi- 
fied epidermal  surroundings,  is  a  convenient  term  in  the  description  of  the  Marchan- 
tiales  and  the  use  of  the  word  in  this  extended  sense  is  sanctioned  by  Lindberg,  Schiff- 
ner,  Stephani,  and  others. 


Marchantiales 


13 


canals  or  enlarged  air-chambers  in  the  subjacent  chlorophyll-bear- 
ing layer  and  are  often  bounded  by  specialized  epidermal  cells. 
Ventral  surface  mostly  provided  with  scales  arranged  in  longitud- 
inal rows.  Root-hairs  of  two  forms  upon  the  same  plant,  the  one 
with  smooth  walls,  the  other  with  interior  peg-like  papillae.  Sex- 
ual organs  aggregated  in  specialized  parts  of  the  thallus  or  on 
modified  branches,  or  in  the  lower  forms  irregularly  scattered. 

The  first  four  segments  of  the  embryo  sporogonium  arranged 
like  the  quadrants  of  a  sphere.  Matured  sporogonium  consisting 
of  a  capsule  without  stalk  or  foot,  all  of  the  interior  cells  forming 
spores,  or  of  capsule,  foot,  and  short  seta  (rarely  attaining  length 
of  1-2  mm.),  when  some  of  . the  interior  cells  produce  elaters. 

The  order  Marchantiales  comprises  the  two  families  Ricciaccae 
(see  below)  and  MarcJiantiaceae  (see  page  34.) 

Family  I.  RICCIACEAE. 

Gametophyte  a  carnose,  dichotomously  branched  or. rarely  sub- 
simple  thallus  ;  chlorophyll -bearing  tissues  of  suberect  lamellae 
enclosing  very  narrow  and  inconspicuous  subvertical  air-canals  or 
more  ample  chambers,  these  unoccupied  by  a  specialized  assimi- 
lative tissue.  Stomata  rudimentary,  rarely  well-developed.  Ven- 
tral scales  mostly  present  but  sometimes  obscure.  Sexual  organs 
arising  singly  from  the  dorsal  surface  just  back  of  the  growing 
apex,  soon  becoming  deeply  immersed  in  the  thallus,  the  elongated 
archegonium-neck  commonly  exserted,  the  walls  of  the  efferent 
canal  of  the  antheridial  cavity  often  produced  into  a  conico-cylin- 
drical  elevation. 

Sporophyte  a  capsule,  without  foot  or  seta,  always  enclosed  by 
the  calyptra,  in  which  the  spores  come  to  lie  at  maturity  through 
the  disappearance  of  the  delicate  capsule-wall.  Inner  cells  all 
producing  spores. 

I.  RICCIA  L.  Sp.  PI.  1 1 38.     1753.    Ex  Mich.  Nov.  PI.  Gen. 

106.  1729. 

Plants  terrestrial,  or  rarely  aquatic,  often  forming  rosettes  or 
half-rosettes,  with  linear,  ovate,  or  cordate  lobes,  usually  closely 
attached  to  the  soil  by  root-hairs,  the  latter  either  smooth-walled 
or  furnished  with  peg-like  protuberances  from  the  wall  into  the 


14  RiCCIACEAE 

lumen,  thallus-segments  commonly  with  a  distinct  median  sulcus  on 
dorsal  surface  ;  lamellae  of  the  chlorophyll-bearing  layer  enclos- 
ing very  narrow  subvertical  air-canals  or  less  commonly  larger 
chambers,  the  latter  when  present  often  breaking  through  to  the 
surface,  giving  the  thallus  a  spongy  or  lacunose  appearance  ;  ventral 
portions  (costa)  of  the  thallus  composed  of  parenchymatous  cells 
nearly  or  wholly  destitute  of  chlorophyll ;  a  rather  poorly  defined 
layer  of  somewhat  prosenchymatous  starch-bearing  cells  sometimes 
distinguishable  between  the  chlorophyll-bearing  and  ventral  strata  ; 
cells  of  the  primarily  superficial  layer  of  the  dorsal  epidermis  often 
soon  collapsed  and  more  or  less  disintegrated.  Stomata  repre- 
sented by  small  pores  surrounded  by  slightly  or  not  at  all  modified 
epidermal  cells.  Scales  hyaline-albescent,  brownish-violet,  or  dark- 
purple,  often  fugacious,*  rarely  surpassing  the  margins  of  the 
thallus,  originating  in  a  single  f  median  series,  soon  ruptured  along 
the  median  line  and  becoming  apparently  two-ranked.  Monoicous 
or  dioicous.  Sexual  organs  scattered,  the  antheridia  irregularly 
intermingled  with  the  archegonia  in  the  monoicous  species.  J  In- 
volucre none.  Capsules  immersed.  Spores  large,  for  the  most 
part  distinctly  tetrahedral,  the  surface  marked  with  free  or  most 
frequently  mesh-forming  ridges,  the  angles  of  the  areolae  often  with 
papilliform  elevations,  or  the  inner  (plane)  spore-faces  sometimes 
simply  punctate. 

Key  to  tlie  Species. 

Thallus    solid,"  i.  e.,  with  very  narrow  vertical  or  subvertical  air-canals  which  com- 
municate only  inconspicuously  with  the  dorsal  surface. 
Thallus  furnished  with  scales  underneath,  not  ciliate-margined. 

Scales  extending  considerably  beyond  the  margin,  whitish-hyaline ;  spores 
immarginate,  obscurely  angular,  90-126 in  maximum  diameter. 

6.  R.  Americana. 

*  R.  crystallina  L.  is  said  to  have  no  traces  of  scales  even  as  rudiments.  They 
are  scarcely  distinguishable  in  some  of  the  other  Californian  species. 

f  Herr  Stephani,  contrary  to  the  usually  accepted  view,  states  (Bull.  Herb.  Boiss. 
6  :  311.  1898)  that  doubtless  most  Ricciae  have  an  originally  double  series  of  scales. 
In  all  the  Californian  species  which  we  have  been  able  to  subject  to  microtome  section- 
ing, viz.,  RR.  Americana^  nigrella,  minima ^  trichocarpa,  Calif ornica,  and  Camp- 
belliana,  the  younger  scales  are  clearly  seen  in  a  transverse  section  to  extend  uninter- 
ruptedly across  the  median  line  and  to  be  attached  to  the  body  of  the  thallus  on  either 
side  ;  later  they  become  biseriate  by  a  median  longitudinal  rupture. 

%  Professor  Campbell  has  found  in  his  researches  on  the  development  of  the  sexual 
organs  in  the  Californian  Ricciae  that  **  several  of  one  sort  or  the  other  would  be  formed 
in  succession."  We  have  sometimes  had  difficulty  in  finding  anything  but  archegonia 
in  species  that  could  at  other  times  be  clearly  shown  to  be  monoicous. 


RiCCIA 


15 


Scales  not  reaching  margin  or  rarely  slightly  exceeding  it  at  thallus-apex. 

Scales  blackish-purple;  thallus-segments  l-I-S  mm.  wide;  antheridial 
ostioles  scarcely  or  not  at  all  elevated.  8.  li.  nigrella. 

Scales  brown  or  nearly  colorless,  usually  inconspicuous  ;  thallus-segments 
1-3  mm.  (mostly  1.5-2  mm.)  wide;  antheridial  ostioles  prominent, 
conico-cylindrical.  7.  R.  Campbelliana. 

Scales  hardly  distinguishable  unless  at  thallus-apex,  whitish  or  hyaline  ; 
thallus-segments  mostly  1-2  mm.  wide  ;  ostioles  elevated  about  .08 
mm.;  superficial  epidermal  cells  soon  becoming  cup-like  by  disappear- 
ance of  the  upper  portion  of  their  wall ;  plane  faces  of  the  spores 
merely  punctate  or  marked  with  numerous  low  mostly  free  ridges. 

5.  R.  minima. 

Thallus  ciliate  on  margins  or  at  apex. 

Thallus-segments  .75-1.5  mm.  (mostly  I  mm.)  wide,  the  margins  densely 
ciliate  ;  I-I2  cilia,  with  rare  exceptions,  springing  from  the  dorsal  epidermis 
above  each  capsule  ;  spores  soon  black  and  very  opaque,  90-120^  in  maxi- 
mum diameter.  2.  R.  trichocarpa. 
Thallus-segments  1-2  mm.  wide,  bearing  few  or  numerous  short  cilia  on  mar- 
gins toward  apex  but  none  on  dorsal  surface  ;  antheridial  ostioles  not  ele- 
vated ;    spores  brown,  70-90  a  in  maximum  diameter,  meshes  of  convex 
face  6-10  //  in  width.  3.  R.  Californica. 
Thallus-segments  2-2.5  i^^ni.  wide,  the  margins  bearing  a  few  short  stout 
cilia  ;  ostioles  conico-cylindrical,  .  I-.2  mm.  high  ;  spores  ^  90-1 18  //  in  maxi- 
mum diameter.  I.  R.  Lescuriana. 
Thallus  naked  on  margins  and  underneath  or  with  very  rudimentary  scales,  very 
rarely  with  rudimentary  cilia  toward  apex  ;  antheridial  ostioles  conico-cylindrical ; 
plane-faces  of  the  spores  reticulate-areolate,  meshes  of  the  convex  face  10-15 /z 
in  width.  4.  R.  glauca. 
Thallus  with  large  air-chambers  which  finally  communicate  conspicuously  with  the  dor- 
sal surface,  giving  thallus  a  spongiose  or  vesiculose-alveolate  appearance. 
Thallus  light-green,  rather  loosely  attached  to  the  soil,  very  thin,  the  transverse 
sections  at  least  six  times  broader  than  high  ;  spores  90-108  //  in  maximum 
diameter,  with  a  margin  4-10  /z  wide.  9.  R.  Catalinae. 
Thallus  often  yellowish-green,  very  closely  adherent  to  the  soil,  transverse  sections 
of  the  segments  2.5-4  times  broader  than  high;  spores  60-90^  in  maximum 
diameter,  with  a  narrow  (sometimes  almost  deficient)  margin  3-5  //  in  width. 

10.  R.  crystallina. 

I.  RicciA  Lescuriana  Aust.  Proc.  Acad.  Nat.  Sci.  Philad.  1869: 

232.  1869. 

Thallus  light  green  and  reticulate  above,  concolorous  below, 
or  tinged  with  purple  at  margins,  orbicular  or  semiorbicular  in 
general  outline ;  the  principal  divisions  few,  with  a  wide-angled 
(30°  90°)  dichotomy  1-3  times  repeated  or  subcruciately  lobed, 
6-10  mm.  X  2-2.5  mm.,  mostly  linear-obcuneate,  the  shorter 
sometimes  nearly  obcordate  ;  terminal  segments  ovate-elliptical  to 


*  See  remarks  on  the  size  of  the  spores  of  the  Californian  R.  Lesair. 


16 


RiCCIACEAE. 


oblong,  subacute,  with  a  few  usually  quite  inconspicuous  hyaline 
or  whitish  ventral  scales  at  the  apex,  the  flat-bottomed  median 
sulcus  occupying  about  one  third  their  width  but  narrowed  and 
apparently  closed  in  front  by  the  convergence  of  the  abruptly  as- 
cending margins,  these  often  incurv^ed  o'ri  drying,  somewhat  incras- 
sate,  though  commonly  subacute  in  cross-section,  bearing  in  a 
single  or  double  series  a  few  stout'  obtuse  or  sharp-pointed  cilia 
.1-3  mm.  long;  transverse  sections  rounded  ventrally,  their 
wddth  2-3  times  their  height,  about  20  cells  thick  in  median  parts, 
the  air-chambers  narrow  ;  cells  of  the  superficial  layer  of  the  epi- 
dermis soon  collapsed  and  disintegrated  or  irregularly  persisting 
as  cups  attached  to  the  cells  below :  monoicous :  antheridia 
abundant,  their  ostioles  prominent,  conic-cylindrical,  .1— .2  mm. 
high  :  capsules  with  a  naked,  sometimes  purple  thallus-covering ; 
spores  brown,  75-118  ^«  in  maximum  diameter,  angular,  with  a 
slightly  granulate,  more  or  less  interrupted  margin  3—12  ti  in 
width,  the  outer  face  lightly  papillate  or  nearly  smooth  in  profile, 
8  or  9  strongly  defined  areolae  measuring  its  diameter,  the  inner 
faces  a  little  less  distinctly  areolate  or  marked  with  ridges  which 
scarcely  anastomose. 

On  the  bank  of  a  rivulet,  Fort  Ross,  Sonoma  County,  March 
15,  1896. 

The  Californian  plant  differs  from  the  typical  R.  Lescurimia 
chiefly  in  the  larger  spores  (90-1 18  n  in  the  former  ;  75-100  n  in 
the  latter)  which  are  also  more  distinctly  areolate  on  the  inner 
faces.  The  thallus-margins  and  the  thallus-covering  of  the  cap- 
sules, too,  show  less  purple  than  Mr,  Austin's  original  plants  from 
New  Jersey,  but  these  differences,  we  think,  are  not  sufficiently 
important  to  be  considered  specific.  Specimens  collected  at  Jack- 
sonville, Florida,  by  J.  Donnell  Smith  in  1877,  agree  essentially 
with  the  Fort  Ross  plant.  Riccia  Lesciiriana  is  perhaps  more  likely 
to  be  confused  with  R.  Californica  than  with  any  of  the  other 
Californian  Ricciac,  {xom  which,  however,  it  may  be  easily  distin- 
guished by  the  larger  spores,  the  prominent  antheridial  ostioles, 
the  shorter,  stouter  cilia,  etc. 

We  would  place,  for  the  present,  with  the  above  species  a 
somewhat  anomalous  Riccia  collected  in  California  •  by  Bolander 
(in  herb.  Underwood,  from  U.  S.  National  Museum).  The  thallus 
is  but  once  forked  and  unusually  thin,  the  transverse  sections  be- 
ing 4-6  times  as  wide  as  high  ;  the  cilia  are  rudimentar}'  or  want- 
ing ;  the  spores  have  more  numerous  and  smaller  areolae,  10-12 


RiCCIA 


IT 


measuring  the  diameter  of  the  convex  face.  Its  large  spores  (90— 
105//),  the  broad  divergent  thallus-lobes  and  other  minor  charac- 
ters stand  in  the  way  of  its  reference  to  R.  glauca. 

2.  RicciA  TRiCHOCARPA  M.  A.  Howe,  Bull.  Torr.  Bot.  Club,  25  : 
184. //.jj 7.  1898. 

Thallus  in  rosettes  about  2  cm.  in  diameter  or  forming  some- 
what irregularly  radiating  masses  ;  the  principal  divisions  linear, 
2-6  times  dichotomous,  .75-1.5  (mostly  1)  mm.  in  width,  often 
black  below  and  at  margins,  densely  clothed  at  the  sides  with  white 
or  tawny  setae  .3-. 6 5  mm.  long,  those  toward  the  apices  often  in 
as  many  as  8—12  irregular  series,  mostly  distinct  at  insertion,  with 
sharp,  rigid,  rarely  slightly  uncinate  points,  the  terminal  thallus- 
lobes  obcuneate  or  oblong-elliptical,  obtuse  or  subacute,  nar- 
rowly and  rather  deeply  unicanaliculate  toward  the  apices  or  sub- 
bicanaliculate,  the  furrow  at  the  extremities  commonly  concealed 
by  the  trichomes,  the  median  sulcus  sometimes  nearly  vanishing 
toward  the  base,  margins  obtusely  rounded,  tumid,  often  connivent 
on  drying,  the  furrow  then  thatched  by  the  somewhat  forwardly 
directed  setae  ;  dorsal  surface  light  green,  minutely  and  regularly 
reticulate  ;  ventral  surface  nearly  plane  and  flat,  with  a  few  very 
inconspicuous  scales  on  either  side  of  the  median  line  at  the  apices, 
falsely  squamose  in  the  basal  parts  through  delamination  due  to 
marcescence  ;  width  of  transverse  sections  1.6-3  times  their  height, 
the  posterior  subquadrangular  with  a  light  median  sulcus,  the 
margins  becoming  more  tumid  and  rounded  in  proceeding  toward 
the  apex  and  the  sulcus  now  and  then  double,  the  sections  convex 
ventrally  only  in  the  extreme  apical  region  ;  20-28  cells  in  thick- 
ness in  median  parts,  texture  nearly  solid,  the  air-chambers  narrow 
and  vertical  ;  epidermis  primarily  bistratose,  the  superficial  layer  at 
first  papillate,  afterwards  collapsed,  and  in  the  older  parts  reduced 
to  a  flattened  cellulose  membrane  incumbent  on  the  lower  layer  : 
monoicous  :  antheridia  scattered,  the  ostioles  elevated  about  .  I  mm.: 
sporogonia  numerous,  in  a  single  or  double  series,  immersed, 
finally — especially  when  dry — hemispherical-protuberant  above, 
together  with  the  long-unruptured  cov^ering,  or  subconical,  the  cov- 
ering marked  with  a  dark-purple  spot  and  bearing  with  rare  excep- 
tions I— 12  setae;  spores  soon  black  and  very  opaque,  narrowly 
or  not  at  all  margined,  90-120//  in  maximum  diameter,  minutely 
granulose-papillate,  with  9-12  areolae  (visible  only  in  the  younger 
spores)  across  the  convex  face,  the  walls  of  these  with  irregularly 
thickened  and  salient  angles,  thus  often  giving  this  face  in  profile 
the  appearance  of  bearing  columnar  or  wart-like  elevations,  areolae 
across  one  of  the  plane  faces  5-7  in  number,  scarcely  elevated  at 
the  angles. 


18 


RiCCIACEAE 


Exsicc.     Hep.  Bor.-Am.  143''  (as  R.  Uiin'da  Lindenb.). 

Hep.  Am.   138*  (as  R.  arvcnsis  Aust.,  var.  hlrta 
Aust.). 

On  rather  dry  and  rocky  soil.  About  San  Francisco,  "  Mis- 
sion Dolores,"  "Turk  St.,"  "near  Ocean  House"  (Dr.  Bolander), 
Mission  Dolores  (Howe);  Berkeley  (Howe);  Ukiah  (Howe);  near 
Stanford  Universityf  (Prof.  D.  H.  Campbell,  Palo  Alto  (S.  E. 
Brown);  Santa  Ana  Hills  (Parish);  Pasadena  (McClatchie);  San 
Diego  (Mrs.  Katharine  Brandegee). 

The  above  is  the  Californian  Riccia  that  has  been  referred 
variously  by  American  authors  to  Riccia  ciliata  Hoffm.,  R.  tiniiida 
Lindenb.,  R.  inttnncsccns  Aust.  MS.,  Underw.  (R.  ciliata  Hoffm., var. 
iniumescens  Bisch.)  and  more  recently  to  Riccia  Jiiria  Aust.  (/\ 
arvensis  Aust.,  var.  Idrta  Aust.).  But  Riccia  arvcnsis  Jiirta^  known 
only  from  Mr.  Austin's  specimens  from  the  neighborhood  of 
Closter,  New  Jersey,  bears  comparatively  few,  short,  usually  blunt- 
pointed  and  incurved  papilla-like  cilia  .1— .3  mm.  long  (not  setac^, 
the  thallus  is  strongly  incrassate-carinate  ventrally,  the  margins 
are  acute,  the  epidermis  is  very  thick,  of  2  or  3  layers  of  de- 
colorate  or  purple-tinged  cells,  the  somewhat  smaller,  more  dis- 
tinctly margined  spores  are  fuscous-brown  at  maturity  and  always 
show  their  areolae  quite  clearly  in  glycerine  instead  of  being 
densely  black  and  opaque  as  in  R.  tricJiocarpa  under  the  same 
treatment,  the  plants  are  mainly  smaller  and  less  frequently  dichot- 
omous,  the  thallus-segments  are  usually  comparatively  broader, 
reddish-purple  below  and  more  evidently  squamigerous. 

A  much  nearer  ally  of  Riccia  iricliocarpa  is  doubtless  R.  ciliata 
Hoffm.  of  Europe.  P'rom  the  typical  form  of  this,  however,  the 
Californian  plant  is  clearly  distinct  in  the  shorter  and  much  more 
abundant  setae  (these  often  reach  i  mm.  in  length  in  R.  ciliata),  in 
the  presence  of  trichomes  over  the  sporogonia,  in  the  rather  larger 
size  of  the  thallus,  more  acute-angled  dichotomy,  and  in  the  com- 
monly black  margins  and  sides.     From  R.  ciliata  lutimicscciis 

*See  footnote,  Bull.  Torr.  Bot.  Club,  25  :  186.  1898. 

t  Professor  Campbell's  specimens,  which  we  took  as  the  tj^pe  of  this  species,  were 
distributed  in  the  Ilep.  Am.,  by  error,  as  having  been  collected  in  San  Mateo  Co., 
and  this  locality  was  copied  in  connection  with  our  original  description.  We  are  in- 
t"  rmed,  however,  by  Prof.  Campbell,  that  the  specimens  came  from  Santa  Clara  Co., 
near  the  San  Mateo  Co.  line. 


RiCCIA 


19 


Bisch.  known  to  us  only  from  the  excellent  figures  and  descrip- 
tion of  its  author  and  from  a  specimen  communicated  by  Herr  M. 
Heeg,  our  plant  differs  in  the  longer,  narrower  segments,  in  the 
shorter  and  even  more  crowded  lateral  setae,  those  toward  the 
apex  often  in  as  many  as  8-12  irregular  series,  and  in  the  normal 
presence  of  1-12  long  trichomes  above  each  sporogonium,  while 
in  R.  ciliata  intwnescens,  the  ''cilia"  appear  to  be  of  somewhat 
rare  occurrence  on  the  dorsal  surface.  In  R.  ciliata,  the  tissues 
covering  the  mature  sporogonium  soon  become  thin,  scarious,  and 
shining,  and  finally  break  away  in  fragments,  exposing  the  capsule 
and  the  spores,  and  Bischoff  *  remarks  of  the  capsules  of  the  var. 
intuniescens :  post  matiudtatem  ntpti,  foveolas  sporis  replctas  in 
frondis  pagina  siipcriore  relinqiientes''  ;  while  in  R.  tricliocarpa  the 
covering  of  the  capsules  remains  very  long  intact — indeed,  in  only 
one  case  out  of  several  specimens  with  spores  evidently  much  past 
maturity,  have  we  seen  the  contents  of  a  capsule  exposed  by  nat- 
ural agencies. 

Riccia  crinita  Tayl.  from  Swan  River,  Australia  (Drummond, 
no.  42),  the  original  of  which  we  have  seen  through  the  kindness 
of  Dr.  B.  L.  Robinson,  is  close  to  R.  tricJiocarpa  in  character  and 
number  of  setae,  which  also  sometimes  occur  over  the  sporogonia. 
But  R.  crinita  is  a  smaller  plant,  only  2  or  3  times  dichotomous, 
with  shorter,  oblong  rather  than  linear  segments,  the  thallus  is 
comparatively  much  thinner,  the  vertical  sections  of  its  segments 
being  3-7  times  as  wide  as  high,  the  margins  are  acute  and  com- 
monly incurv^ed  ;  the  spores  (possibly  not  arrived  at  full  depth  of 
color)  are  light-brown,  in  maximum  diameter,  with  11-14 

smaller  areolae  across  the  convex  face,  this  scarcely  papillate  in 
profile,  the  plane  faces  similarly  areolate,  the  mesh-forming  ridges 
throughout  and  the  narrow  margin  nearly  smooth. 

Riccia  Michelii  Raddi,  var.  ciliaris  Levier  {=R.  tiunida  Lindenb. 
and  R.  palmata  Lindenb.  fide  Levier)  differs  so  widely  from  our 
species  that  a  detailed  comparison  is  unnecessary. 

R.  tricliocarpa  may  be  found  with  archegonia  and  antheridia  in 
January  and  early  February,  ripening  its  capsules  in  April  and 
May.  Like  all  the  Californian  Ricciae  it  is  practically  invisible 
during  the  summer  months. 


Acta  Acad.  Caes.  Leop. -Carol.  Nat.  Cur.  17  :  1063.  1835. 


20 


RiCCIACEAE 


Plate  88.    Riccia  trichocarpa. 
I  and  2.  Portions  of  the  plant,  natural  size. 

3.  Terminal  segments  of  a  young  thallus,  dorsal  view,  X  5- 

4.  End  of  thallus  lobe  from  fig.  i,  with  ostioles  and  a  sporogonium,  X 

5.  Lateral  view  toward  apex  of  a  thallus- segment,  showing  the  numerous  tri- 
chomes,  X  20. 

6.  Marginal  trichomes,  X  53- 

7.  Portions  of  thallus  exhibiting  the  trichome-bearing  elevations  above  the  cap- 
sules, X  20. 

8-13.  Outlines  of  transverse  sections  about  lo  //  in  thickness  at  selected  points 
from  near  the  apex  to  the  old  and  shriveled  base,  X  23.  The  trichomes  do  not  appear 
so  abundant  as  might  be  expected  from  the  material  used  for  microtome  sectioning, 
which  may  be  explained  from  the  thinness  of  the  sections  and  by  the  fact  that  the  de- 
tached sections  and  fragments  of  the  trichomes  have  not  always  adhered  to  the  glass 
slides.    Fig.  8  shows  small  median  scales. 

14.  Epidermis  and  subjacent  cells  from  a  younger  portion  of  the  thallus,  X  225. 
The  superficial  layer  is  here  represented  as  composed  of  collapsed  cells.  In  the  ex- 
treme apical  region  only  they  are  papilliform. 

15.  Epidermis  in  older  parts  of  the  thallus,  X  225.  The  collapse  and  disintegra- 
tion of  the  cells  of  the  outer  layer  is  so  complete  that  only  a  structureless  cellulose  film 
remains. 

16.  Cross- section  of  thallus  showing  the  cell  structure  and  a  young  sporogonium^ 
X  53- 

17.  Spores,  X  305- 

Figs.  I,  2,  4-7  and  17  from  Hep.  Am.  no.  138  collected  near  Stanford  University, 
April,  1892,  by  Prof.  D.  H.  Campbell  ;  figs.  3  and  8-16  from  material  fixed  with  I  per- 
cent, chromic  acid  and  preserved  in  alcohol,  collected  by  the  author  near  Berkeley^ 
California,  February  8,  1896. 

3.  Riccia  Californica  Aust.  Bull.  Torn  Bot.  Club,  6  :  46.  1875. 

Thallus  forming  light  green,  somewhat  glaucescent  rosettes 
8—18  mm.  in  diameter,  concolorous  beneath,  reticulate  above,  often 
minutely  puberulent  or  scurfy  when  moistened  ;  the  principal  di- 
visions 1-3  times  dichotomous  ;  terminal  segments  short-oblong 
or  obovate,  1—2  mm.  wide,  bearing  toward  the  subtruncate  or  re- 
tuse  apex  few  or  numerous  whitish  or  subfuscous,  mostly  slender 
and  sharp-pointed  cilia  .16-.4  mm.  in  length,  the  commonly  ob- 
tuse margins  elevated  and  somewhat  tumid  when  young,  the 
median  sulcus  broad  and  obtuse  except  at  the  extreme  apex,  van- 
ishing backward,  the  segments  more  or  less  concave  dorsally  on 
drying,  rarely  nearly  plane,  scales  very  rudimentary  (none  in  the 
ordinary  sense);  width  of  transverse  sections  2-5  times  their  height, 
thickness  of  thallus  15—25  cells  in  median  parts,  the  upper  layers 
of  cells  chlorophyllose,  the  lower  often  nearly  or  quite  destitute  of 
contents,  air-chambers  narrow  and  vertical  ;  cells  of  the  superficial 
layer  of  the  epidermis  hemispherical-convex  or  papilliform  near  the 
growing  apex  of  the  thallus,  soon  collapsed  or  disintegrated,  per- 


RiCCIA 


21 


sisting  only  as  imperfect  and  irregular  cups  or  more  often  as  mere 
wall-fragments  adhering  to  the  hyaline  and  nearly  empty  cells  of 
the  primitively  lower  layer  :  monoicous  :  antheridia  infrequent,  the 
ostioles  not  elevated  above  the  surface  of  the  thallus  :  archegonium- 
neck  exserted  and  often  conspicuous  even  at  maturity  of  the  spor- 
ogonium  :  capsules  deeply  immersed,  with  a  naked  long  unruptured 
covering,  in  rare  cases  exploded-protuberant  with  age  ;  spores 
brown,  finally  darkening  but  always  more  or  less  translucent,  70— 
go  H  in  maximum  diameter,  distinctly  angular,  with  a  smooth  yel- 
low or  brownish  irregularly  crenate  margin  3-12/7.  in  width,  the 
diameter  of  the  outer  face  measured  by  10—13  areolae,  these  mostly 
6-10  a  in  width,  the  papilliform  elevations  at  the  angles  of  the 
meshes  3-4  a  or  less  in  height  when  seen  in  profile,  the  inner  faces 
with  similar  but  less  papilliform-angled  areolae. 

On  lightly  shaded  banks.  California  "  (Dr.  Bolander  ;  the 
original  specimens  in  Herb.  Pearson).  Berkeley  (Howe),  Fruit 
Vale  (Miss  Edith  S.  Byxbee) ;  Clarendon  Heights,  San  Francisco 
(530*),  and  near  the  Mission  Dolores  (512);  Ukiah,  Mendocino  Co. 
(783) ;  Silver  Lake,  i\lpine  Co.  (Geo.  Hansen). 

The  cilia  in  this  species  are  occasionally  reduced  and  incon- 
spicuous, but  are  always  present,  we  believe,  even  though  some- 
times only  in  a  rudimentary  form. 

Riccia  siibinerniis  Lindb.  ( 1 88 1),  of  which  we  have  been  allowed 
to  see  authentic  specimens  through  the  kindness  of  Mr.  Harald 
Lindberg  and  specimens  of  which  we  also  owt  to  Herr  M.  Heeg, 
differs  from  R.  Calif  arnica  in  the  usually  much  shorter  and  less 
numerous  cilia,  in  the  elevated  conico-cylindrical  antheridial  ostioles, 
in  the  larger  meshes  of  the  outer  face  of  the  spores,  in  the  less 
regular  areolation  of  the  inner  faces,  etc.  The  areolae  of  the  outer 
face  in  R.  siibinerinis  and  R.  ghntca  are  mostly  io-15/.f  in  width, 
while  in  R.  Calif ornica  they  are  usually  only  wide.  We 

agree  with  Dr.  Levier  and  others  in  thinking  Riccia  siibincrinis 
doubtfully  distinct  from  R.  glauca. 

Pirate  89.    Riccia  Californica. 

1.  The  plant,  natural  size,  from  a  dried  specimen. 

2.  Two  of  the  principal  segments  of  the  thallus,  from  alcoholic  material,  X  5- 

3.  A  portion  of  Austin's  original  plant  (soaked  out),  ex  herb.  Pearson,  X  5- 
4-8.  Outlines  of  transverse  sections  of-a  principal  segment  at  somewhat  regular  in- 
tervals, passing  from  near  apex  toward  the  base,  X  23. 


*  The  numbers  refer  to  specimens  collected  by  the  author. 


22 


RiCCIACEAE 


9.  Transverse  section  near  the  middle  of  a  principal  segment  of  another  plant, 
X23. 

10.  From  same  segment  as  fig.  9,  but  near  the  base,  X  23. 

11.  Epidermal  cells  from  near  the  thallus-apex,  X  225. 

12.  Epidermis  from  an  older  part  of  the  thallus,  showing  remnants  of  the  primitive 
surface-layer,  X  225. 

13.  Marginal  trichomes,  X  53- 

14.  Transverse  section  of  thallus,  X  53- 

15.  Spore,  showing  the  inner  (plane)  faces,  X  3^5- 

16.  Spore,  showing  the  outer  (convex)  face,  X  3^5- 

Figs.  I,  2,  9,  10  and  12  from  no.  530,  Clarendon  Heights,  San  Francisco;  fig. 
3,  from  Mr.  Austin's  original,  ex  herb.  Pearson;  figs,  il,  13,  14-16  from  mateiial 
collected  at  Fruit  Vale,  by  Miss  Edith  S.  Byxbee ;  figs.  2  and  4-14  from  material  fixed 
with  I     chromic  acid  and  preserved  in  alcohol. 

4.  RicciA  GLAUCA  L.  pvo  parte  maxima  Sp.  PL  1 139.  1753. 

Differs  from  R.  Californica  chiefly  in  the  normal  absence  of 
cilia,  the  presence  of  cylindrical  or  conico-cylindrical  antheridial 
ostioles  projecting  about  .  i  mm.  above  the  surface  of  the  thallus, 
and  in  having  the  meshes  of  the  outer  face  of  the  spore  10-15 
width. 

Little  Bear  Valley,  San  Bernardino  Mts.  (S.  B.  Parish,  no. 
1670,  Aug.  1884,  in  herb.  Underwood)  ;  near  Mineral  King, 
Tulare  Co.  (Coville  and  Funston,  Death  Valley  Expedition,  no. 
1509,  Aug.  6,  1 891). 

Mr.  Parish's  plant  differs  from  the  larger  forms  of  the  European 
R.  glauca  in  nothing,  we  think,  but  the  rather  smaller  meshes  of 
the  convex  face  of  the  spore,  which,  in  this  specimen,  are  com- 
monly only  about  10  a  in  width.  The  specimen  from  Tulare  Co., 
while  exhibiting  the  ostioles  of  R,  glauca,  has  now  and  then  rudi- 
mentary cilia  toward  the  thallus-apex  and  would  doubtless  be 
identified  with  the  Riccia  subinermis  Lindb.  of  Europe  by  those 
who  maintain  this  as  a  species. 

It  may  be  remarked  that  according  to  the  drawings  of  Herr 
Waldner"^  one  might  gain  the  impression  that  the  ostioles  of  Riccia 
glauca  are  scarcely  elevated  above  the  general  surface  of  the  thallus, 
but  it  is  probable  that  the  antheridium  in  even  the  most  advanced 
stages  represented  by  Waldner  was  still  far  from  maturity.  The 
Antheridienstifte  "  of  R.  glauca  are  described  by  Limprichtt  as 
"  w^ilzenformig  vortretend  "  and  we  find  them  thus  (.1  mm.  long) 
in  G.  &  R.  Hep.  Eur.  no.  646. 

*  Leitgeb,  Untersuch.  iiber  d.  Lebcrm.  Heft  4.  //.  2.  f.  4-6. 
t  Cohn,  Krypt.-Fl.  v.  Schles.  i  :  349.  1877. 


RiCCIA 


23 


5.    RiCCIA  MINIMA  l^.  p.  p.  Sp.  PI.   I  139.  1753. 

Riccia  sorocarpa  Bisch.  Act.  Acad.  Caes.  Leop. -Carol.  Nat. 
Cur.  17  :  1053.  1835. 

Thallus  at  first  subradiate,  forming  later  irregularly  gregarious 
patches,  of  a  bright,  clear,  subcrystalline  light  green  (not  puberu- 
lent  or  scurfy)  when  living  or  when  soaked  out  with  water,  of  a 
lighter  green  when  dry,  minutely,  regularly,  and  compactly  reticu- 
late above  when  moistened,  sometimes  nearly  smooth  on  drying, 
concolorous  below  ;  the  principal  divisions  4—7  mm.  long,  bifid  or 
once  or  twice  dichotomous  ;  the  terminal  segments  oblong,  .75—2 
mm.  wide,  subacute,  carinate-incrassate,  with  an  acute  median 
furrow  and  naked,  ascending,  for  the  most  part  sharply  acute,  often 
hyaline  and  submembranous  margins,  furnished  beneath  toward  the 
apex  with  small  whitish  or  hyaline  scales,  these  often  reaching  the 
margin  but  not  surpassing  it,  the  margins  commonly  elevated  or 
somewhat  incurved  on  drying  ;  transverse  section  parabolic  or  sub- 
quadrate,  its  width  1-4  times  its  height,  about  25  cells  thick  in 
median  parts,  air-chambers  narrow  ;  epidermis  of  two  or  three  layers 
of  cells,  those  of  the  superficial  .stratum  at  first  papilliform,  the 
lower  portion  of  their  wall  becoming  thickened  and  the  upper  soon 
vanishing,  leaving  behind  persistent  cups  attached  to  the  likewise 
thick-walled  cells  of  the  subjacent  layer  :  monoicous  :  antheridia 
few,  the  ostioles  short-cylindrical,  elevated  about  .08  mm.;  cap- 
sules usually  numerous,  with  age  sometimes  emergent-protuberant 
toward  the  base  of  the  segments  by  rupture  of  the  naked  covering  ; 
spores  dark  brown,  70-90  u.  in  maximum  diameter,  angular,  with 
a  brownish  graaulate-papillate  and  crenulate,  usually  interrupted 
margin  3-6  tj.  broad,  the  outer  face  areolate,  10—12  meshes  measur- 
ing its  diameter,  exhibiting  in  profile  papillae  mostly  3-6  fi  long, 
the  inner  faces  densely  and  rather  minutely  punctate  or  furnished 
with  very  short  and  numerous  low  ridges  which  '^do  not  form 
areolae. 

Exsicc.     Hep.  Am.  i  39  (as  R.  glauca). 

Not  uncommon  in  the  coast  region,  often  associated  with  R. 
Calif 07^/ lie  a. 

Berkeley,  Oakland,  Fruit  Vale  ;  Clarendon  Heights  and  near 
Mission  Dolores,  San  Francisco  ;  Fort  Ross,  Sonoma  Co.;  Ukiah 
(782)  ;  Mitchell  Canon,  Mt.  Diablo;  near  Stanford  University  (D. 
H.  Campbell) ;  Pasadena  (A.  J.  McClatchie). 

The  Californian  plant  is  usually  larger  than  the  European  speci- 
mens that  we  have  seen,  yet  it  very  rarely  attains  the  dimensions 
of  representatives  of  this  species  collected  in  Siberia  by  Dr.  H.  W. 


24 


RiCCIACEAE 


Arnell,  to  which  Herr  M.  Heeg  refers  in  the  Botaniska  Notiser 
for  1898  (p.  21)  and  which  we  have  been  privileged  to  examine 
through  the  courtesy  of  Dr.  Arnell.  The  antheridial  ostioles  in 
the  European  plant  so  far  as  we  have  been  able  to  discover  them 
in  specimens  kindly  communicated  by  Herr  Stephani,  Herr  Heeg, 
and  others,  seem  to  be  very  slightly  or  not  at  all  elevated,  yet  in 
the  Siberian  specimens  to  w^hich  allusion  has  already  been  made 
they  sometimes  attain  the  height  of  .  i  2  mm.  above  the  surrounding 
surface,  which  is  in  excess  of  anything  we  have  observed  in  the 
Californian  specimens. 

In  only  one  case  have  w^e  noticed  the  partial  exsertion  of  the 
sporangium  at  maturity  as  figured  and  emphasized  by  Bischoff  in 
his  original  diagnosis  of  R.  sorocarpa.  In  regard  to  this  character 
Herr  Heeg  remarks  :  die  von  Bischoff  besonders  hervorgehobene 
Berstung  der  Lauboberflache  bei  der  Fruchtreife,  bezeichnet  das 
Endstadium  einer  Entwicklungsperiode  und  wird  nur  selten  gerade 
in  diesem  Zustande  aufgesammelt." 

6.  Riccia  Americana  sp.  nov. 

Riccia  laincllosa  Americana  M.  A.  Howe,  /.  /.  Bull.  Torr. 
Bot.  Club,  25  :  189.  1898. 

Thallus  rather  loosely  subradiate  or  forming  compact  more  or 
less  imbricate  masses;  principal  divisions  5-15  mm.  long,  1-3 
times  dichotomous,  very  rarely  simple,  2. 5—4  mm.  in  greatest 
width,  pale  green  and  regularly  reticulate  above,  concolorous  be- 
low, acutely  and  deeply  canaliculate  toward  the  apex,  25-30  cells 
thick  in  median  parts,  furnished  beneath  with  large  obtuse  trans- 
verse patent  or  somewhat  imbricate  subundulate  hyaline  scales, 
these  extending  considerably  beyond  the  acute  or  thin-membra- 
nous ascending  or,  when  dry,  often  erect-connivent  margins  ;  the 
transverse  sections  1.5-3  times  as  broad  as  high  ;  the  terminal 
segments  emarginate,  oblong  to  obcordate,  mostly  obovate  ; 
epidermis  of  two  layers  of  cells,  the  superficial  at  first  subglobose 
or  oval-papilliform,  later  collapsed  or  disintegrated,  two  or  three 
of  the  subjacent  strata  sometimes  becoming  decolorate  :  monoicous 
(polyoicous  ?)  :  antheridia  infrequent,  their  ostioles  not  at  all  or 
slightly  elevated  :  spores  brown,  90-126  u.  in  maximum  diameter, 
obscurely  angular,  wholly  destitute  of  a  wing-margin,  with  8-12 
clearly  defined  areolae,  each  about  i  5  //  in  width,  across  the  outer 
face,  the  inner  faces  marked  with  much  smaller  and  much  less  dis- 
tinct areolae  or  simply  with  irregular  vermicular  lines. 

*Heeg,  Bot.  Notis.  1898:  20.  1898. 


RiCCIA 


25 


On  exposed  or  lightly  shaded  banks,  often  associated  with  R. 
nigrella,  R.  tricJiocarpa  and  R.  Califoniica.  San  Francisco  (Bo- 
lander),  on  hillsides  between  ^Mission  Dolores  and  Clarendon 
Heights,  San  Francisco  (Howe)  ;  Fruit  Vale  (Miss  Edith  S.  Byx- 
bee.  Howe). 

Riccia  Americana  is  closel}'  related  to  R.  lainellosa  Raddi,  from 
which,  however,  it  differs  very  markedly  in  spore-characters. 
The  European  plant  has  distinctly  wing-margined  and  plainly 
angular  spores  and  the  markings  of  the  inner  and  outer  faces  are 
nearly  uniform  ;  the  ridges  of  the  outer  face  are  more  vermicular 
than  in  R.  Anicricana  and  less  often  form  perfect  areolae,  and 
when  the  areolae  are  well  defined,  they  are  smaller  and  more 
numerous,  14  or  15  measuring  the  diameter  of  the  face.  The 
scales  in  R.  Americana  are  more  prominent  than  in  R.  lamel- 
losa.  For  comparison,  we  have  made  use  of  a  specimen  from 
Florence,  Italy  (the  type  locality),  and  one  from  Sicily,  both 
kindly  communicated  by  Dr.  E.  Levier. 

Riccia  Austini  Steph.  (Bull.  Herb.  Boiss.  6  :  336.  1898)  is  a 
nearer  ally  in  respect  to  spore-characters,  with  the  exception  that 
the  spores  are  smaller  (75-i05/>«  in  maximum  diameter),  but  differs 
in  the  much  thinner  and  mostly  smaller  thallus,this  being  only  14-20 
cells  in  greatest  thickness  and  the  breadth  of  the  transverse  sec- 
tions being  3-4  times  their  height,  and  in  the  less  conspicuous 
scales.  R.  Americajia  seems  to  occupy  a  somewhat  intermediate 
position  between  these  two  species.  In  case  it  should  ever  be 
combined  with  R.  Anstini,  this  latter  name  should  be  employed 
for  the  aggregate,  inasmuch  as  it  was  published,  we  are  informed 
by  Dr.  Eugene  Autran,  editor  of  the  Bulletin  de  L Herbier  Boissier^ 
about  two  weeks  in  advance  of  our  Riccia  lamellosa  Americana. 

We  have  been  able  to  see  only  four  or  five  antheridia  in  this 
species  though  hundreds  of  sections  have  been  made.  In  two  in- 
stances, archegonia  occurred  on  the  plants  bearing  the  antheridia  ; 
and  in  two  other  cases  no  archegonia  were  to  be  discovered,  but 
in  the  latter  cases  we  were  sectioning  somewhat  fragmentary  alco- 
holic material.  The  great  majority  of  the  plants  examined  showed 
archegonia  without  traces  of  antheridia. 


26 


RiCCIACEAE 


Plate  90.    Riccia  Americana. 

1.  Dry  plant,  natural  size. 

2.  Plants  from  material  preserved  in  alcohol,  natural  size. 

3.  Thallus-segment,  alcoholic  material,  X  5- 

4.  Thallus-segment,  from  dry  specimen,  X  lO- 

5.  A  tbick  free-hand  transverse  section  from  near  apex  of  thallus,  showing  por- 
tions of  the  scales,  X  23. 

6.  Outline  of  transverse  section  from  same  segment  as  no.  5  but  farther  back, 
X  12. 

7-9.  Outlines  of  microtome  cross-sections  of  thallus  segment  ;  no.  7,  near  apex, 
no.  8,  well  forward,  no.  9,  toward  base,  X  12. 

10  and  II.    Outlines  of  cross-sections  of  thallus-segment  from  another  plant,  X  12. 

12.  Antheridium,  showing  non-elevated  ostiole,  X  ^2. 

13.  Epidermis  near  thallus-apex,  X  225. 

14  and  15.  Epidermis  from  older  parts,  X  225. 

16.  Ventral  scale,  X  23. 

17.  Portion  of  thin  transverse  microtome  section  of  thallus,  X  53- 

18.  Outer  face  of  spore,  X  3^5 • 

19.  Inner  faces  of  spore,  X  3°5- 

20.  Outline  of  optical  section  of  spore-wall,  showing  its  relative  thickness,  X  305 - 
Figs,  I,  4-6,  and  18-20,  drawn  from  specimen  collected  in  Fruit  Vale,  Alameda 

County,  May  2,  1895  ;  2,  lo,  II,  and  13-16,  San  Francisco,  April,  II,  1896  ;  3,  7-9, 
12,  and  17,  Fruit  Vale,  February  i,  1896. 

7.  Riccia  Campbelliana  sp.  nov. 

Thallus  mostly  once  or  twice,  occasionally  three  times,  dichoto- 
mous,  rarely  subradiate,  reticulate  above,  brown  or  nearly  con- 
colorous  below,  very  rarely  blackening,  4-18  mm.  long,  1-3  mm. 
(mostly  1.5-2  mm.)  wide,  more  or  less  rounded  ventrally,  some- 
times carinate,  furnished  beneath  with  brown  or  nearly  colorless 
scales,  these  usually  inconspicuous,  but  sometimes  at  the  apex 
slightly  exceeding  the  thin,  naked,  membranous,  commonly  yellow- 
ish-brown, now  and  then  rather  abruptly  winged  margins,  median 
sulcus  acute,  often  obscure  toward  basal  parts,  the  margins,  on 
drying  usually  ascending,  erect,  or  inflexed-connivent  ;  terminal 
lobes  oblong  or  obovate,  obtuse,  less  commonly  subacute  ;  trans- 
verse sections,  including  margins,  2-5  (mostly  2^)  times  broader 
than  high,  thickness  of  thallus  25-35  cells  in  median  parts,  air- 
canals  narrow  ;  cells,  often  enlarged,  densely  filled  with  a  finely 
granular,  yellowish-brown  material  or  with  a  homogeneous  yellow 
oily  substance,  sometimes  occurring  abundantly  in  all  parts  of 
the  thallus  ;  epidermis  primarily  unistratose,  the  cells  large,  ob- 
long-elliptical in  vertical  section  or  subquadrate-elliptical,  mostly 
higher  than  broad  (40-75//  x  25-50 /i),  long  intact,  disintegrated 
in  the  older  parts  and  replaced  by  an  irregularly  developed 
subjacent  layer:  spores  finally  yellowish-brown,  75— io8/><  (mostly 


RlCClA 


27 


90-100  fi)  in  maximum  diameter,  distinctly  angular,  with  a  mi- 
nutely granulate  or  nearly  smooth  margin  3-6 /y.  width,  the  outer 
face  lightly  papillate  or  nearly  smooth  in  profile,  marked  with  close 
sinuous  ridges  which  rarely  form  completely  closed  meshes,  the 
imperfect  areolae  mostly  4-7  u  in  width,  inner  faces  similarly  but 
less  strongly  marked. 

Near  Stanford  University  (Professor  D.  H.  Campbell,  Feb.  6 
and  May  i,  1896);  on  hills  above  Mission  Dolores  (534,  April 
II,  1896,  with  R.  nigrclla  and  R.  tricJiocarpd) ;  Fort  Ross,  So- 
noma Co.  (March  15,  1896);  Ukiah  (May  13,  1896).  Collected 
by  Professor  Campbell  also  in  1893  (herb.  Underwood). 

Riccia  Cainpbclliaua  in  its  smaller  forms  simulates  to  a  certain 
extent  R.  nigrella,  with  which  it  is  sometimes  associated,  but  is 
always  readily  distinguished  by  the  wider  and  usually  longer  seg- 
ments, which  are  commonly  brown  underneath,  very  rarely  black- 
ening, by  the  thinner  brown  margins,  by  the  larger  spores  and  the 
elevated  antheridial  ostioles.  The  epidermis,  too,  differs  from  that 
of  R.  nigrel/a,  its  cells  being  larger  and  commonly  having  their 
vertical  axis  longer  than  the  horizontal,  while  those  of  R.  nigj'ella 
are  subquadrate  in  vertical  section  and  are  quite  frequently  broader 
than  high,  measuring  25-40/iin  height  by  25—45/^  in  width. 

The  larger  conditions  of  this  species  are  somewhat  suggestive 
of  Riccia  BiscJwffii  Hiiben.  and  R.  Goiigetiana  Mont.,*  especially 
when,  as  sometimes  happens,  the  thallus  is  rather  abruptly  wing- 
margined,  but  the  Californian  plant  differs  clearly  from  both  these 
in  being  monoicous,  in  the  usually  smaller  size,  in  the  entire  ab- 
sence of  cilia,  in  the  smaller,  lighter  colored,  much  more  translu- 
cent spores,  etc. 

Riccia  lauicllosa  Raddi  is  easily  distinguished  from  the  larger 
forms  of  R.  Cai)ipbcUiaua  by  the  much  more  prominent  scales,  the 
usually  less  attenuate  concolorous  margins,  the  different  epidermis, 
three  or  four  layers  of  cells  next  to  the  dorsal  surface  of  the 
thallus  in  R.  lauicllosa  being  commonly  echlorophyllose,  and  by 
the  entire  absence,  so  far  as  we  have  observed,  of  oil-bodies.  R. 
laincllosa  is  described  by  Herr  Stephanit  as  dioicous  and  our  in- 
vestigations on  Italian  material  point  toward  the  same  conclusion, 
though  we  have  been  able  to  see  no  antheridia. 

*  We  have  seen  an  authentic  specimen  of  this  from  the  Montagne  Herbarium 
through  the  courtesy  of  Mons.  Paul  Hariot  of  the  Museum  d'  Histoire  Naturelle  of  Paris. 
fBuU.  Herb.  Boiss.  6  .  341.  1898. 


28 


RiCCIACEAE 


The  specimen  collected  by  Professor  Campbell  near  Stanford 
University  on  May  i.  1896,  we  consider  the  type  of  the  species. 
This  specimen,  so  far  as  we  have  noticed,  does  not  contain  the  oil- 
bodies  referred  to  in  the  description,  but  such  occur  sparingly  in 
the  previously  collected  specimen  now  preserved  in  the  Underwood 
Herbarium  and  are  very  abundant  in  alcoholic  material  from  the 
same  locality  communicated  by  Professor  Campbell.  These  are 
present  also  in  the  San  Francisco  plant,  but  are  sparingly  developed 
or  absent  in  the  specimens  from  Fort  Ross  and  Ukiah.  Allusion 
is  made  in  our  remarks  under  R.  iiigrclla  to  the  occasional  be- 
havior of  the  contents  of  the  epidermal  cells  when  treated  with 
iron-haematoxylin. 

Plate  91,  Fjgs.  1-15.     Riccia  Campbelliana. 
I.  Plant,  dry,  natural  size. 

2  and  3.  Plants  from  soaked-out  dried  material,  natural  size. 

4.  Terminal  portion  of  thallus-segment,  X  lo. 

5-8.  Outlines  of  transverse  sections  of  thallus-segments,  X  12. 

9.  Outline  of  transverse  section  of  an  unusually  wing-margined  thallus-segment, 
X23. 

10.  Papilliform  elevation  of  antheridial  ostiole,  X  53- 

II  and  12.  Thin  transverse  sections  showing  epidermis  and  cells  containing  "  oil- 
bodies,"  X  53»  t^e  latter  section  from  an  older  part  of  the  thallus. 

13.  Cell  from  older  part  of  thallus  containing  "oil-body"  which  here  appears 
densely  granular  (yellowish-brown  in  the  alcoholic  material),  X  225.  The  adjacent 
cells  contain  chloroplasts  and  starch-grains. 

14  and  15.  Spores,  X  305- 

Fig.  I  drawn  from  specimen  collected  at  Fort  Ross,  March  15,  1896 ;  2,9,  14,  and 
15,  near  Stanford  University  (Professor  Campbell,  May  I,  1S96)  ;  3  and  5,  near  Stan- 
ford Universit}'  (Professor  Campbell,  Februar}*  6,  1896)  ;  4  and  8  (Professor  Campbell, 
1893,  herb.  Underwood)  ;  6,.  7,  and  10-13,  ^""om  alcoholic  material  sent  by  Pro- 
fessor Campbell. 

8.  Riccia  nigrella  DC.  Fl.  Fr.  5:  193.    1815.   Lindenb.  Nova 
Acta  Acad.  Caes.  Leop.- Car.  Xat.  Cur.  18  :  466.  //.  2<^.  1836. 
Riccia  minima  L.  p.p.  Sp.  PI.  i  139.  1753- 
Riccia  aggrcgata  XJnd^vw.  Bot.  Gaz.  19:  275.  1894. 
Thallus  at  first  subradiate,  later  commonly  forming  irregularly 
confluent  masses,  dark  green,  somewhat  polished  and  closely  retic- 
ulate above,  dark  purple  or  nearly  black  beneath,  with  transverse, 
semicircular,  blackish-purple,  nitent  scales,  which  do  not  exceed 
the  margin  ;  principal  divisions  2-8  mm.  long,  at  first  simple  and 
obovate,  finally  linear  or  linear-obcuneate  and  1-3  times  dichoto- 
mous  ;  terminal  lobes  i  — 1.5  mm.  wide,  obcuneate-oblong  or  ellip- 


RiCCIA 


29* 


tical-obovate,  obtuse  or  subacute,  narrowly  canaliculate,  the  mar- 
gins often  brownish,  acute,  entire,  slightly  membranous,  becoming 
rather  obtuse  toward  the  base,  often  erect-connivent  on  drying  ; 
width  of  transverse  sections  1-2.5  (mostly  1.5-2)  times  their 
height,  their  ventral  boundary  nearly  rectilinear,  slightly  convex 
toward  the  apex,  the  air-chambers  narrow  and  inconspicuous  ; 
epidermis  of  a  single  layer  of  cells,  these  lightly  protuberant,  sub- 
quadrate  in  vertical  section  and  often  broader  than  high,  25— 40^^ 
X  25-45  fj.,  filled  with  a  transparent  and  colorless  or  slightly  gru- 
mous,  somewhat  refringent  fluid,  collapsed  and  disintegrated  only 
in  the  oldest  parts  of  the  thallus  :  monoicous  :  antheridial  ostioles 
but  slightly  or  not  at  all  elevated  :  capsules  often  numerous, 
crowded  together  near  the  bases  of  the  segments,  hemispherical- 
protuberant  above  together  with  the  naked,  usually  light  brown, 
long  intact  covering,  this  commonly  with  a  small  purple  spot  about 
the  exserted  archegonium-neck ;  spores  brown,  becoming  darker 
and  more  opaque  with  age  but  usually  translucent  in  glycerine, 
60-78  a  in  maximum  diameter,  angular,  with  a  minutely  granular 
or  nearly  smooth,  sometimes  interrupted  margin  3-6  (J.  in  width,  the 
outer  face  with  irregularly  anastomose-reticulate  ridges,  lightly 
papillose  or  almost  smooth  in  profile,  the  inner  faces  usually  finely 
and  somewhat  regularly  reticulate,  with  thick-walled  meshes. 

Exsicc.    Hep.  Bor.-Am.  140b. 
Hep.  Am.  165. 

In  rather  dry  exposed  places,  especially  about  rocks,  often  ac- 
companied by  Riccia  tricJiocarpa  and  R.  minima. 

San  Francisco  :  near  the  Ocean  House  (Bolander) ;  hills  above 
the  Mission  Dolores  (Howe) ;  Wildcat  Canon,  near  Berkeley 
(Howe),  Fruit  Vale  (Miss  Byxbee)  ;  Mitchell's  Canon,  Mt.  Diablo- 
(Howe)  ;  Pasadena  (McClatchie)  ;  Twin  Oaks,  San  Diego  Co. 
(F.  W.  Koch). 

The  Pasadena  plant  (7v.  aggregata  Underw.)  differs  from  the 
European  and  from  the  specimens  of  the  San  Francisco  region  in 
the  more  regularly  and  finely  reticulate  outer  face  of  the  spore^ 
but  this  character  shows  considerable  variability  even  in  the  Pasa- 
dena specimen  and  hardly  justifies,  we  think,  a  specific  separation. 

The  dorsal  epidermis  in  R.  nigrclla  is  very  different  from  that 
of  any  other  Riccia  we  have  examined.  Its  cells  are  for  a  long 
time  turgid  with  a  nearly  transparent  fluid  which  takes  an  intense 
violet  stain  when  sections  of  the  thallus  are  treated  for  several 
hours  with  solutions  of  haematoxylin.     We  found  especially  fa- 


30 


RiCCIACEAE 


vorable  for  this  purpose  Heidenhain's  iron-haematoxylin  method 
(Zeit.  f.  wiss.  Mik.  9:  204.  1892.  Lee,  Microtomist's  Vade- 
Mecum,  175.  1896  [4th  ed.]).  The  only  European  specimen  of 
i?.  nigrella  (Husnot,  Hep,  Gall.  96)  which  we  have  subjected  to 
this  treatment  shows  the  same  peculiarity  even  in  the  soaked-out 
dried  material.  R.  Canipbelliana,  especially  in  forms  with  numer- 
ous oil-bodies,  occasionally  exhibits  something  which  seems 
similar  at  first  sight,  but  in  this  the  epidermal  cells  which  take  the 
haematoxylin  stain  occur  irregularly,  are  filled  with  a  more  gran- 
ular substance,  and  appear  to  be  simply  cells  with  oil-bodies  like 
those  found  elsewhere  in  the  thallus. 

Herr  Stephani,  who,  we  are  informed,  has  had  the  opportunity 
of  examining  the  original  specimen  of  Riccia  aggregaia  Underwood 
describes*  it  as  dioicous  ;  we,  however,  on  making  microtome  sec- 
tions of  the  same  type  material,  find  it  clearly  monoicoiis.  Herr 
Stephani  describes  the  ''frons"  of  R.  nigrella  as  subtriplo  latoir 
quam  crassa  "f  while  that  of  R.  aggrcgata  is  vix  diiplo  latior  qtiani 
alta,''  but  on  comparing  sections  of  French  and  Italian  specimens 
of  R.  nigrella  with  sections  from  corresponding  regions  in  R. 
aggregata,  we  find  the  differences  slight  and,  we  believe,  of  no 
specific  significance.  The  length  of  the  -frond,  the  number  of 
times  its  forking  is  repeated,  and  the  form  of  the  terminal  lobes 
are  variable  characters  in  both  the  Californian  and  the  European 
specimens,  and,  either  separately  or  combined,  afford  no  reliable 
grounds  for  specific  distinctions. 

9.  Riccia  Catalinae.    Underw.  Bot.  Gaz.  19:  275.  1894. 

Thallus  at  first  stellate  or  radiate,  often  forming  later  some- 
what irregularly  intertangled  masses,  rather  loosely  attached  to  the 
substratum,  light  green,  concolorous  below  and  furnished  with  a 
few  hyaline  very  rudimentary  scales,  the  younger  parts  extremely 
thin  and  exhibiting  suggestions  of  a  wide-meshed  areolation  as  if 
from  interior  air-chambers,  the  cells  of  the  dorsal  surface  large, 
thin-walled,  and  indistinctly  defined  (at  least  in  soaked-out 
herbarium  material),  the  older  parts  of  the  thallus  spongiose- 
alveolate  from  the  bursting  through  of  the  air-chambers  to  the 
upper  surface,  root-hairs  nearly  or  wholly  smooth  within  ;  the 


*  Bull.  Herb  Boiss.  6  :  342.  1898. 
f/-  c.  334. 


RiCCIA 


31 


principal  divisions  obcuneate-oblong  when  young,  becoming  linear, 
3—5  times  dichotomous,  6-15  mm.  x  1-2.5  mm.,  lightly  concave 
above,  with  thin,  naked,  slightly  ascending  or  plane  margins  ;  the 
terminal  segments  commonly  emarginate  :  monoicous  :  ostiolar 
papillae  cylindrical,  .06-  1 5  mm.  high  :  capsules  in  a  single  or 
irregularly  double  series,  rather  prominent,  the  covering  long  in- 
tact ;  spores  brown,  darkening  and  rather  opaque  with  age,  90— 
108  a  in  maximum  diameter,  distinctly  angular,  with  a  granulate- 
crenulate  or  minutely  tuberculate  margin  4-10  /7.  in  width,  the  prom- 
inent ridges  of  the  outer  face  rather  irregularly  anastomosing,  the 
areolae  of  the  middle  of  the  face,  when  well  defined,  usually  very 
large  (  20—45  diameter)  and  often  enclosing  an  isolated  tubercle 
or  a  free-ending  spur,  the  inner  faces  mostly  with  tubercles  and 
short  free  irregular  ridges,  sometimes  with  imperfect  areolae. 

On  wet  soil  in  the  bottom  of  a  deep  cafion,  Santa  Catalina 
Island,  A.  J.  ]\IcClatchie,  Sept.  15,  1893,  no.  441. 

Riccia  Catalinae  is  a  near  ally  of  R.  crystallina  L.  from  which 
it  differs  in  the  larger  size,  in  the  finally  3-5  times  dichotomous 
more  elongated  divisions,  which  are  less  closely  adnate  to  the  soil, 
in  the  larger  spores  (60-90  u.  in  the  various  European  specimens 
of  R.  crystallina  examined),  in  the  much  less  regular  areolation  of 
the  spore -faces,  and  in  the  considerably  larger  size  of  the  central 
meshes  of  the  outer  face,  when  these  are  well-formed. 

The  thallus  is  so  thin  and  so  largely  occupied  by  air-chambers 
that  we  have  been  unable,  from  the  dried  material,  to  obtain  any 
satisfactory  sections. 

Plate  91,  Figs.  16-20.    Ricca  Catalinae. 

16.  Portion  of  thallus,  dry,  natural  size. 

17.  End  of  young  terminal  lobe,  dorsal  surface,  showing  position  of  the  internal 
lamellae  which  bound  the  air-chambers,  X  23.    From  a  soaked-out  fragment. 

18.  Dorsal  view  of  thallus  in  the  older  parts,  X  23. 

19.  Spores,  X  305- 

10.  Riccia  crystallina  L.  Sp.  PI.  1138.  1753. 

Thallus  in  closely  attached  orbicular  or  suborbicular  rosettes 
5-1  5  mm.  in  diameter,  light  green  on  both  sides  or  often  tinged  with 
yellow,  spongiose  with  age  and  vesiculose-alveolate;  the  principal  di- 
visions mostly  lobate-obcordate  or  obcuneate,  or  sometimes  becom- 
ing broadly  linear  and  1-3  times  furcate,  the  segments  often  close 
and  somewhat  crowded,  terminal  lobes  1.5-2.5  mm.  wide,  transverse 
sections  2.5-4  times  broader  than  high,  margins  naked,  subobtuse, 
slightly  elevated  on  drying  or  nearly  plane,  ventral  scales  wanting  ; 


32 


RiCCIACEAE 


air-chambers  large,  epidermis  (in  dried  material)  disorganized  and 
indistinct :  monoicous  :  ostiolar  elev^ations  prominent,  cylindrical, 
.15— .27  mm,  X  .05— .07  mm.:  capsules  rather  deeply  immersed, 
scarcely  prominent  even  at  maturity  ;  spores  soon  dark-brown, 
becoming  nearly  opaque,  60-90  in  maximum  diameter,  distinctly 
angular,  with  a  narrow,  granulate-papillate,  sometimes  almost  de- 
ficient, margin  (mostly  3—5  //  broad),  the  convex  face  with  strong 
anastomose-reticulate  ridges,  about  7  or  8  imperfectly  formed 
areolae  measuring  its  diameter,  those  near  the  middle  usually 
larger  ( 1 5-30//)  and  sometimes  enclosing  a  free-ending  spur,  the 
plane  faces  marked  with  short  free  or  irregularly  anastomosing 
ridges,  the  mature  more  opaque  spores  appearing  tuberculate- 
papillate. 

On  the  banks  of  a  stream  upon  a  rocky  hillside,  southern 
slope  of  the  Dixey  Mts.,  Lassen  Co.  (M.  S.  Baker  and  F.  P. 
Nutting,  July  2,  1894).  We  refer  here  also  a  specimen  in  herb. 
Underwood,  collected  near  San  Francisco  by  Mrs.  Katharine 
Brandegee  in  1892,  which,  in  the  more  solid — less  conspicuously 
vesiculose-alv^eolate — thallus,  approaches  the  closely  allied  R. 
Brandegei  {{vom  the  southern  extremity  of  the  peninsula  of  Lower 
California). 

The  Lassen  County  plant  seems  scarcely  to  differ  from 
European  specimens  of  R.  crystallina  unless  possibly  in  the  greater 
opacity  of  the  spores  when  fully  mature  and  in  their  somewhat 
narrower  margin. 

Riccia  crystallina  is  listed  in  Dr.  Bolander's  Catalogue  of  the 
Plants  growing  in  the  Vicinity  of  San  Francisco." 

Riccia  Frostii  Aust.  (Bull.  Terr.  Bot.  Club,  6:  17.  1875),  orig- 
inally described  from  Nevada  and  Colorado,  and  since  found  to  be 
quite  widely  distributed  in  the  Rocky  Mountain  region  and  to  extend 
as  far  easward  as  the  Mississippi  Valley  is  to  be  expected  in  Cali 
fornia.  It  can  readily  be  distinguished  from  any  of  the  Californian 
species  here  described  by  the  small  narrowly  margined  spores,  45-58 
/7.  in  maximum  diameter,  marked  almost  uniformly  over  the  en- 
tire surface  by  numerous  short,  delicate,  wavy  ridges  which  rarely  an- 
astomose ;  the  thallus,  which  is  thin  in  texture,  with  a  fibrous-reticu- 
late and  minutely  foveolate  surface,  becoming  lacunose  and  spongy  in 
the  older  parts,  forms  flat,  dark-  or  grayish-green  rosettes,  8-15  mm.  in 
diameter,  closely  adherent  to  the  soil,  with  numerous  narrow  usually 
crowded  divisions  .5-1.5  mm.  wide;  the  covering  of  the  abundant 


RiCCIA 


33 


capsules  is  soon  ruptured,  exposing  the  spores.  R.  Frostii  is  dioicous, 
the  male  plant  (^R.  Watsoni  Aust.)  being  smaller  and  more  or  less 
purple  in  color,  the  slenderly  cylindrical  ostioles  elevated  about .  i  mm. 

RicciA  FLUiTANS  L.  Sp.  PL  1139.  1753- 

Thallus  thin,  green,  floating  or  immersed,  often  forming  inter- 
tangled  mats,  destitute  of  rhizoids,  or  becoming  terrestrial  and  attached 
by  root-hairs,  repeatedly  dichotomous,  the  segments  narrowly  linear 
(.5-1.5  mm.  wide,  the  entire  thallus  1-5  cm.  long),  the  median  axis 
thickened,  the  mostly  flat  margins  only  2-  or  3-stratose,  air-chambers 
large,  oblique,  usually  elongated,  communicating  with  the  dorsal  sur- 
face by  small  pores  surrounded  by  slightly  modified  epidermal  cells  : 
capsules  (found  only  in  the  terrestrial  or  semiterrestrial  conditions)  pro- 
tuberant below  with  the  enclosing  tissues  of  the  thallus,  the  spores 
usually  set  free  ventrally,  these  yellowish-brown,  translucent,  70-75  ij. 
in  maximum  diameter,  with  a  margin  3-6  ,a  broad,  the  outer  face  with 
rather  smooth  meshes  12-15  p.  in  width,  inner  faces  less  distinctly 
areolate. 

This  species  was  collected  by  Messrs.  Coville  and  Funston  in  Vegas 
Valley,  Lincoln  Co.,  Nevada  (Death  Valley  Expedition,  no.  400)  and 
probably  occurs  within  the  limits  of  California. 

RICCIOCARPUS  Corda;  Opiz,  Beitr.  I  :  651.  1829.^ 

Plants  lemna-like,  floating  or  finally  attached  to  the  soil  by  the  sub- 
sidence of  the  water,  dark  green,  sometimes  purple-margined,  dichoto- 
mous, mostly  2-4-lobed,  with  a  very  pronounced  median  furrow,  the 
lobes  obcordate  or  obovate,  the  ventral  surface  bearing  numerous  long, 
pendant,  linear  or  linear-lanceolate,  reddish-violet,  dentate  scales ; 
colorless  ventral  layer  of  thallus  much  reduced,  the  chlorophyll-bearing 
layer  composed  of  large  irregularly  polyhedral  chambers  separated  by 
mostly  unistratose  lamellae  ;  epidermis  with  small  but  distinct  stomata. 
Dioicous.  Antheridia  in  an  elongated  ridge-like  androecium  situated 
in  the  median  furrow.    Capsule  immersed. 

The  only  recognized  species  is 

RicciocARPUs  NATANS  (L. )  Corda,  /.  c. 

Riccia  7iataiis  L.  Syst.  Nat.  1339.    1759  [Ed.  10]. 

Plants  about  i  cm.  long,  the  lobes  4-9  mm.  wide  :  spores  45-55  p-, 
papillate-areolate,  dark-brown  at  maturity,  the  areolae  becoming  obscure 
and  the  papillae  more  prominent. 

Rarely  found  with  capsules.  This  species  has  not  yet  been  collected, 
so  far  as  is  known,  within  the  limits  of  the  state,  but  it  is  found  in  Ore- 
gon and  is  likely  to  occur  in  California. 

*  It  would  appear  from  a  statement  of  M.  Auguste  Lejolis  (Mem.  Soc.  nation. 
Sci.  nat.  et  math.  Cherbourg,  29:  167.  1894)  that  a  separate  of  Corda's  "Genera 
Hepaticarum  "  was  published  in  1828 — in  advance  of  its  appearance  in  Opiz'  Bei- 
trage  zur  Naturgeschichte. 


34 


Marchaxtiaceae 


Family  II.    M ARCH ANTI ACE AE. 

Chlorophyll-bearing  layer  of  the  thallus  (exc.  in  Diiuwrtiera) 
with  well-developed  air-chambers,  these  in  most  cases  containing  a 
loose  somewhat  filamentous  assimilative  tissue  or  soon  divided  into 
smaller  secondary  chambers  by  chlorophyll-containing  lamellae. 
Stomata  (exc.  in  Dmnortierd)  always  present,  simple  (/.  the  pore 
surrounded  by  four  or  more  somewhat  modified  epidermal  cells, 
all  in  a  single  stratum)  or  dolioform  (/.  e.,  the  pore  surrounded  by 
a  barrel-shaped  wall,  several  cells  in  height,  which  projects  into 
the  underlying  air-chamber).  Sexual  organs  nearly  always  ag- 
gregated, often  borne  on  long-stalked,  capituliform  receptacles. 

Capsule  short-stalked,  breaking  through  the  calyptra  at  ma- 
turity, dehiscing  irregularly  or  by  a  special  lid,  rarely  valvate. 
Spores  always  accompanied  by  sterile  cells,  which  (exc.  in  the 
Corsi?iioideae'^)  are  developed  as  elaters. 

Key  to  the  Genera  of  tbe  Itlarcliautiaceae. 

Sporogonium  single,  underneath  the  apical  margin  of  the  thallus,  enclosed  by  an  invo- 
lucre consisting  of  two  rigid,  brown  or  dark  purple  valves  which,  previous  to  the  ma- 
turity of  the  capsules,  are  united  in  a  vertical  keel-like  suture.  2.  Targionia. 
Sporogonia  in  groups  on  stalked  receptacles. 

Stomata  dolioform  ;  antheridia  on  a  disk-shaped  receptacle  ;  9  receptacles  (in 
our  species)  with  8-ll  finger-like,  usually  decurved  rays;  gemmae  in  dorsal, 
cup-shaped,  dentate-margined  receptacles.  lo.  Marchantia. 

Stomata  of  the  thallus  simple 

Always  gemmiferous,  the  gemmae  in  dorsal,  crescentic,  entire-margined  recep- 
tacles.   Found  in  or  about  greenhouses.  9.  Luniilaria. 
Never  with  gemmae-cups. 

Thallus  large  (5-25  cm.  X  -75-1 -5  cm.)  areolae  very  distinct,  the  ele- 
vated stomata  easily  visible  without  magnification.  The  uppermost 
cells  of  the  thin,  sharp H'  defined,  chlorophyll-bearing  layer  each  with  a 
conical  or  long-cylindrical  colorless  beak.  8.  Conocephalum. 

Thallus  smaller  (less  than  5  cm.  long  in  the  Californian  species),  cells  of 
the  chlorophyll-bearing  stratum  never  with  a  colorless  beak. 
Sporogonia  each  surrounded  by  a  membranous,  white  or  sometimes 
violet  pseudoperianth,  this  at  first  somewhat  conical,  usually  much 
exserted,  finally  more  or  less  completely  divided  longitudinally 
into  3-18  narrow  segments.  7.  Asterella. 

Pseudoperianths  wanting  or  ver}'  rudimentary. 

9  branch  arising  from  the  dorsal  surface  of  thallus,  the  peduncle 
without  a  root-hair  furrow;  stomata  normally  stellate  (/.  e.,  the 
cells  bounding  the  pore  have  the  radial  walls  strongly  thick- 
ened). 3.  Clrc'ca. 


*  This  subfamily  is  not  represented  in  North  America,  so  far  as  is  now  known. 


Targioxia 


35 


9  branch  arising  from  apex  of  thallus  or  thallus-lobe,  the  pe- 
duncle with  a  single  root-hair  furrow,"^  stomata  not  stellate. 
Thallus  thin  ;  air-chambers  large  and  empty,  separated 
by  mostly  unistratose  lamellae  ;  antheridia  immersed  in 
the  thallus  in  an  elongated,  single  or  irregularly  double 
median  row  just  back  of  the  peduncle,  ostioles  incon- 
spicuous ;   9  receptacle  finally  disciform. 

6.  Cryptojiiitrium. 

Thallus  somewhat  firm  and  rigid,  the  primary-  air-chambers 
soon  filled  by  secondary  walls  ;   9  receptacle  hem- 
ispherical or  subconical. 
Capsule  at  maturity  completely  filling  the  broadly 
campanulate  involucre  and  protruding. 

5.  Gfimaldia. 

Involucre  conchoid-bivalved,  its  cavity  only  partially 
filled  by  the  capsule.  4.  Reboulia. 

2.  TARGIONIA  L.  Sp.  PL  1136.     1753.    Ex  xMich.  Nov.  PL 
Gen.  3.  //.  J.  1729. 

Thallus  usually  rather  thick  and  coriaceous,  simple  or  once  or 
twice  dichotomous,  innovating  at  apex  or  often  latero-ventrally, 
broadly  costate,  areolae  mostly  indistinct,  stomata  simple,  the 
white -margined  pores  more  or  less  conspicuous  ;  chlorophyll-bear- 
ing layer  sharply  defined,  the  chambers  filled  with  branched  con- 
ferva-like "  filaments,  epidermal  cells  thick,  their  walls  with  trigones  ; 
ventral  scales  reaching  to  margin  or  much  reduced.  Gemmae 
none.  Androecium  small,  disciform,  terminating  a  short  latero- 
ventral  shoot.  Archegonia  in  a  group  of  several  arising  just  back 
of  the  apex  of  the  thallus,  becoming  apparently  ventral  by  the 
over-arching  of  the  surrounding  dorsal  tissues,  without  a  special 
receptacle,  a  single  archegonium  of  the  group  ordinarily  maturing 
a  sporogonium,  this  soon  surrounded  by  a  somewhat  laterally 
compressed,  obovoid,  subglobose,  or  hemispherical  2-valved  invo- 
lucre, the  anterior  margins  of  the  latter  uniting  to  form  a  median 
vertical  keel-like  suture,  finally  gaping  widely  and  exposing  the 
capsule,  the  valves  rigid,  entire-margined,  brown  or  dark  purple, 
at  times  iridescent.  Pseudoperianth  none.  Sporogonium  con- 
sisting of  a  subglobose  capsule,  a  very  short  pedicel,  and  a  bulbous 
foot ;  capsule  dehiscing  irregularly,  its  wall  of  a  single  layer  of 
cells  with  annular  or  spiral  thickenings.  Spores  opaque,  nearly 
spherical,  the  outer  membrane  thrown  into  ridges  and  wart-like 
folds,  loosely  investing  the  firmer  inner  coat  and  occasionally  cadu- 
cous, the  entire  surface  minutely  and  irregularly  reticulate  or  granu- 
late.   Elaters  2-4-spiral,  often  branched. 


*  See  footnotes  on  following  pages  under  generic  diagnoses  of  Reboulia  and  GtI- 
maldia  and  also  remarks  upon  Cryptomitriiwi  in  Erythea,  5  :  87.  1897. 


36 


Marchantiaceae 


I.  Targionia  hypophylla  L.  /.  c. 

Thallus  mostly  simple,  sometimes  dichotomous,  obovate  to 
sublinear,  commonly  linear-obcuneate,  progressing  chiefly  by 
latero-ventral  or  apical  innovations,  4-20  mm.  long,  2-5  mm.  in 
maximum  width,  rounded  or  slightly  emarginate  at  apex,  reddish- 
brown  or  dark-purple  beneath,  the  margins  usually  involute  or 
erect-connivent  on  drying,  colorless  stratum  of  about  25  layers  of 
cells  in  axile  parts,  rather  suddenly  reduced  to  the  ventral  epider- 
mis toward  the  lateral  margins  ;  scales  densely  imbricate,  the  su- 
perior extending  to  the  margin,  often  abruptly  contracted  to  a 
narrow,  sometimes  twisted,  subentire  or  ciliate-dentate  apical  ap- 
pendage :  monoicous  :  spores  brown,  58-80/^;  elaters  180-330// 
x6-I2/^  subacute  or  obtuse. 

Exsicc.  Hep.  Bor.-Am,  137  (as  Targionia  MicJielii  Corda). 
Hep.  Am.  159. 

On  shaded  banks,  mostly  in  rocky  soil.  Common,  particularly 
in  the  Coast  Range  Mountains  from  San  Diego  to  Humboldt 
County,  and  extending  northward  to  Vancouver  Island  (Macoun). 
Berkeley ;  San  Francisco  ;  Sausalito,  Mill  Valley  ;  Sonoma  (F» 
T.  Bioletti),  Duncan's  Mills ;  Ukiah  (746)  and  Big  River  Boom 
(681),  Mendocino  Co.;  Blue  Lake  (997),  Humboldt  Co.;  Oak 
Run,  Shasta  Co.  (Baker  and  Nutting);  Jackson,  Amador  Co. 
(George  Hansen) ;  Ojai,  Santa  Barbara  Co.  (W.  F.  Hubby) 
Nordhoff,  Ventura  Co.  (Miss  Jacqueline  K.  Newton)  ;  Pasadena 
(A.  J.  McClatchie) ;  San  Bernardino  (S.  B.  Parish) ;  San  Diego 
(W.  G.  Farlow)  and  Twin  Oaks  (F.  W.  Koch),  San  Diego  Co.; 
Surprise  Cafion,  Inyo  Co.  (Coville  and  Funston,  Death  Valley  Ex- 
pedition, no.  619) ;  also,  by  the  last  named  collectors,  in  Lincoln 
Co.,  Nevada.    First  found  in  California  by  Dr.  Bolander. 

3.  CLEVEA  Lindb.  Not.  ur  Sallsk.  pro  Fauna  et  Flora  Fenn.  9  : 

289.  1868. 

Thallus  usually  small,  simple  or  once  dichotomous,  sometimes, 
innovating  from  the  apex,  distinctly  areolate,  the  areolae  but 
slightly  elevate-pulvinate,  the  cells  bounding  the  stomata  with 
(typically)  strongly  thickened  radial  walls  ;  air-chamber  layer  of 
numerous  empty  cavities  with  mostly  unistratose  dissepiments, 
these  cavities  appearing  in  several  series  in  the  median  portions  iiva 
cross  section,  occupying  the  greater  part  of  the  axial  thickness  of 
the  thallus  and  composing  the  whole  of  the  wings,  being  gradu- 


Clevea 


37 


ally  or  rather  abruptly  reduced  to  a  single  series  toward  the  thin 
thallus-margins  ;  postical  scales  numerous,  hyaline,  at  least  at  their 
apices,  or  rarely  purple  throughout,  obtuse,  acute  or  long-acuminate, 
the  anterior  usually  exceeding  the  margin  of  the  thallus  and  often 
conspicuous.  Dioicous.  Antheridia  immersed,  scattered  near  the 
median  line  of  the  thallus,  with  slender  subcylindrical  ostiolar  pa- 
pillae. C  branch  from  the  median  dorsal  surface  of  the  thallus,  com- 
monly arising  at  a  considerable  distance  back  of  the  apex,  single  or 
in  a  linear  series  of  1-4.  Peduncle  subpellucid,  without  a  root-hair 
furrow.  9  receptacle  consisting  simply  of  the  4  (-1)  obovoid 
somewhat  downwardly  inclined  involucres,  these  lightly  com- 
pressed laterally  and  bilabiate  about  to  the  middle  by  a  vertical 
cleft,  affixed  directly  to  the  top  of  the  peduncle  or  slightly  connate 
at  the  bases.  Pseudoperianth  none  ;  capsule  nearly  sessile  and 
included  or  subexserted  by  a  somewhat  elongate  pedicel,  its  walls 
on  dehiscence  cleft  to  the  middle  or  below  into  3-8  irregular 
valves,  the  cells  with  numerous  annular  or  spiral  thickenings. 
Spores  yellowish,  red,  or  bj  own,  densely  covered  with  rather  large 
obtusely  conical  papillae.    Elaters  2-4-spiral. 

I.  Clevea  hyalixa  (Sommerf)  Lindb.  Not.  ur  Sallsk.  pro  Fauna 
et  Flora  Fenn.  9  :  291.  1868. 

Marchantia  hyalinaSommQvL  Mag.  Naturv.  II.  i  :  284.  1833 
{Jide  Lindberg). 

Plagiochasina  erythrosperina  Sulliv.;  Aust.  Proc.  Acad.  Nat. 
Sci.  Philad.   1869  •  229.  1869. 

Thallus  strongly  concave  or  canaliculate  above,  narrowly  ob- 
long to  obovate,  obcordate  at  the  apex,  simple  or  once  forked,  5— 
I  5  mm.  long,  2-6  mm.  in  greatest  width,  scales  mostly  numerous 
and  persistent,  conspicuously  projecting,  with  rare  exceptions,  and 
the  anterior  often  inflexed,  hyaline,  sometimes  purple  at  base, 
rarely  purple  throughout,  acute  or  acuminate,  the  occasionally 
subtubulose  points  with  crenulate  margins :  peduncle  usually  sin- 
gle, 2-15  mm.  high  ;  9  receptacle  2.5-4  mm.  broad,  with  long 
and  numerous  white  or  purplish  paleae  beneath  :  capsule  dehiscing 
by  3-6  irregular  valves,  the  wall-cells  with  annular  thickenings  ; 
spores  44-60  reddish-brown,  rarely  verging  toward  yellow  or 
darkening;  elaters  120-300//  long,  8-15  a  in  greatest  width,  often 
bispiral  at  the  extremities,  in  the  middle  mostly  tri-,  rarely  quadri- 
spiral. 

About  rocks  in  mountainous  regions,  especially  northward. 
<  Near  a  lake  on  the  trail  to  the  White  Chief  Mine,  1 100  ft.  above 
Mineral  King,  Tulare  Co."  (Coville  and  Funston,  Death  Valley 


38 


Marchantiaceae 


Expedition,  Aug.  6,  1891,  no.  i  5  1 1);  also,  by  the  same  collectors^ 
''near  White  Chief  Mine"  (no.  1528). 

lb.  Clevea  hyalina  Calif ornica  var.  nov. 

Santeria  linibata  *  Aust.  /.  /.  Proc.  Acad.  Nat.  Sci.  Philad. 
1869  :  229.  1869. 

Thallus  often  once  dichotomous,  with  strongly  pronounced 
radial  ridges  or  undulations  on  the  ventral  surface  toward  the  thin 
margin,  very  distinctly  areolate,  pores  in  the  median  parts  large 
and  elliptical,  often  50-150/^  in  length,  the  radial  walls  of  the 
bounding  cells  unthickened,  the  pores  near  the  margin  smaller, 
with  the  radial  w^alls  of  the  surrounding  cells  more  or  less  clearly 
incrassate,  scales  purple  or  slightly  decolorate  at  their  extremities, 
exceeding  the  margin  :  peduncle  delicate,  slender,  25-32  mm.  high, 
with  few  and  rather  inconspicuous  paleae  at  the  apex  ;  involucres 
for  the  most  part  strongly  discrete  :  capsule  finally  subexserted,  the 
pedicel  attaining  a  length  of  2  mm. 

Doolan's  Creek  Canon,  near  Ukiah,  May  13,  1896  (Howe,  no, 
765);  a  specimen  collected  by  A.  J,  McClatchie  on  the  trail  to 
Wilson's  Peak,  San  Gabriel  Mts.,  April,  1893,  has  more  in  com- 
mon with  this  variety  than  with  the  typical  form,  though  the  pedun- 
cles are  shorter,  the  capsules  sometimes  subsessile,  the  areolation 
is  less  distinct,  and  the  guard-cells  of  the  stomata  have  more 
strongly  thickened  radial  walls.  Professor  McClatchie's  plant 
differs,  too,  from  both  the  Ukiah  specimen  and  the  typical  C.  hya- 
lina in  the  very  slender  inconspicuous  scales  with  almost  filiform 
points. 

The  Ukiah  plant  is  a  remarkable  one,  and  it  is  with  misgivings 
that  we  refer  it  to  Clcvca  Jiyalina  as  a  mere  variety,  but  the  Wil- 
son's Peak  specimen,  though  somewhat  aberrant  in  scale-characters, 
bridges  over  the  gap  in  respect  to  the  stomata  and  in  some  other 
particulars,  as  noted  above.  The  form  possibly  needs  comparison 
with  Clevca pediccllata  {PlagiocJiasma pcdicellatus  Griff  Not.  ad  PI. 
Asiat.  1:  pi.  68.  f.  2.  1849)  from  Afghanistan,  though  the 
latter  would  appear  from  figure  and  description  to  differ  at  least  in 
8-  or  9-  instead  of  4-  or  5-valved  capsule. 

Mr.  Austin,  who  published  Plagiochasina  crythrospejina  "  from 
a  plant  thus  labeled  (with,  however,  a  query-mark  after  the  ''n.  sp."), 

*  For  remarks  on  the  use  of  the  name  limbata^  see  footnote  under  Asterella  Cali- 

fornica. 


Reboulia 


39 


in  the  Sullivant  collection,  was  apparently  the  first  to  recognize 
that  the  name  was  a  synonym  for  Clevea  Jiyalina,  as  is  attested  by 
a  pencil  inscription  in  Austin's  handwriting  on  the  pocket  contain- 
ing the  original  specimen,  now  in  the  herbarium  of  Harvard  Uni- 
versity. The  several  plants  since  collected  in  the  mountainous 
regions  of  the  western  United  States  and  British  America,  and 
identified  with  this  species  of  Sullivant's,  though  quite  variable, 
cannot,  we  think,  be  satisfactorily  distinguished  from  the  Clevea 
Jiyalina  of  Europe.  The  spores,  however,  often  incline  more  to 
red,  and  the  thallus  is  sometimes  proportionally  broader. 

Of  true  Sauteria,  we  have  seen  the  following  three  American 
specimens,  probably  all  referable  to  Saiiteria  alpina  (Nees  &  Bisch.) 
Nees  :  "under  rocks.  Sulphur  Mts.,  Banff,  J.  Macoun,  i6  July, 
1891,  no.  345  ;"  under  rocks.  Lake  Agnes,  7000  ft.,  J.  Macoun, 
19  Aug.,  1 89 1,  no.  364"  ;  St.  Paul  Islands,  Pribiloff  Group,  Beh- 
ring  Sea,  1897  (Mr.  Trevor  Kincaid). 

4.  REBOULIA  Raddi  (as  Reboiiillia)  Opusc.  scientif  di  Bologna, 

2:  357.  1818.* 
Aster  clla  Pal.  de  Beau  v.  /.  /.  Encyc.  ]Meth.  Bot.  Suppl.  i  : 
502.     1 8 10. 

Thallus  coriaceous  and  rigid,  dichotomous,  usually,  also,  inno- 
vating at  the  apex,  broadly  costate,  scale-bearing  and  dark-purple 
or  brownish  beneath,  the  dorsal  surface  without  evident  areolation, 
provided  with  small  scattered  simple  stomata,  epidermal  cells  with 
trigones  :  chlorophyll-bearing  layer  strongly  developed,  passing 
gradually  into  the  colorless  stratum,  the  latter  limited  to  the  region 
of  the  costa,  cells  with  oil-bodies  occurring  here  and  there  in  both, 
air-chambers  at  first  simple,  their  limits  afterwards  obscured  by 
more  or  less  numerous  secondary  walls  ;  root-hairs  numerous, 
colorless.  Gemmae  none.  Androecium  sessile  or  somewhat 
sunken,  in  the  median  line  of  the  thallus,  disciform,  suborbicular  or 
with  a  lunate  sinus  in  front,  often  surrounded  by  a  few  narrow 
scales.  O  receptacle  conical  or  hemispherical,  becoming  at  times 
nearly  plane  above,  paleaceous-barbate  beneath,  containing  airr 
chambers  and  furnished  with  dolioform  stomata,  cleft  about  to  the 
middle  into  1-6  (commonly  4  or  5  )  thick,  often  spreading  lobes, 
the  membranous  ventral  margins  of  these  forming  the  conchoid- 
bivalved  involucres,  each  of  the  latter  enclosing  a  single  sporo- 


*  This  citation  is  as  given  by  authors  We  have  been  unable  to  see  the  original 
paper. 


40 


M  ARCH  AXTIACEAE 


cronium.  Pseudoperianth  wanting.  Peduncle  from  the  apex  of  the 
thallus-lobe,  with  a  single  root-hair  furrow,  destitute  of  assimila- 
tive tissue,  surrounded  at  the  base  by  a  few  narrow,  whitish  or 
purple  paleae,  the  receptacle  conspicuously  involucrate  when  young 
and  sessile.  Sporogonium  consisting  of  a  subglobose  capsule,  a 
scarcely  perceptible  pedicel,  and  a  large  foot.  Capsule  only  par- 
tially filling  the  involucral  cavity,  yellowish-green,  the  apical  por- 
tion of  its  wall  falling  away  in  fragments  at  dehiscence,  leaving  be- 
hind a  subhemispherical  urn  with  an  erose-dentate  margin,  the 
cells  without  annular  or  spiral  fibers,  those  in  the  apical  region 
with  trigones  or  sometimes  exhibitmgj  irregular  nodulose  thicken- 
ings in  the  lateral  walls.  Spores  }'ellow  or  brownish,  more  or  less 
distinctly  tetrahedral,  all  the  faces  with  rather  large  often  irregular 
areolae,  the  angles  pellucid-margined,  membrane  finely  granulate. 
Elaters  2-4-  (mostly  3-)  spiral. 

I.  Rebol'lia  hemisphaerica  (L.)  Raddi,  /.  c. 

Marchantia  JicmispJiacrica  L.  Sp.  PI.  1138.  1753. 

Aster ella  JieinispJiaerica  Pal.  de  Beau  v.  Encyc.  Meth.  Bot. 
Suppl.  I  :   502.     1 8 10. 

Thallus  6-30  mm.  long,  rather  pale  green  above,  the  segments 
emarginate  at  apex,  obcuneate  or  obcordate,  less  commonly 
elongated,  margins  undulate-crenulate :  monoicous  :  peduncle 
mostly   1.5-3  ^^"^   length,  rarely  longer:    spores   65-80 /i ; 

elaters  300—450  n.  long,  10—12  //  in  maximum  width,  the  apices 
subobtuse. 

A  cosmopolitan  species,  but  the  only  Californian  station  we  can 
cite  is  Folsom  (Katharine  Brandegeej. 

5.  GRIMALDI At  Raddi,  Opusc.  scientif  di  Bologna,  2:  356. 

i8i8.: 

Thallus  somewhat  firm  and  rigid,  broadly  costate  or  some- 
times sharply  carinate,  dichotomous  and  often  innovating  at  the 

*  Schift'ner  in  his  key  to  the  Marchantiaceous  genera  in  Engler  <S:  Prantl,  Nat. 
Pflanzenfam.  i^;  25,  denies  a  "  \Vurzelrinne  "  to  A'eOouIio,  but  we  rind  it  always  pres- 
ent, as  described  by  Xees,  Linipricht,  and  others,  both  in  European  and  American  forms 
of  the  plant. 

t  We  are  not  unmindful  of  the  fact  that  Grinialdia  is  said  to  have  been  used  by 
Schrank  in  1805  and  1808  as  a  generic  name  for  certain  species  of  phanerogams,  which 
by  later  writers  have  been  almost  universally  referred  to  Cassia  or  to  Chamaecrista 
(Breyne,  1678;  Moench,  1794).  There  can,  therefore,  be  no  doubt  that  a  literal 
carrying  out  of  Article  W .  of  the  Rochester  Code  would  call  for  the  disappearance  of 
Gritnaldia  from  among  the  names  of  genera  of  Hepaticae,  and  indeed  from  all  l>otanical 
literature  save  the  strictly  historical,  unless,  perchance,  some  one  should  rind  grounds 
for  recognizing  again  the  Grinialdia  of  Schrank  as  a  genus.  But  until  some  one  actu- 
ally proposes  to  revive  the  older  Grinialdia — if  that  time  ever  comes — it  seems  to  us  to 
be  conducive  to  clearness  to  retain  in  use  the  only  "  Grinialdia  "  that  has  been  gener- 
ally known  to  botanists  for  the  past  eighty  years. 

\  See  footnote  under  Reboulia. 


Grimaldia 


41 


apex  or  latero-ventrally,  mostly  dark  purple  beneath,  dorsal  sur- 
face ver\'  indistinctly  areolate,  furnished  with  simple  inconspicuous 
stomata,  epidermis  thick,  the  walls  of  its  cells  more  or  less  thick- 
ened at  the  angles,  chlorophyll-bearing  layer  deep,  passing  gradu- 
ally into  the  colorless  stratum,  somewhat  inflated  cells  with  oil- 
bodies  often  occurring  in  both,  the  primary  air-chambers  soon 
filled  by  secondary  w^alls  :  scales  extending  to  the  margin  or  pro- 
jecting and  inflexed  at  the  thallus-apex,  purple  or  vinous-red, 
sometimes  conspicuously  hyaline-pointed  or  nearly  colorless 
throughout.  Gemmae  none.  ]\Ionoicous,  dioicous,  or  polyoicous. 
Antheridia  immersed  near  the  median  line  of  the  thallus,  some- 
times at  about  the  middle  of  a  segment,  but  mostly  forming  a 
papillate  androecium  near  an  apical  sinus,  the  androecium  sur- 
rounded by  the  slightly  elevated  adjacent  tissues  or  wholh'  desti- 
tute of  a  special  involucre,  covered  occasionally  by  the  inflexed 
thallus  -margins.  O  receptacle  with  a  single  archegonium  in  each  of 
the  3  or  4  short  lobes,  maturing  1—4  sporogonia,  finally  long-stalked, 
hemispherical-umbonate  or  subconoidal,  commonly  papillate  abo\e, 
the  lobes  at  maturity  mostly  directed  downward,  rarely  somewhat 
spreading,  often  obscurely  defined.  Peduncle  from  apex  of  the 
thallus  or  from  a  small  latero-ventral  innovation,  with  a  single 
root-hair  canal, more  or  less  paleaceous-involucrate  at  base, 
clothed  underneath  the  receptacle  with  pendant,  often  white,  pa- 
leae,  these  sometimes  obsolete  or  wholly  wanting.  Involucres 
broadly  campanulate,  continuous  with  the  lobes,  the  margins  re- 
pand.  Pseudoperianth  none  or  rudiamentary.  Capsule  globose, 
completely  filling  the  involucres  and  protruding,  circumscissile 
above  the  middle,  cells  of  its  wall  destitute  of  annular  thickenings 
or  with  traces  of  such  at  the  margin  of  the  thicker  operculum, 
pedicel  very  short.  Spores  yellow,  brown  or  dark  violet-purple, 
tuberculate,  rugose,  or  sometimes  rather  regularly  areolate. 
Elaters  2-4-spiral,  often  somewhat  attenuate  toward  the  extremi- 
ties. 

I.  Grimaldia  Californica  Gottsche ;  Underw.  Bot.  Gaz.  13: 
1 14.  1888.  Bolander,  Gal.  Med.  Gaz.  1870  :  184  (40)  (name 
only).  1870. 

Grimaldia  Californica  Steph.  Bull.  Herb.  Boiss.  6  :  794.  1898. 
Thallus  linear  or  spatulate-oblong,  6-20  mm.  x  1.5-4  mm., 

^  The  peduncle  of  Gi-imaldia  is  said  by  Schififner  in  his  key  to  the  genera  of  the 
^larchantiaceae  to  be  without  a  "  Wurzelrinne  "  (Engler  &  Prantl,  Nat.  Pflanzenfam. 

:  25),  but  it  is  surely  present  in  G.  Calif oi'iiica,  G.  pilosa,  G.  niidrogyna  [Mar- 
chajitia  androgyiia  L.  pro  parti:  maxima,  Gi-imaldia  dichotoma  Raddi)  and  in  Amer- 
ican specimens  of  Grimaldia  fragrans  ;  it  seems,  however,  to  be  sometimes  only 
rudimentary  in  European  specimens  of  G.  fragrans,  though  attributed  to  the  species 
without  reservation  by  Nees  and  by  Limpricht. 


42 


Marchantiaceae 


dichotomous,  or  progressing  by  latero-ventral,  very  rarely  apical 
innovations,  broadly  costate,  25-30  cells  thick  in  median  parts, 
mostly  crescentic  in  cross  section  with  broad,  thin,  often  erect  or 
inflexed,  commonly  purplish  margins,  the  air-chambers  almost 
wholly  filled  by  secondary  walls,  leaving  very  small  lacunae  ;  ventral 
scales  purple,  closely  imbricate,  the  inferior  reniform  or  obliquely 
ovate,  subentire  or  irregularly  dentate,  extending  to  the  margin,  the 
superior  ovate  or  lanceolate,  rather  abruptly  narrowed  to  a  slender, 
acuminate,  often  bifid,  subnodose  or  repand  point,  slightly  exceed- 
ing the  margin  and  inconspicuously  inflexed  at  the  apical  sinus,  their 
tips  rarely  decolorate :  polyoicous  :  antheridia  immersed,  with 
short-papilliform  ostiola,  somewhat  scattered  near  the  middle  of 
a  thallus-segment  or  aggregated  near  the  apex,  without  special 
involucre,  the  androecium  when  subapical  often  more  or  less  cov- 
ered by  the  inflexed  thallus-margins  :  peduncle  from  the  apex  of  a 
small  obovate  or  cordate  latero-ventral  innovation,  1.5-2.5  cm. 
high,  mainly  reddish,  pale  above,  with  a  few  small  purple  scales  at 
base,  naked  beneath  the  receptacle  or  obsoletely  palaceous  ;  9 
receptacle  small,  1.5-2.25  mm.  in  maximum  width,  hemispherical- 
conoidal,  lightly  papillate  above,  obscurely  lobed,  maturing  1-4 
sporogonia  :  capsule  purplish  ;  spores  dark  violet-purple^,  55-75  u., 
obscurely  angular,  tuberculate-verrucose  especially  on  the  outer 
face,  the  often  loose  and  saccate  exterior  membrane  of  the  inner 
faces  commonly  rugose  or  irregularly  areolate,  entire  surface  mi- 
nutely papillate  or  granulate  ;  elaters  2-4-  (mostly  3-)  spiral,  purple 
both  as  to  the  bands  and  the  unthickened  wall,  often  considerably 
attenuate  at  one  or  both  extremities,  200-300 long,  9-15 //in 
greatest  width. 

Yosemite  Valley,  "  on  rocks  in  the  spray  of  Bridal  Veil  Fall, 
June,  1866"  (Bolander)  ;  also  in  the  region  of  the  Yosemite  Val- 
ley by  C.  M.  Cooke,  Jr.,  1896  (in  herb.  A.  W.  Evans)  ;  on  trail  to 
Wilson's  Peak,  San  Gabriel  Mts.  (A.J.  McClatchie,  Apr.  21, 
1893)- 

A  portion  of  Bolander's  original,  communicated  by  Dr. 
Gottsche  to  Professor  Underwood,  is  in  the  Underwood  Her- 
barium. Other  specimens  from  the  first  collection  are  in  the 
herbaria  of  Professor  Underwood,  of  Columbia  University,  and  of 
the  U.  S.  National  Museum. 

Grinialdia  Californica  is  somewhat  nearly  allied  to  Grinialdia 
aiidrogyiia  (L.)  Lindb.  of  Southern  Europe,  but  clearly  differs  in  the 
purple  elaters,  the  dark  purple  spores,  the  broader  thallus  with 
thinner  margins,  the  rather  smaller,  more  conoidal,  less  evidently 


Cryptomitrium 


43 


lobed  9  receptacles,  in  the  origin  of  the  peduncle  from  a  small 
latero -ventral  innovation,  which  seems  to  occur  rarely  in  G. 
aiidrogyna,  and  in  other  characters  of  less  moment. 

Plate  92.    Grimaldia  Califormca. 

1.  Plant,  natural  size. 

2.  Thallus,  with  androecia  and  vegetative  latero-ventral  innovations,  X  3- 

3.  Thallus,  with  androecia  and  peduncle-bearing  innovation,  X  3- 

4.  Thallus,  with  peduncle-bearing  innovations,  X  3- 

5.  $  thallus,  X  3- 

6.  Transverse  section  of  thallus,  X  12. 

7.  Transverse  section  of  thallus  through  androecium,  X  12. 

8.  Transverse  sections  of  mature  peduncles,  X  29. 

9.  -1 3.  Outlines  of  ventral  scales,  mostly  from  near  thallus  apex,  X  29. 
14.  Stoma  and  adjacent  epidermal  cells,  X  305- 

15-17.  9  receptacles  (containing  dehisced  capsules),  X  4- 

18.  Ventral  view  of  receptacle  with  dehisced  capsules,  X 

19.  Spores,  X  305- 

20.  Elater,  X  305- 

All  figures  drawn  from  soaked  out  dried  specimens  ;  I-4,  6,  8,  9,  II,  and  14-20, 
from  portions  of  the  original  material  collected  by  Dr.  Bolander  and  preserved  in  herb. 
Underwood  (partly  communicated  by  Dr.  Gottsche  and  partly  from  U.  S.  National  Her- 
barium); 5,  7,  10,  12,  and  13,  from  specimen  collected  by  Professor  A.  J.  McClatchie 
in  the  San  Gabriel  Mts. 

6.  CRYPTOMITRIUM  Aust.  ;  Underw.  Bull.  111.  State  Lab.  Nat. 
Hist.  2:  36.  1884. 

Platycoaspis  Lindb.  Kongl.  Sv.  Vet.  Akad.  Handl.  23^:  ii. 
1889. 

Thallus  thin,  green  and  often  somewhat  polished  above,  con- 
colorous  or  purple  beneath,  6-15  mm.  long,  1-3  times  irregularly 
dichotomous,  with  the  inner  margins  of  the  terminal  segments  now 
and  then  connate,  sometimes  progressing  by  apical  or  lateral  in- 
novations from  the  rather  thick  costa,  the  segments  obovate,  ob- 
cordate,  or  oblong,  3-9  mm.  in  maximum  width,  venulose-areo- 
late,  stomata  simple,  the  minute  pores  surrounded  by  several 
finally  oblong  cells  with  unthickened  radial  walls  ;  scales  small, 
violet-purple,  irregularly  biseriate,  broadly  ovate,  variously  lobed, 
extending  half  way  to  the  margin  or  fragmentary  and  inconspicu- 
ous ;  air-chamber  layer  of  large  empty  cavities  separated  by  mostly 
unistratose  lamellae,  these  chambers  in  3-5  layers  over  the  costa 
in  transverse  section,  occupying  the  whole  of  the  veiy  thin  alar 
expansion  and  reduced  to  a  single  layer  at  the  margin,  the  latter 
green  or  brownish-purple,  undulate-repand  or  crenate,  sHghtly  as- 


44 


]\  I A  RC  H  AXTI A  C  E  AE 


cending  when  dry,  furnished  with  a  unistratose  border  1-3  cells 
in  width  ;  costa  (exclusive  of  air-chamber  layer)  1 5-20  cells  in 
thickness,  gradually  attenuate  to  the  wing  on  either  side.  Monoi- 
cous.  Antheridia  immersed  in  the  thallus  just  posterior  to 
the  base  o  the  $  branch,  in  a  single  or  irregularly  double, 
often  much  elongated,  row,  each  completely  filling  the  loculus  at 
maturity  and  adnate  to  its  walls,  ostioles  papilliform,  inconspicu- 
ous. 0  branch  from  the  apex  of  the  costa.  Peduncle  slender, 
naked  throughout,  pale  or  sometimes  brownish-purple  below, 
irregularly  sulcate,  the  dorsal  (posterior)  face  without  assimila- 
tiv^e  tissue,  the  ventral  (anterior)  face  with  a  single  root-hair  fur- 
row.* O  re::eptacle  subhemispherical  in  early  stages,  soon  be- 
coming circular-disciform,  lightly  convex  and  somewhat  papulose 
above,  plane  and  naked  beneath,  much  attenuated  toward  the  cre- 
nate  margin,  furnished  with  3-7  (commonly  5)  radiating  root-hair 
canals  in  communication  with  the  root-hair  furrow  of  the  peduncle, 
these  reaching  a  little  more  than  halfway  to  the  margin,  surrounded 
by  a  compact  tissue,  and,  after  drying,  sometimes  appearing  costa- 
like ;  air-chambers  large,  in  a  single  layer,  with  dolioform  stomata  ; 
archegonia  in  groups  of  four,  alternating  with  the  root-hair  canals, 
one  archegonium  of  each  group  (very  rarely  two)  producing  a 
sporogonium.  Sporogonia  (3—7,  usually  5)  occupying  ellipsoidal 
radiately  disposed  locuH  reaching  commonly  a  little  more  than  half 
the  distance  to  the  periphery  of  the  receptacle,  the  margins  of  the 
loculus  early  conniv^ent  and  enclosing  the  capsule,  separating  later 
by  an  elongated  radial  cleft,  this  finally  gaping  and  widely  expos- 
ing the  capsule,  the  lips  becoming  mambranous-scarious,  remain- 
ing somewhat  elevated,  and  functioning  as  the  only  involucre. 
Calyptra  inconspicuous.  Capsule  nearly  spherical  or  oblately 
ellipsoidal,  the  apical  third  or  fourth  of  its  wall  consisting  of  two 
layers  of  cells  and  falling  away  as  an  operculum  in  dehiscence,  the 
wall  otherwise  unistratose,  the  cells  without  annular  or  spiral  thick- 
enings, those  of  the  operculum  with  the  walls  slightly  thickened 
at  the  angles.  Seta  very  short,  foot  bulbous.  Spores  brown, 
more  or  less  distinctly  tetrahedral,  irregularly  areolate-lamellate, 
with  a  pellucid  margin.  Elaters  attenuate,  contorted,  often 
branched,  closely  2 -(3-)  spiral. 
The  only  recognized  species  is 


*  Herr  Stephani  (Bot.  Gaz.  17  :  58.  1892)  has  described  the  peduncle  of  Cali- 
fornian  specimens  of  Ciyptomitrium  as  bicanaliculate.  In  his  latest  paper  on  the  sub- 
ject  (Bull.  Herb.  Boiss.  7  :  222.  1899)  he  reports  that  in  Chilian  plants  he  finds  the 
peduncle  with  one  or  with  two  root-hair  furrows  in  one  and  the  same  specimen.  In 
our  investigations  upon  Californian  material  and  the  Hookerian  type,  we  have  as  yet 
been  unable  to  find  any  peduncle  with  more  than  one  furrow  containing  rooi-hairs. 


Cryptomitrium 


45 


I.  Cryptomitrium  texerum  (Hook.)  Aust.  1.  c.    Stephani,  Bot. 

Gaz.   17:58.      1892.      Bull.   Herb.   Boiss.   7:  222.  1899. 

Howe,  Erythea,  5  :  87. //.  j.  1897. 

Marchantia  tencra  Hook.;  Kunth,  Syn.  PI.  i  :  46.  1822. 

Duvalia  tenera  Gottsche  ;  G.  L.  &  X.  Syn.  Hep.  554.  1846. 

Peduncle  I  5-30 mm.  X  .3  — .5  mm.:  spores  3 5-50//;  elaters  30c- 
450 long,  7-1 1  u.  in  greatest  width. 

On  moist  shaded  banks  Fruit  Vale,  Alameda  Co.  (Miss  Edith 
S.  Byxbee,  Howe);  Navarro,  Mendocino  Co.  {fide  Miss  Byxbee); 
Menlo  Park  (Blasdale);  Folsom  (Mrs.  Brandegee);  Jackson,  Ama- 
dor Co.  (Hansen);  San  Bernardino  (Parish,  no.  2221);  Pasadena 
(McClatchie).  Also  collected  in  CaHfornia  by  Parry,  Bigelov/, 
Bolander,  and  Torrey  {^fidc  Austin). 

The  original  plant  of  Hooker  was  collected  near  Ario  [State 
of  Michoacan  (?)  ]  ^Mexico,  by  Humboldt.  A  portion  of  this,  bear- 
ing the  legend  "  no.  236,  Marchantia  tencra,  n.  sp.,"  Legit  Hum- 
boldt," has  been  sent  to  Professor  Underwood  through  the  kind- 
ness of  Dr.  W.  T.  Thiselton-Dyer,  the  Director  of  the  Royal 
Botanic  Gardens  at  Kew,  and  is  deposited  in  the  herbarium  of 
Columbia  Univ^ersity.  This  fragment,  rather  unfortunately,  bears 
only  very  young  capsules,  so  we  have  been  unable  to  compare 
spores  and  elaters,  but  it  seems  to  agree  essentially  with  the  Cali- 
ifornian  plant  in  structure  of  thallus  and  peduncle,  form  of  re- 
ceptacle, position  of  antheridia,  etc.  The  peduncles  are,  however, 
much  shorter,  being  only  3  or  4  mm.  high,  but  this  may  be  due, 
in  part,  to  the  immaturity  of  the  receptacle  ;  they  are,  moreover, 
only  about  half  as  thick  as  in  the  Californian  specimens,  which 
may,  perhaps,  be  partly  accounted  for  by  the  possibility  that  after 
so  many  years'  drying  they  may  not  fully  regain  their  natural  size 
on  soaking  out. 

Duvalia  brevipeduncidata  Mont,  from  Chili  {legit  Gay)  is  thought 
by  authors  to  be  the  same  species.  The  Californian  plant  is,  how- 
ever, in  need  of  further  comparison  with  those  of  ]\Iexico  and  Chili. 

Plates  93  and  94. 

Cryptomitrium  tenerum. 

I-4.  Principal  segments  of  thallus,  natural  size  ;  fig.  2,  with  a  fully  developed  9 
receptacle  and  peduncle. 


46 


Marchantiaceae 


5.  Ventral  view  of  a  thallus -segment,  X  3- 

6.  Thallus-lobe  showing  young  9  receptacle  and  position  of  antheridia,  X  ^• 

7.  Margin  of  thallus-lobe,  dorsal  view,  showing  position  of  the  internal  lamellae 
which  mark  the  boundaries  of  the  air-chambers,  X  23  (from  alcoholic  material). 

8.  Outline  of  median  longitudinal  section  of  thallus  with  antheridia  and  young  9 
receptacle,  X  23. 

9.  Portion  of  a  transverse  section  of  a  thallus-lobe,  passing  through  androecium 

X  23. 

10.  Median  region  of  a  transverse  section  (about  lo /a  in  thickness),  passing  through 
androecium,  X  4i- 

11.  Marginal  region  of  the  same  transverse  section,  X  41- 

12.  Anterior  margin  of  thallus-lobe  from  a  slightly  oblique  longitudinal  section,  X 

225. 

13  and  14.  Outlines  of  cross-sections  of  mature  peduncles,  X  53  (dried  material). 

15.  Outlines  of  cross-sections  of  a  peduncle  from  the  original  material  of  "  Mar- 
chanlia  tenera'''  collected  in  Mexico  by  Humboldt,  X  53- 

16.  Section  of  young  peduncle,  X  S^- 

17.  Stoma,  surface  view,  X  225. 

18.  Transverse  section  of  thallus-apex  with  young  9  receptacle,  X  23. 

19.  Vertical  section  through  9  receptacle  with  mature  sporogonium  (slightly  sche- 
ma'ized),  X  23. 

20.  Vertical  section  of  a  young  9  receptacle  about  half  way  between  the  vertical 
axis  and  periphery,  traversing  one  of  the     costae"  v/ith  its  root-hair  canal,  X  23. 

21.  A  thin  horizontal  section  of  a  young  9  receptacle,  showing  the  archegonia  in 
cross-section  in  groups  alternating  with  the  root-hair  canals,  X  3^' 

22.  A  young  antheridium,  free  in  its  loculus,  X  225. 

23.  A  mature  antheridium,  with  wall  more  or  less  blended  with  that  of  the  loculus, 
X53- 

24-27.  Receptacles  with  mature  sporogonia,  X  4  5  24  and  25,  lateral  views,  the 
former  from  alcoholic,  the  latter  from  dry  material  ;  26  and  27,  dorsal  views,  the  latter 
from  a  pressed  specimen  and  showing  externally  the  position  of  the  *'  costae";  28,  ven- 
tral view. 

29.  Capsule,  apical  view,  showing  operculum,  X  23. 

30  and  31.  Vertical  sections  of  capsules-wall,  showing  the  bistratose  operculum, 
X27. 

32.  Spore,  outer  face,  X  3^5 • 

33.  Elater,  X  305- 

Figs.  1-3,  6-12,  16-24,  26,  28-31,  from  material  collected  at  Fruit  Vale,  Ala- 
meda County,  fixed  at  l%  chromic  acid,  and  preserved  in  alcohol  ;  4  from  Folsom 
{Mrs.  Brandegee)  ;  5,  dry  material.  Fruit  Vale;  13,  14,  32  and  33,  dry  material, 
Menlo  Park  (Mr.  W.  C.  Blasdale)  ;  15,  Mexico  (Humboldt)  ;  27,  Pasadena  (Mc- 
Clatchie). 

7.  ASTERELLA  Pal.  de  Beauv.  /.  /.  Encyc.  Meth.  Bot.  SuppL 

I  :  502.  1810. 

Fimbraria  Nees,  Hort.  Phys.  Berol.  44.  1820.  [Afterward 
amended  to  Fimbriaria.'^ 


ASTERELLA 


47 


Hypenantron  Corda  ;  Opiz,  Beitr.  i:  648.  1829.*  Schiffn.; 
Engler  &  Prantl,  Nat.  Pflanzenfam.      :  33.  1893. 

Thallus  simple  or  somewhat  sparingly  dichotomous,  some- 
times innovating  in  front,  or  laterally  from  near  the  ventral  aspect 
of  the  more  or  less  distinctly  defined  costa,  ver>^  thick,  usually 
with  a  thin  often  colored  margin,  commonly  dark  purple  beneath, 
pores  and  areolae  of  the  dorsal  surface  distinct  or  somewhat  ob- 
scure in  drying,  air  chambers  deep  or  rather  small  and  shallow, 
scales  mostly  violet-purple,  rarely  white.  Stomata  of  the  thallus 
simple,  the  cells  bounding  the  pore  with  unthickened  radial  walls  ; 
stomata  of  9  receptacle  dolioform.  Monoicous  or  less  commonly 
dioicous.  Antheridia  in  slightly  elevated  or  indistinctly  defined 
receptacles  borne  in  the  median  line  of  the  thallus  or  on  short 
specialized  lateral  branches.  9  receptacle  obtusely  conical  or  sub- 
hemispherical,  sometimes  depressed-umbonate  or  flattened  with 
age,  maturing  1-6  (usually  3  or  4)  sporogonia,  these  enclosed 
singly  in  the  strongly  or  obscurely  defined  lobes.  Peduncle  a 
direct  continuation  of  the  thallus  or  of  a  thallus-lobe,  the  latter 
sometimes  a  small  innovation  springing  from  one  side  of  the 
median  ventral  line  of  the  costa,  the  dorsal  (posterior)  face  of  the 
peduncle  without  assimilative  tissue,  the  ventral  (anterior)  face  with 
a  single  root-hair  furrow,  in  communication  with  which  at  the  apex 
are  root-hair  sinuses  in  the  receptacle  alternating  with  its  lobes. 
Sporogonium  and  the  commonly  inconspicuous  calyptra  surrounded 
by  a  special  membranous  involucre,  the  pseudoperianth,  this  at 
first  somewhat  conical,  becoming  usually  much  exserted,  finally 
with  3-18  longitudinal  clefts,  the  narrow  segments  coherent  at  the 
apex  or  soon  becoming  free.  Capsule  globose,  ovoid,  or  obovoid, 
more  or  less  regularly  circumscissile  above  the  middle  or  (in  A. 
Califovnicd)  the  apical  third  or  half  of  the  wall  falling  away  irregu- 
larly in  dehiscence,  leaving  a  fimbriate-lacerate  margin,  the  wall — 
for  the  most  part,  at  least — of  a  single  layer  of  cells,  without  an- 
nular or  spiral  thickenings,  but  often  exhibiting  distinct  trigones, 
especially  in  the  apical  portion.  Seta  very  short,  foot  bulbous. 
Spores  rather  large,  yellow,  brown,  fuscous,  or  black,  obscurely 
tetrahedral  or  flattened,  usually  with  a  pellucid  margin,  areolate- 
reticulate,  verrucose,  or  irregularly  rugose.    Elaters  1-4-spiral. 

Key  to  the  Species. 

9  branch  from  apex  of  thallus  or  of  one  of  its  main  segments. 

9  receptacle  strongly  lobed,  paleaceous-barbate  beneath  ;  segments  of  pseudo- 
perianth  coherent  at  apex  ;  spores  yellow,  elaters  mostly  unispiral  ;  dioicous. 

I.  A.  Californica. 

9  receptacle  conic-ovate,  scarcely  lobed,  naked  beneath  ;  segments  of  pseudo- 
perianth  coherent  at  apex  ;  spores  fuscous  or  black,  elaters  1-3-spiral  ;  monoi- 
cous. 4.  A.  Palmeri. 

*  See  foot-note  under  Ricciccm-piis,  p.  33. 


48 


Marchantiaceae 


9  receptacle  subhemi spherical,  obscurely  lobed,  naked  beneath ;  segments  of 

pseudoperianth  soon  free  ;  spores  yellow  or  light  brown,  elaters  2-4-spiral  ; 

monoicous.  5.  A  gracilis. 

9  branch  from  apex  of  a  small  lateral  innovation,  the  latter  springing  from  the  costa  at 

one  side  of  its  median  ventral  line. 
9  receptacle  subhemispherical  or  depressed-umbonate,  2-3.5  maximum 

width;  pseudoperianth  white,  lo-l2-cleft;  spores  65-85//,  elaters  8-lo//  broad. 

2.  A.  Bo  andtri 

9  receptacle  subconoidal,  2.5-5  '^n^-       maximum  width  ;  pseudoperianth  com- 
monly violet,  l2-i8-cleft ;  spores  60-125      elaters  9-16  //  broad. 

3.  A  violacei. 

I.  AsTERELLA  Californica  (Hampe)  Underw.  Bot.  Gaz.  20  :  60. 

.  1895. 

Fimbriaria  Californica  Hampe;  Aust.  Hep.  Bor.-Am.  135. 
1873.    Underw.  Bull.  111.  State  Lab.  Nat.  Hist.  2  :  41.  1884. 

Satiteria  Ihnbata  *  Aust.  p.p.  Proc.  Acad.  Nat.  Sci.  Philad. 
1869  :  229.  1869. 

"^Through  the  kindness  of  Professor  F.  E.  Weiss  of  the  Owens  College,  Manches- 
ter, England,  we  have  enjoyed  the  privilege  of  being  able  to  examine  the  original 
material  upon  which  the  Saiitcria  limbata  of  Austin  was  founded.  We  find,  as  does 
Count  Solms-Laubach,  who  has  also  studied  the  type  specimen  (See  Bull.  Herb  Boiss. 
6  :  773.  1898),  that  Mr.  Austin's  diagnosis  was  drawn  from  two  entirely  distinct 
species,  but  we  would  differ  from  Count  Solms  as  to  the  identity  of  the  plant  from  which 
the  first  and  greater  part  of  Mr.  Austin's  description  was  made  and  from  which  the 
character  embodied  in  the  specific  name  was  derived.  This  we  find  to  be  the  $  thal- 
lus  of  the  always  dioicous  Asterella  Californica.  We  were  assisted  to  this  view  by  the 
discover}'  of  an  androecium,  which  apparently  escaped  the  notice  both  of  Mr.  Austin 
and  Count  Solms.  The  antheridia  were  found  to  be  united  in  a  definitely  circumscribed 
and  slightly  elevated  area  and  to  have  short  ostiolar  papillae  instead  of  being  scattered 
and  having  slenderly  subcylindrical  ostiolar  papillae  as  in  Sauteria  and  Clei<ea,  The 
ventral  scales,  stomata,  dorsal  epidermis,  and  internal  structure  were  then  compared 
with  the  corresponding  parts'of  Asterella  Californica  and  were  found  to  agree  so  accu- 
rately as  to  remove  all  doubts  of  the  specific  identity  of  the  two  plants.  And  we  could 
discover  no  evidence  of  admixture  save  with  the  fertile  plant  from  which  Mr.  Austin 
drew  the  description  of  the  9  receptacle  and  peduncle  of  his  Sanie7'ia  litnbata.  This 
fertile  plant  we  refer  to  our  Clevea  hyalina  Californica.  Count  Solms,  however  {fide 
Stephani,  /.  r. )  found  the  stomata  of  the  **  sterile"  plant  to  conform  to  those  of  Clevea 
hyalina.  We,  on  the  other  hand,  found  that  the  radial  walls  of  the  cells  bounding  the 
stomata  were  sometimes  slightly  thickened,  but  no  more  so  than  in  Asterella  Californica 
and  never  in  any  such  way  as  in  the  typical  stomata  of  the  Astroporae,  Asterella 
Californica  was  found  growing  at  Ukiah  in  rather  close  association  with  our  Clevea 
hyalina  Californica. 

Count  Solms,  in  his  MS.  notes  left  with  the  type-material,  retains  the  specific 
name  lirtibata  for  the  "sterile"  [^]  plant,  and  it  would  seem  to  us  that  the  name 
should  be  maintained  for  this  plant  if  for  either,  inasmuch  as  the  description  of  it  occu- 
pies the  first  and  by  far  the  greater  part  of  the  original  diagnosis  and  inasmuch  as  it  was 


ASTERELLA 


i9 


C/evsa /Mata  Solms  p.p.\  Stephani,  Bull.  Herb.  Boiss.  6:  773. 
1898. 

Thallus  8-25  mm.  long,  usually  1-3  times  dichotomous,  rarely 
simple,  the  ultimate  segments  emarginate,  obovate,  obcordate,  or 
broadly  oblong,  indistinctly  areolate,  4-12  mm.  in  maximum  width, 
abruptly  carinate,  lightly  concave  above,  sometimes  narrowly  cana- 
liculate toward  apex,  25-35  cells  thick  in  axile  parts,  with  rather 
large  air-cavities,  margins  undulate-crenate,  commonly  broad,  thin, 
and  brownish-purple,  ascending  on  drying,  or  occasionally  erect 
and  connivent,  the  reddish-purple  scales  subovate  or  ovate-lanceo- 
late, not  reaching  to  the  margin  or  sometimes  inconspicuously  pro- 
jecting and  inflexed  toward  the  apex,  each  with  1-4  slenderly 
acuminate  or  subfiliform,  nearly  entire,  often  decolorate,  apical 
processes  :  dioicous  :  $  and  9  plants  mostly  in  separate  patches  ; 
antheridia  in  clearly  circumscribed,  slightly  elevated,  strongly  pap- 
illate, oval  to  linear-oblong,  sometimes  forked  discs,  situated  in 
the  median  line  of  the  thallus  at  some  distance  back  of  the  apex  : 

9  receptacle  from  the  apex  of  the  thallus,  bearing  2  or  3  arche- 
gonia  in  each  of  the  4  or  5  groups,  maturing  1-5  (commonly  4) 
sporogonia,  large  (4-7  mm.  in  maximum  width),  finally  subhemi- 
spherical,  convex-umbonate,  nearly  smooth  above,  containing  large 
air-chambers  and  becoming  vesicular-rugose  in  drying,  paleaceous- 
barbate  beneath,  with  strongly  defined,  rather  turgid,  often  spread- 
ing, entire  or  crenate-margined  lobes  ;  peduncles  stramineous,  stout, 

10-30  mm.  X  .5-6  mm.,  naked  or  very  sparingly  pilose,  without 
paleae  at  base,  sulcate,  angled,  and  slightly  twisted  when  dry  ; 
pseudoperianth  white,  large,  ovate,  12-16  cleft,  the  segments  co- 

evidently  the  "broad,  wavy,  dark  purple,  membranaceous  margin  of  the  frond"  of 
this  that  suggested  the  name.  Others,  however,  might  contend  that  the  name  should  go 
with  the  more  complete  capsule-bearing  plant.  If  retained  for  the  former  (the  Asterella)^ 
it  would  appear  to  have  four  years  priority  over  the  name  Californica^  and  would  thus, 
in  accordance  with  the  Rochester  Code,  replace  this  latter  name  But  the  determina- 
tion of  sterile  and  even  of  antheridiiferous  thalli  of  the  Marchanticeae  is  sometimes  a 
difficult  and  delicate  matter,  and  it  is  to  be  feared  that  in  this  particular  case  the  iden- 
tity of  the  scanty  material  upon  which  Satiteria  linibata  "  was  chiefly  founded  migl  t 
always  be  more  or  less  of  a  bone  of  contention."  The  unsatisfactoriness  of  this 
material  may  be  seen  from  the  fact  that  while  we  identify  it  without  hesitation  with 
Asterella  Californica,  a  botanist  of  the  experience  and  high  standing  of  Count  Solms- 
Laubach  refers  it  without  reserve  to  the  genus  Clevea.  Under  these  circumstances  it 
seems  to  us  the  wiser  course  to  leave  aside  altogether  the  name  linibata 

vSince  the  above  note  was  written.  Count  Solms-Laubach,  to  whom  we  had  sent  a 
specimen  of  Asterella  Californica  for  comparison,  admits  (Bot.  Zeit  57'  :  30.  16 
1899)  that  we  have  shown  "  ziemlich  zweifellos  "  its  identity  with  the  sterile  portion 
of  the  original  ' '  Satit  'Ha  limbata  ' '  material 


50 


Marchantiaceae 


hering  at  the  rather  obtuse  apex,  their  margins  plane  or  lightly 
reflexed  ;  capsule  prolate-ellipsoidal  or  subglobose,  the  apical  third 
or  half  of  its  wall  falling  away  irregularly  in  dehiscence  leaving 
behind  a  cup  with  a  fimbriat'e-lacerate  margin  ;  seta  very  short ; 
spores  yellow,  90-135  y.,  obscurely  tetrahedral,  furnished  when 
mature,  especially  at  the  angles  and  on  the  outer  face,  with  con- 
spicuous irregular  ridges  or  wings,  the  entire  surface  minutely 
reticulate  (meshes  1-5  /y.);  elaters  150-240//  x  12-18 /^  obtuse, 
slightly  flexuous,  with  one  or,  more  rarely,  two,  yellow  spiral 
bands. 

Exsicc.  Hep.  Bor.-Am  135  (as  Fimbriaria  Calif oniica). 
Hep.  Am.  119  (as  Fimbriaria  Californica). 

On  open  or  lightly  shaded  banks,  often  about  rocks,  from  San 
Diego  as  far  north,  at  least,  as  Mendocino  and  Shasta  counties. 
Berkeley  ;  San  Leandro  (Underwood) ;  San  Francisco  (Bolander, 
Brandegee,  Howe)  ;  Mill  Valley  ;  Duncan's  Mills,  Sonoma  Co.; 
Ukiah  and  Half- Way  House,  Mendocino  Co.;  Oak  Run,  Shasta 
Co.  (Baker  &  Nutting);  Jackson,  Amador  Co.  (Geo.  Hansen);  Mur- 
phy's, Calaveras  Co.  (J.  Burtt  Davy);  Mt.  Diablo,  Contra  Costa 
Co.;  Knight's  Landing,  Stainslaus  Co.  (F.  \V.  Bancroft);  Fresno 
(F.  P.  Nutting);  Kaweah  River,  Tulare  Co.  (Coville);  Lake  San 
Andreas,  San  Mateo  Co.;  Palo  Alto  (Campbell);  Santa  Cruz  (Far- 
low);  Santa  Barbara  (Farlow);  Nordhoff,  Ventura  Co.  (Miss  Jac- 
queline K.  Newton);  Santa  Monica  (Dr.  Hasse);  Pasadena  (Un- 
derwood, McClatchie,  Miss  Teed);  Santa  Ana  Hills  (Parish,  no. 
1728);  Twin  Oaks,  San  Diego  Co.  (F.  W.  Koch),  San  Diego  (Far- 
low).    First  collected  by  Bolander  and  by  Bigelow. 

Somewhat  resembling  in  size  and  habit  Aster clla  Lvidciibcrgiana 
Lindb.  (Fimbriaria  Lindcnbcrgiana  Corda),  but  very  distinct  in 
being  dioicous,  in  the  broader,  obovate  thallus-segments,  in  the 
more  strongly  lobed  receptacle,  the  white  pseudoperianth,  the 
much  larger,  more  prominently  rugose-winged,  more  minutely  re- 
ticulate, yellow  spores,  the  yellow  spiral  band  of  the  elaters,  the 
irregular  dehiscence  of  the  capsule,  etc. 

Plates  95  and  96.    Astkrb:lla  Californica. 
I.   9  plant,  natural  size,  with  fully  developed  receptacle  and  peduncle. 
2  and  3.  Male  plants,  natural  size. 

4.  A  large  sterile  thallus,  natural  size. 

5.  9  plant  with  young  receptacles,  natural  size. 


ASTERELLA 


51 


6.  Ventral  view  of  thallus,  natural  size. 

7.  Outline  of  transverse  section  of  $  thallus,  near  apex,  X  23. 

8.  Outline  of  transverse  section  through  androecium,  X  23. 

9.  Outline  of  longitudinal  section  through  androecium,  X  ^2. 
1,0.  A  very  thin  median  longitudinal  section  of  thallus,  X  48- 
II-13.  Ventral  scales,  X  12. 

14.  Cross-section  of  young  peduncle,  X  4.1. 

15.  Cross-section  of  a  mature  peduncle,  X  23. 

16.  Surface  view  of  stoma  from  near  thallus-apex,  X  3^5 • 

17.  Outline  of  portion  of  longitudinal  section  of  thallus  passing  through  a  young 
receptacle,  X  23. 

18.  Section  of  9  receptacle  in  a  plane  perpendicular  to  the  peduncle,  showing 
two  archegonia  in  transverse  section  in  each  lobe,  one  of  each  group  fertilized  and  sur- 
rounded by  the  developing  pseudoperianth,  X  23. 

19-21.   9  receptacles,  the  first  young,  the  others  with  matured  sporogonia,  X  4- 

22.  Sectional  view  of  a  dolioform  stoma  from  9  receptacle,  X  225. 

23.  Outline  of  mature  sporogonium,  X  ^2. 

24.  Capsule  after  dehiscence,  X  ^2. 

25.  Spore,  outer  face,  X  225.  (The  folds  and  ridges  of  the  epispore  become 
sometimes  more  numerous  than  here  represented. ) 

26.  Spore,  inner  faces,  X  225. 

27.  A  portion  of  the  wing-margin  of  spore,  showing  the  minute  reticulation,  X 

305- 

28.  Sections  of  fully  matured  spores,  showing  the  great  development  of  the  epi- 
sporic  layer,  X  225.  The  section  on  the  left  passes  considerably  to  one  side  of  the  cen- 
ter of  the  spore  and  consequently  gives  a  somewhat  exaggerated  idea  of  the  thickness  of 
epispore  in  comparison  with  the  width  of  the  lumen. 

29.  Elaters,  X  225. 

Figs.  II-13  drawn  from  material  collected  by  Mr.  J.  Burtt  Davy  in  Calaveras 
County  ;  25-27,  and  29,  from  Oak  Run,  Shasta  County  (M.  S.  Baker  and  F.  P.  Nut- 
ting); the  remaining  from  specimens  collected  by  the  author  in  Mill  Valley  and  near 
Berkeley. 

2.  AsTERELLA  BoLANDERi  (Aust.)  Underw.  Bot.  Gaz.  20:  6i. 

1895. 

Fimbriaria  Bolanderi  Aust.  Proc.  Acad.  Nat.  Sci.  Philad. 
1869 :  230.  1869. 

Aromatic  :  thallus  oblong  to  narrowly  linear,  10-45  n^"^-  X 
1.5-4  mm.,  costate-carinate,  25-35  cells  thick  in  the  middle,  be- 
coming very  thin  at  the  usually  desiccated  border,  innovating  in 
front  or  laterally  from  the  costa,  very  rarely  dichotomous,  indis- 
tinctly porose,  scarcely  areolate,  light  green  above,  usually  purple 
below,  rounded  or  very  slightly  emarginate  at  the  apex,  nearly 
flat  or  lightly  concave  when  living,  the  membranous,  undulate- 
crenate,  brown  or  purple  margins  often  erect  and  appressed- 
connivent  on  drying,  air-cavities  small,  scales  dark  purple,  not 
reaching  the  margin,  the  costa  densely  clothed  with  matted  root- 


52 


Marchantiaceae 


hairs  :  monoicous  :  androecia  and  9  branches  on  short  lateral  in- 
novations, the  latter  1.5-3  length,  springing  from  the  sides 
of  the  costa,  those  bearing  a  9  branch  expanded,  emarginate, 
bilobed,  or  somewhat  obcordate,  1-3  mm.  in  maximum  width,, 
those  bearing  the  androecia  subclavate,  only  slightly  expanded  at 
the  distal  extremity,  .75-1  mm.  in  greatest  width,  all  furnished 
beneath  with  scales  and  root-hairs :  $  receptacles  with  4—10 
antheridia,  lightly  papillate :  9  receptacle  with  a  single  archegonium 
in  each  of  the  4  lobes,  maturing,  commonly,  3  or  4  sporogonia, 
subhemispherical  at  first,  becoming  subconical,  depressed-umbo- 
nate,  or  muc  hflattened,  with  lobes  somewhat^divergent,  2—3.5  irim. 
in  maximum  width,  containing  small  air-chambers,  rugose  on 
drying,  surrounded  and  more  or  less  covered,  when  young  and 
sessile,  with  narrow  paleae,  these  springing  mostly  from  its  ventral 
surface  or  from  the  top  of  the  peduncle,  inconspicuous  at 
maturity  ;  peduncle  slender,  10-25  mm.  x  .2 5-. 5 5  mm., 
brownish-purple  below,  pale  above,  sparingly  pilose  toward  the 
base,  apex  of  the  innovation  often  with  a  few  inflexed  scales  ; 
pseudoperianth  white,  conical  and  acute  when  young,  finally  sub- 
globose,  with  10-12  widely  gaping  clefts,  the  segments  cohering 
at  the  apex,  rarely  free  with  age,  mostly  with  revolute  margins  : 
capsule  subglobose,  sometimes  oblately  flattened,  nearly  black  at 
first,  becoming  brown,  circumscissile  somewhat  above  the  middle^ 
the  margin  irregularly  toothed  ;  spores  obscurely  tetrahedral, 
65-85  fi,  yellowish-brown,  becoming  darker,  alveolate-areolate, 
with  4-6  areolae  across  each  face,  the  angles  of  the  spore  with  a 
yellow  pellucid  margin  ;  elaters  200-250  /j.  x  8-10  /i,  closely 
2-(3-)  spiral. 

Exsicc.  Hep.  Bor.-Am.  I36d  (as  Fimbriaria  Bolanderi). 
Hep.  Am.  158  (as  Fimbriaria  Bolanderi). 

On  lightly  shaded  banks.  Fruit  Vale,  Alameda  Co.  (Miss 
Edith  S.  Byxbee)  ;  Mill  Valley,  Marin  Co.  (Howe)  ;  Hood's  Peak, 
Sonoma  Co.  (F.  T.  Bioletti)  ;  Howell  Mountain,  Napa  Co.  (W.  A. 
Setchell)  ;  Jackson,  Amador  Co.  (Geo.  Hansen)  ;  Twin  Oaks,  San 
Diego  Co.  (F.  W.  Koch).  The  species  was  originally  collected 
at  San  Rafael  by  Dr.  Bolander  in  1865  ;  it  was  also  found  by 
Dr.  Bolander  in  the  same  year  in  dry  ditches  near  the  American 
River  at  Auburn. 

The  lateral  innovations  in  this  species  show  a  tendency  to 
group  in  such  a  way  that  nearly  all  on  one  side  of  the  thallus  bear 
antheridial  discs  and  nearly  all  on  the  other  side  9  branches. 
In  the  San  Diego  County  specimen  the  elaters  are  sometimes 
unispiral. 


ASTERELLA 


53 


Plate  97.    Asterella  Bolanderi. 

1.  Plant,  natural  size. 

2.  A  sterile  thallus,  natural  size,  somewhat  unusually  elongated. 

3.  Thallus,  branching  by  latero-ventral  innovations,  X  3- 

4.  Thallus  with  the  small  ^  and  9  latero  ventral  innovations  and  an  apical 
vegetative  innovation,  X  3- 

5.  Receptacle  with  mature  sporogonia,  X4  (fiom  alcoholic  material). 

6.  Young  receptacle  X  4  (alcoholic). 

7.  Mature  receptacle  (dry),  X  4- 

8.  Mature  receptacle  (dry),  X  ^• 

9.  Longitudinal  section  of  thallus  a  little  one  side  of  the  median  line,  showing  in- 
sertion of  the  $  and  9  innovations,  X23. 

10.  Median  longitudinal  section  of  the  thallus,  X  47- 

11.  Transverse  section  of  the  thallus,  with  androecium,  X  23. 

12.  Innovation  with  young  9  receptacle,  X  ^• 

13.  Innovation  bearing  androceium,  X  8. 

14.  Stoma,  surface  view,  X  225. 

15.  Longitudinal  section  of  innovation  with  young  9  receptacle,  X  23. 

16.  Portion  of  transverse  section  of  thallus  with  longitudinal  section  of  a  (J 
branch,  X  4i- 

17.  Transverse  section  of  one  of  the  innovations  near  the  base,  X4i- 

18.  Cross-section  of  peduncle,  X  4i- 

19.  Sporogonium,  X  ^2. 

20.  Capsule  after  dehiscence,  y^i2. 

21.  Spores,  X  225. 

22.  Elater,  X  225. 

The  figures  are  drawn,  for  the  most  par',  from  specimens  collected  in  Mill  Valley, 
Marin  County. 

3.  Asterella  vioLACEA  (Aust.)  Underw.  Bot.  Gaz.  20  :  61.  1895. 
Firnbriaria  violacea  Aust.  Bull.  Torr.  Bot.  Club,  3  :  17.  1872. 

Aromatic  :  thallus  oblong  to  narrowly  linear,  8-25  mm.  x 
1.5-5  nim.,  broadly  costate-carinate,  30-40  cells  thick  in  median 
parts,  becoming  very  thin  at  the  border,  innovating  in  front,  or 
more  often,  laterally,  from  the  costa,  sometimes  dichotomous,  ob- 
scurely areolate,  indistinctly  porose,  rounded  or  sHghtly  emarginate 
at  the  apex,  densely  radiculose  along  the  costa,  concave  or  canal- 
iculate above  when  moist,  the  usually  broad  brownish  mem- 
branous undulate-crenate  margins  commonly  erect  and  appressed- 
connivent  on  drying,  air-chambers  small,  scales  dark  purple, 
sometimes  reaching  the  margin  :  monoicous  :  androecia  and  9 
branches  borne  on  short  innovations,  the  latter  springing  from  the 
sides  of  the  costa  near  the  median  ventral  line,  1 . 5-4  mm.  long, 
those  bearing  a  9  branch  bilobed,  obcordate,  or  emarginate -sub- 
quadrate,  2-4  mm.  in  maximum  width,  those  bearing  an  androe- 
cium clavate,  slightly  expanded  at  apex,  .75-1  mm.  in  maximum 


54 


Marchantiaceae 


width  :  androecia  with  6-12  antheridia,  Hghtly  papillate  :  9  recep- 
tacle enclosing  a  single  archegonium  in  each  of  the  four  lobes, 
maturing  2-4  (usually  3)  sporogonia,  subconoidal  with  lobes  di- 
rected downward  or  only  slightly  spreading,  very  rarely  becoming 
depressed-umbonate,  2.5-5  in  maximum  width,  containing 

small  air-chambers,  nearly  smooth,  or,  on  drying,  subrugose, 
densely  surrounded  when  young  and  sessile  by  narrow  paleae 
springing  from  its  ventral  surface  or  from  the  upper  part  of  the 
peduncle,  these  usually  persistent  and  conspicuous  at  maturity  ; 
peduncle  10-40  mm.  x  .35-  55  mm.,  brownish-purple,  becoming 
abruptly  pale  near  the  receptacle,  pilose  toward  the  base  ;  pseudo- 
perianth  commonly  violet,  conical,  finally  obovoid-subpyriform, 
or  subglobose,  with  12-18  narrow  or  somewhat  gaping  clefts,  the 
segments  cohering  at  the  apex  or  often  becoming  free  with  age, 
sometimes  flexuous  and  contorted,  the  margins  commonly  revolute  : 
capsule  nearly  spherical,  the  apical  third  or  half  of  its  wall  falling 
away  in  dehiscence  in  irregular  fragments,  leaving  a  subhemi- 
spherical  cup  with  an  erose-dentate  margin  ;  spores  yellow,  be- 
coming brownish,  60-125//,  alveolate-areolate,  with  4-7  areolae 
across  each  face,  the  pellucid  margins  finally  broad  ;  elaters  180— 
^20  fjt  X  9-16  fJt,  closely  2-  or  3-spiral,  the  bands  yellow  or  slightly 
brown. 

On  exposed  or  somewhat  shaded  banks.  Olema,  Marin  Co. 
(Jepson) ;  St.  Helena,  Napa  Co.  (Jepson)  ;  Ukiah,  Mendocino  Co. 
(Howe) ;  Oak  Run,  Shasta  Co.  (Baker  and  Nutting) ;  Jackson, 
Amador  Co.  (Hansen)  ;  Pasadena  (McClatchie,  Feb.  18,  1893 — 
mixed  with  A.  Palmeri).  Collected  also  by  Coulter  and  by  Bo- 
lander,  the  precise  locality  unknown.  Not  known  to  occur  out- 
side of  California. 

Asterclla  violacea  seems  occasionally  to  approach  A.  Bolandcri^ 
but  is  usually  very  distinct  in  the  violet  1 2-1 8-cleft  pseudoperianth, 
the  larger  more  conical  9  receptacle  with  less  spreading  lobes 
and  more  abundantly  paleaceous-barbate  beneath,  and  in  the 
commonly  larger  spores  and  broader  elaters.  The  violet  colora- 
tion sometimes  disappears  from  the  pseudoperianth,  but  in  such 
cases  a  trace  of  purple  is  often  to  be  detected  on  the  outer  in- 
volucre near  its  margin. 

Plate  98.    Astkrella  violacea. 

1.  Plant,  natural  size. 

2.  Thallus,  showing  branching  and  the  $  and  9  innovations,  X  3- 

3.  Thallus,  natural  size,  with  the  somewhat  unusual  dichotomous  branching. 


ASTERELLA 


55 


4-6.  9  receptacles  with  matured  sporogonia,  X  4- 

7-3.  Transversa  s32tiDns  of  thallus,  sho.vinj  in  the  latter  a  $  ini:)vatiDn,  X 

9.  Transverse  section  of  peduncle,  X  4^ 

10.  Capsule  in  dehiscence,  X  '2. 

11.  Capsule  after  dehiscence,  12. 

12.  Elater,  X  225. 

13.  Spore,  outer  fa:e,  X  225.  (The  walls  of  the  areolae  appear  somewhat  too 
rigid.) 

Figures  all  from  soaked-out  dried  material  ;  I-3,  5,ani  7-9  from  specimens  col- 
lected by  Mr.  Geo.  Hansen  near  Jackson,  Amadjr  Co.;  4.  St.  Helena  (Mr.  \V.  L. 
Jepson),  6,  Olema,  Marin  Co.  (Jepson), 

4.  AsTERELLA  Palmeri  (Aust.)  Uiidcrw.  Bot.  Gaz.  20  :  63.  1895. 

Fimbriaria  Palmeri  Aust.  Bull.  Torr.  Bot.  Club,  6  :  47.  1875. 
Steph.  Bull.  Herb.  Boiss.  7:  213.  1899. 

Fimbriaria  nudata  M.  A.  Howe,  Erythea,  i  :  112.  1893. 
Steph.  Bull.  Herb.  Boiss.  7:  212.  1899. 

Asterell a  nudata  Underw.  Bot.  Gaz.  20:  61.  1895. 

Thallus  simple  or  once  dichotomous,  5-10'mm.  long,  2-5  mm. 
in  maximum  width,  the  segments  or  entire  thallus  obcordate 
to  obcuneate,  rather  indistinctly  areolate  above,  dark  purple  be- 
neath, broadly  carinate,  30-40  cells  thick  in  median  parts,  with 
numerous  and  deep  air-chambers,  the  brown  or  purple,  usually 
broad,  undulate-crenate  margins  ascending,  commonly  appressed- 
connivent  on  drying,  scales  dark  purple,  very  rarely  decolorate, 
often  extending  slightly  beyond  the  margin,  a  few  sometimes  in- 
conspicuously inflexed  at  the  apical  sinus  :  monoicous  :  antheridia 
occupying  a  lightly  elevated  oval  or  oblong,  papillate  area  just 
posterior  to  the  base  of  the  9  branch  :  receptacle  maturing  2-6 
(commonly  3  or  4)  sporogonia,  becoming  obtusely  conic-ovoid, 
scarcely  lobed,  sometimes  constricted  near  the  middle  in  drying, 
2.5-4  mm.  in  maximum  width,  nearly  smooth  or  subrugose  when 
dry, -naked  beneath,  air-chambers  large;  peduncle  from  apex  of 
the  thallus,  light  brown  or  stramineous,  12-40  mm.  (mostly  i  5-20 
mm.)  X  -35-65  mm.,  naked,  without  involucre  at  base,  sulcate, 
rather  brittle  ;  pseudoperianth  white,  the  exserted  portion  globose- 
ovoid,  often  strongly  constricted  at  zone  of  emergence,  with  8-12 
narrow  or  somewhat  gaping  clefts,  the  segments  coherent  at  the 
apex  :  capsule  nearly  spherical,  irregularly  circumscissile  above  the 
middle  ;  spores  60-80 at  first  almost  colorless,  marked  only  with 
faint  ridges  or  hnes,  at  maturity  very  opaque,  fuscous  or  black, 
with  a  rather  narrow  warty  border,  the  faces  irregularly  rugose- 
areolate  ;  elaters  i-3-(mostly  2-)  spiral,  120-270//.  x  10-18 /i, 
sometimes  branched,  finally  yellowish-brown. 


56 


Marchantiaceae 


Mill  Valley  (Howe),  St.  Helena  (Jepson\  Jackson  (Hansen), 
Pasadena  (McClatchie),  San  Bernardino  (Parish),  Santa  Monica 
(Hasse) ;  also  at  San  Diego  by  an  unknown  collector  and  at  Witch 
Creek,  (San  Diego  Co.  ?)  (R.  D.  Alderson). 

The  original  specimens  were  collected  by  Dr.  Palmer  on 
Guadalupe  Island,  off  the  coast  of  Lower  California.  Portions 
of  what  appear  to  be  the  original  plants  are  found  in  herb.  Un- 
derwood and  herb.  Columbia  University. 

Aster ella  fragrans  (Schleich.)  Tre\'is. — to  which  A.  Palmcri  is 
probably  most  nearly  allied — has  been  found  in  New  Mexico, 
Idaho  and  Alaska,  and  is  likely  to  occur  in  California.  This  differs 
from  Aster clla  Palineri  in  the  conspicuous  white  scales  projecting 
beyond  the  margin  of  the  thallus,  in  the  presence  of  an  involucre 
of  narrow  whitish  paleae  at  the  base  of  the  peduncle,  in  the  yellow, 
more  broadly  margined  spores,  etc.  The  spores  of  Astcrclla  Palmcri 
show  a  tendency  to  cohere  in  large  masses  until  very  late  in  the 
process  of  ripening — a  character  which  seems  also  to  be  exhibited 
by  A.  fragrans. 

Plate  99. 

I -1 5.  Asterella  ^'ahtieri. 

1.  Plant,  natural  size. 

2.  Thalius,  showing  peduncle  and  position  of  androecium,  X  3- 

3.  Transverse  section  of  thallus  through  androecium,  X  23. 

4.  Median  longitudinal  se:tion  of  thallus,  passing  through  androecium  and  base 
of  peduncle,  X  23. 

5  and  6.  Fully  developed  9  receptacles,  dry,  X  4- 

7-10.  Series  of  cross-sections  of  peduncle  foni  near  apex  to  base,  X  4^- 

11.  Capsule  in  dehiscence,  X  ^2. 

12.  Elater,  X  225. 

13.  Young  spore,  X  225. 

14.  Mature  spore,  X  225. 

15.  Section  of  mature  spore,  X  225. 

Figs.  I,  2,  6,  12  and  13  drawn  from  Guadalupe  Island  material  (legit  Palmer)  ; 
3-5,  7-10  and  15,  St.  Helena  (Jepson);  II  and  14,  Jackson,  Amador  Co.  (Hansen). 

5.  Asterella  gracilis  (Web.  /.)  Underw.  Bot.  Gaz.  20:  61. 

1895. 

Marchaiitia  gracilis  Web.  f.  Hist.  Muse.  Hep.  Prod.  105. 
1815. 

MarcJiantia  pilosa  Wahl.  Fl.  Lapp.  399.  18  12.  Not  M.  pilosa 
Horn.  Fl.  Dan.  8:  7.  pi.  1426.     18 10. 


COXOCEPHALUM 


57 


MarcJiantia  LudicngiiSchwdiGgv.  (?)  Hist.  Muse.  Hep.  Prod.  33. 
1814. 

Fimbriaria  gracilis  Lindb.  Not.  ur  Sallsk.  pro  Fauna  et  Flora 
Fenn.  9:  282.  1868. 

Thallus  simple  or  once  dichotomous,  linear-obcuneate,  some- 
times oblong  ox  obovate,  4-10  mm.  x  1.5-3  nim.,  rather  dis- 
tinctly porose  and  areolate,  canaliculate  or  somewhat  broadly  con- 
cave, dark  purple  underneath,  25—35  cells  thick  in  median  parts, 
with  deep  air-chambers,  the  narrow,  purple  or  concolorous,  undu- 
late-crenate  margins  ascending,  or,  on  drying,  sometimes  erect  and 
connivent,  scales  purple,  reaching  the  margin  or  now  and  then 
projecting  a  little  beyond  :  monoicous  :  antheridia  few,  immersed 
in  the  thallus  just  posterior  to  the  base  of  the  9  branch,  loculi 
with  papilliform  ostioles  :  9  receptacles  maturing  1-4  (mostly  3) 
sporogonia,  subhemispherical,  1.5—3  iriiri-  in  diameter,  nearly 
smooth,  becoming  rugose  in  drying,  naked  beneath,  the  obscurely 
defined  lobes  with  a  thin  membranous  border  ;  peduncle  from  the 
apex  of  the  thallus,  10-40  mm.  x  .3-5  mm.,  light  brown  to  brown- 
ish-purple, naked,  surrounded  at  the  base  by  a  few,  often  inconspic- 
uous scales  or  paleae  ;  pseudoperianth  white,  5— 12- (mostly  8-) 
cleft,  the  segments  early  becoming  free,  most  frequently  somewhat 
inclined  together,  sometimes  spreading  :  capsule  quite  regularly 
circumscissile  above  the  middle  ;  spores  yellow  or  slightly  brown, 
48-64  /i,  narrowly  margined,  areolate -reticulate  or  with  shallow 
alveolae,  the  meshes  5—8  across  each  face,  with  somewhat  thick- 
ened walls  and  angles;  elaters  150-275  fi  x  7-15  /^-,  2-4-spiral, 
more  or  less  attenuate  at  the  ends,  commonly  flexuous  or  con- 
torted, sometimes  branched. 

Near  Mineral  King,  Tulare  Co.  (Coville  and  Funston,  Death 
Valley  Expedition,  nos.  1420  and  15  10,  August  1891). 

8,  CONOCEPHALUM*  Wigg.  Prim.  Y\.  Hols.  82.  1780. 

Fcgatella  Raddi,  Opusc.  scientif.  di  Bologna,  2  :  356.  i8i8.t 
Thallus  large,  dichotomously  branched,  with  long  root-hairs 
springing  in  tufts  from  the  narrow  clearly  defined  costa,  areolae 
very  distinct,  mostly  hexagonal,  stomata  simple  (except  on  $  and 
9  receptacles),  large,  forming  whitish  pustule-like  elevations,  the 
pores  easily  visible  to  the  unaided  eye  ;  chlorophyll-bearing 

*We  have  been  unable  to  verify  this  citation.  A  transcription  of  Wiggers'  diag- 
nosis is  given  by  M.  Auguste  Le  Jolis  (Mem.  Soc.  nation.  Sci.  nat.  et  math.  Cherbourg 
29:  125.  1894). 

t  See  footnote  under  Relwidia. 


58 


Marchaxtiaceae 


stratum  thin,  the  uppermost  cells  in  the  air-chambers  with  a 
conical  or  long-cylindrical  colorless  rostrum  ;  costa  clothed  with 
closely  imbricate  scales,  those  at  the  apex  often  with  a  violet 
orbicular  or  reniform  apical  expansion,  marginal  scales  absent. 
Gemmae-cups  wanting.  Androecium  disciform,  strongly  papillate, 
sessile  in  a  depression  bounded  by  a  somewhat  scarious-mem- 
branous  elevation  of  the  dorsal  layers  of  the  thallus,  termi- 
nating a  short  branch,  this  becoming  apparently  lateral  by  the 
continued  growth  of  the  other  fork.  Q  receptacle  obtusely  con- 
ical, indistinctly  or  not  at  all  lobed,  long-stalked,  surrounded  when 
still  sessile  by  a  slightly  elevated  membranous  sheath  ;  peduncle 
with  a  single  root-hair  furrow,  its  dorsal  (posterior)  surface  desti- 
tute of  assimilative  tissue.  Involucres  4-1 1,  tubular,  confluent, 
constituting  almost  the  whole  of  the  matured  receptacle,  each 
surrounding  a  single  sporogonium.  Pseudoperianth  wanting.  Cap- 
sule oblong-pyriform,  on  a  thick  pedicel  mostly  a  little  more  than 
one  half  as  long,  dehiscing  by  cleavage  about  to  the  middle  into 
4-8  recurved  irregular  valves,  its  final  exsertion  usually  accom- 
panied by  the  detachment  of  the  pedicel  from  the  receptacle,  cap- 
sule-wall of  mainly  a  single  layer  of  thick  cells  with  annular 
and  spiral  thickenings  ;  spores  large,  papillate,  multicellular  at  time 
of  dehiscence  ;  elaters  rather  short  and  thick,  2-4-spiral. 

I.  CoNCCEPHALUM  coxicuM  (L.)  Dumort.  [as  Conoce/}/ia/its~\  Comm. 
Bot.  115.     1822.    Underw.  Bot.  Gaz.  20  :  67.  1895. 

Marchantia  conica  L.  Sp.  PI.  1138.  1753. 
Fegatclla  conica  Corda  ;  Opiz,  Beitr.  i  :  649.  1829.* 
Thallus  oblong  or  linear,  5-25  cm.  x  .75-1.5  cm.;  colorless 
stratum  in  region  of  the  costa  of  20-25  layers  of  small  thick- 
angled  cells,  enclosing  3-5  large  longitudinal  (finally  empty)  muci- 
lage canals  and  passing  abruptly  at  the  sides  into  7-4  layers  of 
larger  cells,  becoming  bi-  (rarely  uni-)  stratose  at  the  extreme 
margin  :  dioicous  :  peduncle  3-6  cm.  high,  naked  :  spores  yel- 
lowish-brown, 66-95  }i  ;  elaters  200-270  x  9-30  /i,  often  dilated 
in  the  middle. 

In  moist  deeply  shaded  places,  especially  on  stones  and  rocks 
beside  streams. 

P'elton,  Santa  Cruz  Co.  (Underwood);  on  stones  in  stream-bed 
near  Lake  Pilarcitos,  San  Mateo  Co.;  on  moist  ground  in  woods, 
"Big  Riv^er  Boom,"  near  ]\Iendocino  ;  and  on  shaded  dripping 
rocks  near  pAireka.  Not  conmion  in  California  ;  first  collected 
by  Professor  Underwood  in  1888. 

*  See  footnote  under  Ricciccarpit  ,  p.  33. 


LUNULARIA 


59 


None  of  the  Californian  specimens  seen  bear  9  receptacles. 
The  measurements  of  spores  and  elaters  have  been  taken  from  a 
plant  collected  near  Seattle,  Washington,  by  C.  V.  Piper. 

Conoccplialiiui  in  a  sterile  state  may  be  readily  distinguished 
from  MarcJiantia  polyuiorpJia  by  the  usually  longer  thallus,  the 
very  distinct  mostly  hexagonal  areolae,  the  simple  (not  com- 
pound "  or  "  dolioform  ")  stomata  with  pores  easily  visible  to  the 
unaided  eye,  the  conical  or  elongate-cylindrical  rostrum  of  the 
upper  green  cells  in  the  air-chambers,  the  absence  of  marginal 
scales,  the  narrow  clearly  defined  costa,  etc. 

9.  LUNULARIA  Adans.  Fam.  PL  15.     1763.    Ex  Mich.  Nov. 
PI.  Gen.  4.  //.  ^.  1729. 

Thallus  moderately  large,  furcate  or  often  progressing  by  in- 
novations from  the  apex,  with  an  effuse  median  thickening,  the 
segments  oblong  or  obcuneate,  with  a  hyaline  unistratose  margin 
1-4  cells  in  width,  areolae  mostly  indistinct  after  drying  ;  stomata 
simple,  the  pores  invisible  to  the  unaided  eye,  the  surrounding 
elevated  cells,  however,  often  whitish  and  readily  visible  ;  chlo- 
rophyll-bearing stratum  narrow  in  a  sectional  view,  occasional  cells 
of  the  colorless  layer  containing  a  large  oil-body  in  the  form  of 
an  emulsion.  Gemmae  in  crescentic  receptacles,  the  thin  entire 
margin  of  the  latter  wanting  in  front.  Dioicous  (sometimes 
monoicous?).  Androecium  sessile,  oval-disciform,  papillate,  sur- 
rounded by  a  slight  elevation  of  the  adjacent  parts.  O  receptacle 
arising  from  a  deep  sinus  of  the  thallus,  surrounded  when  young 
and  sessile  by  a  tubular-ovate  sheath,  consisting  of  numerous 
scales,  the  inner  of  these  membranous,  hyaline,  ciliate-fimbriate  ; 
archegonia  in  commonly  four  groups  of  three  or  four,  only  one  in 
each  group  developing  a  sporogonium  ;  receptacle  destitute  of 
stomata  and  root-hairs,  consisting  at  maturity  almost  wholly  of 
the  slightly  thickened  top  of  the  peduncle  and  of  the  1-6  (usually 
4)  spreading  tubular  involucres  each  surrounding  a  single  sporo- 
gonium. Pseudoperianth  wanting.  Peduncle  without  a  root- 
hair  furrow,  delicate,  pellucid,  pilose,  involucrate  at  base.  Capsule 
dark-brown,  obovoid,  rather  long-stalked,  exserted  from  the 
bilabiate  or  repand-mouthed  involucre,  quadrivalved  to  the  base 
in  dehiscence,  the  valves  often  2 -parted,  more  or  less  twisted  in 
drying,  the  cells  of  the  wall  without  annular  or  spiral  thickenings. 
Spores  yellow  or  brownish,  smooth.  Elaters  bispiral,  very  long 
and  slender,  often  adhering  a  long  time  to  the  ends  of  the  valves. 

Only  one  species  of  the  genus  is  ordinarily  recognized  : 


60 


Marchantiaceae 


I.  LuNULARiA  CRUCiATA  (L.)  Dumort.  Comm.  Bot.  ii6.  1822. 

MarcJiantia  cniciata  L.  Sp.  PI.  1137.  1753- 

Thallus  1-5  cm.  x  .5-1  cm.,  colorless  layer  about  20  cells 
thick  along  the  axis,  becoming  gradually  thinner  to  the  unistratose 
margin  :  peduncle  1.5-3  high  :  spores  15-18  fi  ;  elaters  300- 
650 /i  long,  5-6  fi  broad  in  the  middle. 

In  and  about  greenhouses.  Oakland,  San  Jose,  and  San 
Francisco  (Underwood,  1888);  Berkeley  and  San  Francisco 
(Howe) ;  also  in  the  Golden  Gate  Park,  San  Francisco,  at  con- 
siderable distance  from  any  greenhouse ;  Pasadena  i^fidc  Mc- 
Clatchie).  Introduced  from  Europe.  Only  a  sterile  gemmiferous 
condition  is  known  to  occur  in  America,  but  the  plant  is  easily  rec- 
ognizable by  the  crescentic,  entire-margined  gemmae-cups,  which 
are  almost  always  present.  Our  diagnosis  has  been  completed 
from  European  material  and  from  the  descriptions  of  authors. 

10.  MARCHANTIA  L.  Sp.  PI.  1137.  1753. 
Ex  March,  fii  Act.  Ac.  Paris.  229.  171 3.* 
Thallus  large,  usually  several  times  dichotomous,  with  a  broad 
effuse  costa,  the  root-hairs  abundant,  areolae  mostly  rhombic, 
stomata  dolioform  throughout,  often  not  readily  visible  to  the  un- 
aided eye  ;  air-chamber  stratum  sharply  defined  against  the  under- 
lying colorless  tissue,  of  a  single  layer  of  caverns,  these  filled  with 
branched,  chlorophyll-bearing  filaments.  Gemmae  in  cup-shaped, 
dentate-margined  receptacles  arising  from  the  dorsal  surface  of 
the  costa.  Androecium  long-stalked,  disciform,  with  a  thin 
crenate  margin  or  stellately  or  palmately  lobed,  furnished  with 
scales  on  the  ventral  surface.  0  receptacle  stellate  or  rarely 
subentire,  usually  somewhat  ecceatric,  with  4- 11  often  much 
elongated  and  finger-like,  deflexed  or  spreading  rays,  each  of  the 
latter  containing  a  single  root-hair  canal  communicating  with  its 
ventral  surface  or,  at  distal  extremity,  wholly  enclosed.  Peduncle 
with  two  (very  rarely  three)  root-hair  furrows,  provided  on  its 
dorsal  (posterior)  surface  with  stomata  and  an  air-chamber  layer.! 
Archegonia  commonly  in  8  groups  of  several  each  (sometimes  in 
4  groups,  occasionally  in  other  numbers  by  suppression,  abortion, 

*  Citation  from  Pfeiffer,  Nomenclator  Botanicus. 

f  This,  in  effect,  is  asserted  of  the  genus  Marchantia  by  Stephani  (Bot  Gaz.  17  : 
60  1892.  Bull.  Herb.  Boiss.  7  :  384.  1899).  Most  of  the  numerous  species  of  J/r//  - 
cJiantia  are  exotic  and  we  are  unable  to  assert  from  any  personal  investigations  that  this 
character  belongs  to  all.     It  certainly  holds  good  for  M.  polyniorpho. 


Marchantia 


61 


or  irregular  multiplication)  alternating  *  with  the  rays,  these,  how- 
ever, one  in  excess  of  the  archegonia-groups.  Each  group  of 
archegonia  usually  maturing  several  sporogonia,  these  enclosed 
by  a  common  membranous  2-valved,  fimbriate  involucre,  and 
each  surrounded  by  a  cleft  pseudoperianth.  Capsule  borne  on  a 
seta  as  long  as  the  pseudoperianth,  dehiscing  by  several  revoluble 
teeth  or  short  valves.  Spores  small,  smooth  or  nearly  so.  Elaters 
long  and  narrow,  2-(i-3-)  spiral. 

I.  Marchantia  polymorpha  L.  Sp.  PI.  1137.  1753. 

Thallus  prostrate  or  ascending,  rarely  suberect,  often  in  widely 
extended  masses,  oblong  or  linear,  2-10  cm.  x  .5-2  cm.,  com- 
monly several  times  dichotomous,  about  16  cells  thick  in  the 
middle  of  the  costa,  becoming  gradually  thinner  to  the  narrowly 
unistratose,  mostly  somewhat  undulate-crisped  or  irregularly  short- 
lobed  margin,  areolae  mainly  rhombic,  the  pores  scarcely  visible 
to  the  unaided  eye  ;  the  outer  ventral  scales  oblong,  obtuse,, 
slightly  exceeding  the  margin,  colorless  or  tinged  with  brown  or  pur- 
ple :  dioicious  :  androecium  peltate,  with  commonly  8  short  rounded 
lobes  or  merely  crenate  :  9  receptacle  with  8—1 1  (commonly  9) 
finger-like,  usually  decurved  rays  ;  involucre  fimbriate,  its  segments 
subulate-acuminate  and  ciliate-laciniolate,  often  rufous  ;  peduncle 
of  9  receptacle  2-7  cm.  long,  more  or  less  pilose,  that  of  the  an- 
droecium usually  shorter :  capsule  finally  exserted,  its  wall  uni- 
stratose, with  annular  thickenings  ;  seta  attaining  length  of  1-1.5 
mm.;  spores  yellow,  12-15  «,  nearly  smooth;  elaters  300-600  fJt 
long,  attenuate,  3-5  [jl  in  greatest  width,  bispiral. 

On  wet  banks,  in  bogs,  beside  streams,  about  greenhouses,  etc. 

Widely  distributed  in  California  but  hardly  common.  We  can 
name  only  the  following  localities  :  Berkeley;  San  Francisco  ; 
Mill  Valley,  Olema ;  near  Duncan's  Mills,  Sonoma  Co.;  Mendo- 
cino (577);  near  Eureka;  near  Douglas  City  (1161),  Trinity  Co.- 
Sisson  ;  near  Redding  (Baker  and  Nutting)  ;  Silver  Lake,  Alpine  Co. 
(Geo.  Hansen);  Jackson,  Amador  Co. (Geo.  Hansen); San  Bernardino 
Mts.  (S.  B.  Parish,  nos.  1445,  1446,  1693,  1694,  3369);  vicinity 
of  Pasadena  (A.  J.  McClatchie).  Also  collected  in  California  by 
Dr.  Bolander.  We  have  found  no  mature  capsules  in  any  CaH- 
fornian  specimens  and  our  description  has  been  supplemented  from 
material  collected  in  Europe  and  in  the  eastern  portions  of  the 
United  States. 


*  Exc.  M.  gemifiata  N.  R.  &  Bl.  See  Schiffner,  Ueber  exotische  Hepaticae,  Nova 
Acta  Ksl.  Leop. -Carol.  Deutsch.  Akad.  Naturforsch.  60:   280,  281.  1893. 


62 


J  UXGERM  ANN  I  ALES 


MarcJiantia  Orcgoncnsis  was  described  b}^  Herr  Stephani  in 
1 89 1  (Bot.  Centralb.  45  :  203)  from  $  plants  collected  by  Roll  on 
Mt.  Hood.  The  only  distinctive  character  emphasized  is  a  reni- 
form  irregularly  dentate  and  spinose  appendage  to  the  postical 
scales.  In  a  portion  of  the  original  plant  communicated  by  Herr 
Stephani  to  Professor  Underwood,  we  have  been  able  to  detect  a 
few  such  scales  immediately  bounding  the  costa  at  the  apical  sinus, 
but  there  are  also  oblong  or  linguiform,  obtuse,  entire  scales  toward 
the  margin  as  in  JM.  polymorpJia ;  as  we  have  observed  quite 
similarly  appendaged  scales  at  the  apex  of  the  costa  in  European 
specimens  of  M.  polyniorpha  [c.g.y  Rab.  Hep.  Eur.,  no.  6  ;  Massal, 
Hep.  It.  Ven.  Exs.,  no.  21)  and  also  in  American  specimens,  we  be- 
lieve that  this  peculiarity  has  no  specific  significance.  These  ap- 
pendages are  usually  of  a  violet  or  brownish  color  and  are  formed 
by  a  more  or  less  pronounced  narrowing  of  the  scale  accompanied 
by  a  slight  twisting  or  convolution  in  the  zone  of  contraction. 

Order  II.  JUNGERMANNIALES. 

Gametophore  varying  in  different  genera  from  a  wholly  leafless 
thallus,  with  or  without  a  midrib  but  without  other  differentiation 
of  tissues,  to  a  cylindrical  leaf-bearing  stem.  Root-hairs  always 
with  smooth  walls.  Stomata  wanting.  Sexual  organs  usually  in 
groups,  often  on  more  or  less  modified  branches,  yet  never  on 
special  stalked  receptacles,  rarely  immersed. 

The  first  division  of  the  fertilized  egg  transverse,  the  lower  of 
the  two  cells  usually  taking  no  further  part  in  the  development  of 
the  embryo.*  Sporogonium  provided  with  foot  and  seta,  the  latter 
commonly  much  elongated  ;  calyptra  ruptured.  Capsule  nearly 
always  opening  by  four  valves.  Sterile  cells  always  accompany- 
ing the  spores,  usually  developed  as  elaters.f 

The  order  Jungermanniales  comprises  the  two  somewhat  arti- 
ficial families,  the  Metzgeriaceae  (see  below)  and  the  Jungerman- 
niaceae  (see  page  82). 

*See  Campbell,  The  Structure  and  Development  of  the  Mosses  and  Ferns,  72. 
1895. 

f  Among  the  Californian  genera,  the  sterile  cells  in  Sphaerocarptis  (which  has  sev- 
eral points  of  contact  with  the  Ricciaceae)  maybe  easily  overlooked  unless  the  contents 
of  somewhat  immature  caj^sules  are  studied.  The  sterile  cells  are  here  starch-laden, 
are  destitute  of  spiral  thickenings  and  become  disorganized  or  inconspicuous  by  the  time 
the  spores  are  fully  ripe. 


Metzgeriaceae 


63 


Family  III.  METZGERIACEAE.* 

Apical  cell  of  the  gametophore  never  directly  transformed  into 
an  archegonium.  Sporogonia  arising  from  the  dorsal  surface  of  the 
gametophore  or  only  apparently  from  an  apex.  This  family  con- 
tains all  the  Jungcrmannialcs  in  which  the  gametophore  is  persist- 
ently thallose  and  also  the  forms  transitional  between  the  thallo- 
phytic  and  cormophytic  types.  The  aberrant  genera  Haploinitriinn 
and  Calobryujfi  do  not  occur,  so  far  as  is  known,  on  the  North 
American  continent.  The  involucres  of  the  sexual  organs  and  of 
the  sporogonium  very  rarely,  if  ever,  represent  true  leaves. 

Key  to  the  Genera  of  Metzgeriaceae 
Seta  very  short,  less  than  y'^mm.  long  ;  spores  accompanied  by  sterile  cells  without 
spiral  thickenings. 

9  gametophore  suborbicular,  the  somewhat  leaf-liice  peripheral  lobes  almost  en- 
tirely concealed  by  the  numerous  subglobose,  cylindrical-ovoid,  or  fusiform- clavate 
involucres  ;  annual  II.  Sphaerocarpus. 

Gametophore  a  somewhat  elongated  thickened  axis,  with  conspicuous  crowded  ir- 
regularly succubous  foliar  expansions  on  either  side  ;  perennial  through  the  con- 
version of  a  large  portion  of  the  axis  into  a  tuber. 

12.  Geothallus. 

Seta  4-50  mm.  long  ;  elaters  always  developed. 

Gametophore  with  quite  clearly  defined  stem  and  leaves  ;  root-hairs  very  long, 

mostly  violet  or  vinous-purple.  16.  Fossombronia . 

Gametophore  thalloid,  without  ventral  scales  and  without  special  gemmae-recep- 
tacles. 

Thallus  rather  fleshy,  pinnately,  palmalely,  or  subdichotomously  branched,  the 
unistratose  marginal  lamina,  if  present,  narrow  and  passing  gradually  into 
the  costa  ;  antheridia  and  archegonia  on  short  lateral  branches ;  spores 
unicellular.  13.  Riccardia.-\ 

Thallus  irregularly  dichotomous,  the  wide  unistratose  margin  passing  gradu- 
ally into  the  indistinct  costa  ;  antheridia  in  loculi  on  the  back  of  the  costa, 
archegonia  in  a  cavity  near  its  apex  ;  spores  multicellular  at  time  of  de- 
hiscence. 14.  PelIia.-\ 
Thallus  with  a  narrow  costa  sharply  limited  from  the  unistratose  marginal 
laminae,  mostly  dichotomous  ;  antheridia  and  archegonia  on  short  ventral 
branches.  Metzgeria  (p.  75). 
Gametophore  thalloid,  but  the  thin  margin  often  with  more  or  less  leaf -like  lobes  ; 
cavities  near  the  bases  of  the  lobes  commonly  occupied  by  Nostoc  colonies  ; 
ventral  surface  bearing  small  dentate  scales  ;  gemmae  in  long-necked,  flask- 
shaped  receptacles  arising  from  the  dorsal  surface  near  the  ends  of  the  lobes  ; 
.spores  unicellular.                                                       15.  Bla  ia. 

*  Underwood,  Bot  Gaz  19:  361.  1894.  The  "  Jungermanniaceae  anakrogynae  " 
of  Leitgeb  and  of  Schiflher. 

f  Sterile  conditions  of  RiccarJia  pingiiis  and  Pellia  sp.  may  be  easily  confused. 
See  remarks  under  R.  pingui<. 


64 


Metzgeriaceae 


II.  SPHAEROCARPUS*  Adans.  Fam.  PL  2:  14.    1763.  Ex 

Mich.  Nov.  PI.  Gen.  4.  //.  j.  1729. 

Gametophore  small,  orbicular,  oblong,  or  cuneate,  thallus-like 
and  irregularly  lobed  or  now  and  then  subfoliose,  of  a  few  layers 
of  nearly  similar  cells  in  central  or  median  parts,  the  lobes  unistra- 
tose  unless  at  base,  ascending  or  flat ;  root-hairs  smooth,  colorless, 
long  and  numerous.  Dioicous.  Sexual  organs  in  most  cases 
numerous  and  thickly  covering  the  dorsal  surface  of  the  thallus, 
surrounded  singly,  rarely  in  groups  of  2  or  3  {fide  Leitgebt),  by 
unistratose  involucres.  plants  much    smaller  than  the  9 , 

scarcely  visible  to  the  unaided  eye,  commonly  tinged  with  purple, 
oblong  or  cuneate,  often  once  dichotomous  or  subpalmately 
lobed.  Antheridia  oval,  on  very  short  stalks,  the  involucres 
flask-shaped  or  subcorneal.  Archegonial  involucres  usually 
crowded  and  more  or  less  completely  concealing  the  thallus,  from 
tubular  or  clavate  to  pyriform  or  subglobose,  orifice  reduced  to  a 
small  apical  pore,  this  often  enlarged  at  maturity.  Calyptra  of  2 
or  3  layers  of  cells,  usually  ruptured  early,  the  upper  portion, 
with  the  archegonium  neck,  long  persisting  on  the  apex  of  the 
capsule. 

Sporogonium  consisting  of  a  nearly  spherical  capsule,  a  very 
short  or  almost  obsolete  seta,  and  a  well-developed  foot.  Cap- 
sule indehiscent,  its  wall  of  a  single  layer  of  cells  destitute  of 
spiral,  annular,  or  other  local  thickenings.  Spores  permanently 
adherent  in  4's,  or  (5.  Do?tnellii  Anst.)  becoming  free  at  full  matur- 
ity, or  (5.  cristatus  M.  A.  Howe)  separating  early  in  the  develop- 
ment of  the  sporogonium  ;  tetrads,  when  persistent,  distinctly 
areolate,  appearing  crenulate,  papillate,  or  subechinulate  in  optical 
section  ;  spores,  when  separating,  compressed-tetrahedral,  rounded - 

*  Mr.  James  Britten  (Journal  of  Botany,  36  :  397-399.  1898)  has  recently  called 
attention  to  the  use  in  1759  and  1763  of  Sphaerocarpus  as  a  name  for  the  cruciferous 
genus  commonly  known  as  Neslia.  The  name  is  said  to  occur  in  both  the  first  and 
second  editions  of  Fabricius'  "  Enumeratio  Plantarum  Horti  Helmstadiensis,"  pub- 
lished at  the  dates  mentioned,  respectively.  In  the  first  edition,  which  alone  has  clear 
priority  over  Adanson's  work,  *'  Sphaerocarpus  Heister  "  appears  simply  as  a  synonym 
of  **  Rapistrum  arvense  fo  io  auriculato  T."  It  would  seem  that  the  usual  interpreta- 
tion of  the  Rochester  Code  [the  interpretation  which  denies  the  publication  of  Dalca  as 
a  genus  in  the  Species  Plantarum  of  Linnaeus  (1753)]  i^^st  be  unable  to  recognize 
the  publication  of  Sphaerocarpus  as  a  genus  in  the  above  use  of  the  word  in  the 
first  edition  of  Fabricius'  "  Enumeratio."  In  any  event,  it  would  be  a  gross  violation 
of  the  spirit  of  the  priority  principle  to  annul  the  original  Sphaerocarpus  of  Micheli, 
grandly  described  and  figured  in  1729,  in  order  to  validate  the  application  thirty  years 
later  of  the  same  name  to  a  totally  different  genus. 

f  Leitgeb,  Untersuchungen  iiber  die  Lebermoose,  4  :  67,  68.  1879. 


Sphaerocarpus 


65 


lenticular,  or  sometimes  concavo-convex,  areolate  or  cristate,  in 
.S".  Dvinellii  prom'.nenLly  tuberculate-papillate  in  the  basilar  cir- 
cumference. Accompanying  the  spore-mother-cells  are  smaller 
starch-bearing  sterile  cells  probably  equivalent  to  elaters  morpho- 
logically, but  lacking  a  spiral  band,  these  more  or  less  shriveled 
and  obscure  at  the  maturity  of  the  spores. 

Key  to  tlie  Species. 

Spores  permanently  united  in  tetrads,  the  surface  of  these  regularly  areolate  ;  9  invo- 
lucre 1.2-2.6  mm.  high,  oblong-cylindrical,  fusiform-clavate,  or  very  rarely  subpyri- 
form,  more  or  less  pointed  at  the  apex,  cells  near  the  middle  90-150,//  X  30-50 

S.  Ca  ifornicus. 

Spores  separating  long  before  the  maturity  of  the  capsule,  their  surface  t}-pically  cristate, 
less  commonly  irregularly  and  sparingly  areolate;  9  involucre  .85-1.2  mm.  high, 
subglobose  or  obovoid,  rounded  at  the  apex,  cells  near  the  middle  45-80  a  X  30-40 

S.  cristatus. 

I.  Sphaerocarpus  Califorxicus  Aust.  Bull.  Torr.  Bot.  Club,  6: 

305.  1879. 

Sphaerocarpus  Bej'teri  Aust.  p.  p.  Hep.  Bor.-Am.  138.  1873. 
Not  Mont.  Ann.  Sci.  Nat.  Bot.  II.,  9:  39.  1838. 

Sphaerocarpus  Michelii  Bell.,  var.  Califoniiciis  Aust.;  Underw. 
Bull.  111.  State  Lab.  Nat.  Hist.  2  :  30.  1884. 

Sphaerocarpus  tcrrestris  Mich.,  var.  Californiciis  Underw.  Zoe, 
I  :  364.  1890. 

O  thallus  suborbicular,  somewhat  stipitate,  mostly  4-8  mm. 
in  diameter,  commonly  in  thickly  gregarious  masses,  the  often  leaf- 
like lobes  almost  entirely  concealed  by  the  involucres,  these  1.2- 
2.6  mm.  high,  oblong-cylindrical,  fusiform-clavate,  very  rarely 
subpyriform,  sessile,  more  or  less  acuminate,  the  orifice  usually 
inconspicuous  ;  marginal  cells  of  the  thallus  about  45  fi,  medial 
surface  cells  60—90  /v.,  cells  near  mouth  of  9  involucre  45-60  /i, 
near  the  middle,  90-1 50/>«x  30-50  :  ^  thallus  oblong,  commonly 
furcate  or  subpalmately  parted,  the  flask-shaped  antheridial  invo- 
lucres .27—36  mm.  high  :  capsule  .5—7  mm.  in  diameter,  seta 
scarcely  perceptible  at  maturity,  the  foot  commonly  remaining  in 
thallus  when  the  capsule  is  detached  ;  spores  permanently  united 
in  tetrads,  these  105-140 /7.  (mostly  120/^)  in  diameter,  dark 
brown,  -areolate,  the  meshes  almost  regularly  pentagonal  or  hexa- 
gonal or  somewhat  elongated,  15-30  a  in  diameter,  the  tetrad  ap- 
pearing crenulate  or  slightly  papillate  in  optical  section  owing  to 
the  somewhat  increased  elevation  of  the  ridges  at  the  angles  of 
the  meshes,  the  entire  surface  very  minutely  granulate  ;  sterile 
cells  subspherical  to  irregularly  oblong-cylindrical,  30-60  a  in 
longest  diameter,  evanescent  at  maturity  of  the  spores. 


66 


Metzgeriaceae 


Exsicc.  Hep.  Bor.-Am.  138  {p.  p.). 

On  flat  compact  soil  in  meadows  and  besides  paths,  less  com- 
monly on  moist  banks.  Berkeley  ;  San  Francisco  ;  Mill  Valley  ; 
Stanford  University  (Campbell),  Reche  Canon,  vicinity  of  San  Ber- 
nardino (Parish,  no.  3613);  Pasadena  (McClatchie).  Collected 
also  by  Brewer  and  by  Bolander. 

SpJiaerocarpus  Californiais  is  closely  related  to  Sphacrocarpus 
Sphaerocarpiis  (Dicks.),*  differing  chiefly  in  the  longer  and  more 
pointed  cylindrical-oblong  or  fusiform-clavate  involucres,  which  in 
the  latter  species  are  commonly  pyriform. 

Plate  100. 
Figs.  9-12,  Sphaerocarpiis  Californiciis. 

All  figures  are  drawn  from  specimens  collected  in  ]Mill  Valley,  Marin  Co. 

9.  9  plant,  with  matured  sporogonial  involucres,  natural  size. 

10.  $  plant,  X  12. 

11.  Portion  of  9  thallus  with  involucres  of  various  ages,  X  23. 

12.  Spore- tetrad,  X  305- 

2.  Sphacrocarpus  cristatus  sp.  nov. 

9  thallus  suborbicular,  3-8  mm.  in  diameter,  marginal  cells  of 
the  unistratose  lobes  3C-45  //.,  medial  45-60 /v. :  involucres  .85- 
1.2  mm.  high,  sessile,  thickly  aggregated,  at  first  cylindrical-ob- 
long, soon  becoming  subglobose  or  obovoid,  rounded  at  the  apex, 
irregularly  circumscissile  with  age  at  about  the  middle  or  the  upper 
portion  falling  away  in  fragments,  cells  near  the  usually  conspicu- 
ous orifice  30-40/.?,  near  the  middle  45-80/^  x  30-40  «:  $  plant 
often  once  furcate  with  oblong-ovate  segments  :  capsule  .5-. 8  mm. 
in  diameter,  on  a  seta  about  40  in  length,  the  bulbous  foot  usu- 
ally accompanying  the  capsule  when  this  is  detached  from  the 
thallus  ;  spores  separating  long  before  the  maturity  of  the  capsule, 
never  persisting  in  tetrads,  compressed,  rounded-biconvex  or  some- 
times concavo-convex,  yellowish-brown,  65-80  (t  in  maximum 
diameter,  cristate,  the  crests  sinuous,  4-7  (i.  high,  subcrenulate, 
t8-22  in  number  in  the  basilar  circumference,  mostly  radiating 
from  near  the  middle  of  each  of  the  two  faces,  often  1-3  times 
furcate,  sparingly  anastomosing,  forming  sometimes  1-6  (rarely 
more)  completely  closed  meshes  in  most  cases  near  the  middle  of 

*  Tar^ionia  Sphacrocarpus  Dicks.  PI.  Crypt,  i  :  8.  1785. 
Sphaerocarpiis  tcrrcs/ris,  minima,  Mich.  Nov.  PI.  Gen.  4.  //.  j.  1729. 
Spaerocarpiis  Michc/ii  Pell.  App.  ad  Fl.  Pedem.    52.     1792  (  aiiciornm). 
Mem.  Acad.  Roy.  Sci.  Turin,  5  :  25S.  1793. 

Sphacrocarpus  tcrrcsfris  Sm.  Eng.  P)0t.  //.  2g'/.  1796. 


Sphaerocarpus 


67 


the  face  ;  sterile  cells  cubico-spherical  to  oblong-cylindrical,  45-70 
u.  in  longest  diameter,  obscure  at  maturity  of  the  spores. 

Exsicc.  Hep.  Am.  160  (as  Sphaerocarpus  terrestris  Mich.,  var. 
Calif orniciis  Aust.,  a  little  of  which  is  intermingled  in  the  three  sets 
examined). 

California:  near  Stanford  University,*  May,  1892  (Professor 
D.  H.  Campbell);  Berkeley,  Feb.  15,  1896  (Howe). 

Sphaerocarpus  cristatiis  is  a  ver}'  remarkable  species  with  no 
near  relatives,  so  far  as  can  be  ascertained  from  accessible  litera- 
ture and  specimens.  The  spores  separate  very  early — at  a  time 
when  they  are  still  colorless  and  only  40-50  in  diameter  and 
when  the  surface  markings  have  become  scarcely  visible.  This 
early  dissolution  of  the  tetrad  is  a  character  of  almost  generic 
importance  yet  the  gap  in  this  respect  between  Sphaerocarpus  cris- 
tatus  on  the  one  hand  and  Sphaerocarpus  Californicus  Aust.  and 
Sphaerocarpus  Sphaerocarpus  (Dicks.)  on  the  other,  in  which,  so 
far  as  we  have  observed,  the  spores  are  permanently  united  in  4's, 
is  bridged  over  somewhat  by  vS.  Donnellii  Aust.,  from  Florida,  in 
which  the  spores  separate  at  maturity.  But  the  spores  of  ^.  Don- 
nellii are  quite  different  in  their  more  angular  outline  and  in  their 
markings,  being  areolate  instead  of  typically  cristate  and  being  also 
prominently  tuberculate-papillate  in  the  basilar  circumference. 

Professor  Campbell  (Erythea  4 :  73-77.  1896)  was  the  first 
to  observe  and  record  the  separation  of  the  spores  in  Californian 
specimens  of  Sphaerocarpus  and  it  seems  probable  from  his  de- 
scription and  drawings  that  his  interesting  Notes  on  Sphaero- 
carpus "  (/.  r.)  were  based,  chiefl}^  at  least,  upon  5.  cristatus. 

Hep.  Am.  160  in  the  herbarium  of  Professor  Underwood  we 
would  name  as  the  type  of  the  species. 

Plate  100. 
Figs.  1-8,  Sphaerocarpus  cristatus. 
All  figures  are  drawn  from  the  type  material. 

1.  9  P^ant,  with  matured  involucres  and  sporogonia,  natural  size. 

2.  Portion  of  thallus  with  involucres  of  various  ages,  X  23. 

3.  Sporogonium,  X  23. 

*The  locality  given  in  Hep.  Am.  160  is  "  San  INIateo  Co,,"  but  Professor  Camp- 
bell writes  in  reply  to  a  request  for  a  more  precisely  defined  station  that  nearly  all  his 
Hepaticae  have  been  collected  within  the  limits  of  Santa  Clara  County,  in  which  Stan- 
ford University  is  situated. 


68 


Metzgeriaceae 


4  Antheridial  involucre,  X  23. 

5  and  6.  Opposite  faces  of  a  single  spore,  X  3^5  (^^^  former  is  thought  to  be 
what  was  originally  the  outer  face,  chiefly  because  the  ridges  are  here  a  little  the 
stronger     The  number  of  completely  closed  meshes  in  the  latter  is  unusually  great). 

7  and  8.  Spores,  X  3^5 »  ^^^^  latter  showing  the  flattening. 

12.  GEOTHALLUS  Campb.  Bot.  Gaz.  21:  13.        2.  1896. 
Ann.  Bot.  10:  489.  //.  2/j.,  2j.  1896. 

Gametophore  transitional  in  character  between  the  thalloid  and 
foHose  types  of  Hepaticae,  consisting  of  a  simple  or  dichotomous 
flattened  axis  several  layers  of  cells  in  thickness,  passing  some- 
what abruptly  at  the  sides  into  large  crowded  irregularly  succubous 
mostly  unistratose  leaf-like  expansions  and  often  bearing  smaller 
irregularly  disposed  leaf-like  laminae  on  the  dorsal  surface  ;  cells 
large,  thin-walled  ;  root-hairs  numerous,  long,  colorless.  Dioicous. 

and  9  plants  similar,  or  the  former  slightly  smaller.  Sexual 
organs  usually  few,  often  standing  near  the  axils  of  the  leaf-like 
lobes,  enclosed,  in  most  cases  singly,  by  unistratose  involucres. 
Antheridia  ovoid,  on  very  short  stalks,  these  composed  of  more 
than  a  single  vertical  series  of  cells,  involucre  flask-shaped. 
Archegonial  involucres  at  first  cylindrical  or  somewhat  flask - 
shaped,  becoming  saccate  or  campanulate  at  maturity  of  the 
sporogonium,  the  orifice  large.  Calyptra  mostly  of  two  layers  of 
cells,  ruptured  rather  late,  the  upper  part,  with  the  archegonium- 
neck,  persisting  on  the  apex  of  the  capsule. 

Sporogonium  consisting  of  a  nearly  spherical  indehiscent  cap- 
sule, a  very  short  seta,  and  a  bulbous  foot.  Capsule  wall  of  a 
single  layer  of  large,  dark- colored  cells,  without  annular  or  other 
local  thickenings.  Spores  separate  at  maturity,  very  large,  thick- 
walled,  the  inner  face  reticulate,  otherwise  smooth,  accompanied  by 
oval  or  ellipsoidal-cylindric,  thin-walled,  sterile  cells. 

I.  Geothallus  tuberosus  Campb.  /.  c. 

Plant  simple  or  once  dichotomous,  5-7  mm.  long,  3-5  mm. 
wide,  perennial  through  the  conversion  of  a  large  portion  of  its 
axis  into  an  oval  or  flattened  tuber  :  leaf-like  lobes  nearly  hori- 
zontal or  somewhat  ascending,  extremely  variable  in  form,  mostly 
oblong,  obovate,  or  linear-ligulate,  1.5-2  mm.  in  length,  rarely 
broader  than  long,  sometimes  cristate-laminate  at  base,  the  mar- 
gins entire  or  slightly  sinuate  ;  marginal  cells  pentagonal,  or  sub- 
quadrate-oblong,  50-65  X  35  ft,  the  submarginal  hexagonal-oblong 
or  irregularly  pentagonal,  50-100  //  x  35-60  a,  those  toward  the 
base  of  the  lobes  often  becoming  over  200  ft  in  length  :  $  invo- 
lucre .45  mm.  in  height:  capsule  .8  mm.  in  diameter,  seta  about 


Geothallus 


69 


90  a  long  ;  spores  nearly  black,  1 20-1  50//  in  maximum  diameter, 
spDre-wall  smooth  or  very  minutely  punctulate  and  very  thick 
(8-12  a)  except  as  to  the  comparatively  small  inner  face  where  it 
is  thinner  and  bears  reticulate  ridges,  the  meshes  about  i  5  fi  in 
width  ;  sterile  cells  48-108  ti  long. 

On  sandy  soil,  near  San  Diego  (Mrs.  Katharine  Brandegee  ; 
March,  April,  1895),  associated  with  OpJiioglossuin  nudLcaii'c. 

A  remarkable  plant,  clearly  allied  to  SpJiaerocarpus,  yet  as 
clearly  distinct  from  it  generically.  Living  laboratory-grown 
specimens,  bearing  antheridia  and  archegonia,  and  mounted  micro- 
tome sections  of  the  mature  sporogonium,  have  been  furnished  us 
through  the  generosity  of  Professor  Campbell.  Mature  9  invo- 
lucres we  have  not  seen  and  our  description  of  these  is  adopted 
from  the  author's  description  with  the  assistance  of  his  figure  5 
(Bot.  Gaz.  /.  <:.).  We  have  seen  the  spores  only  in  microtome 
sections,  yet  it  seems  certain  from  these  that  the  spore-surface  is 
reticulate  upon  the  inner  face  and  otherwise  practically  smooth,  as 
described  by  Professor  Campbell — a  unique  character,  so  far  as 
our  observations  upon  the  Hepaticae  go.  In  Anthoceros,  Riccia, 
Fossoinbronia,  etc.,  it  is  always  the  outer  face  of  the  spore  that  is 
the  more  conspicuously  roughened.  Doubtless  the  restriction  of 
the  ridges  in  Geotlialhis  to  the  inner  face  is  correlated  in  some  way 
with  the  unusual  thickness  and  rigidity  of  the  spore -wall  in  the 
region  of  the  outer  face,  especially  as  the  ridges  of  the  inner  face 
are  seen  in  section  to  be  formed  by  a  folding  of  the  exospore,  the 
contour  of  which  is  followed  by  the  episporic  coating. 

The  gametophore  of  Geothallus  (as  grown  in  the  laboratory) 
much  resembles  in  general  appearance  that  of  Fossonibronia  longi- 
seta,  though  the  "leaves"  are  stiffer,  less  crisped,  and  more  hori- 
zontal. 

The  formation  of  tubers  is  a  character  which  this  hepatic 
shares  with  AntJioccros  dicJwtoinus  Raddi,  Anthoccros  Donnellii 
Aust.,  Anihoceros  tubcrosus  Tayl.,  AntJioccros  pJiymatodcs  M.  A, 
Howe,  Riccia  vcsicata  Tayl.  (=  R.  cancellata  Tayl.  fide  Stephani), 
Riccia  pcrennis  Steph.,  Riccia  bulbifera  Steph.,  Petalophylhnn 
Prcissii  Gottsche,  PetalopJiylluin  laincllatiiin  (Hook.)  Lindb.,  and 
Fossonibronia  tuberifcra  Goebel.  The  stem  of  Fossonibronia 
longiscta  also  is  often  more  or  less  tuberously  thickened  at  the 
apex,  especially  in  specimens  from  the  southern  part  of  Cah- 


70 


Metzgeriaceae 


fornia.  This  adaptation  for  carrying  the  plant  over  a  season  of 
drought  will  doubtless  be  found,  when  the  species  from  the  more 
arid  regions  of  the  earth  come  to  be  better  known,  to  be  more 
common  among  the  Hepaticae  and  Anthocerotes  than  has  been 
generally  supposed,  as  has  already  been  remarked  by  Herr 
Stephani.  From  the  above  showing  it  will  be  seen  that  the  occur- 
rence of  tubers  in  GeotJiallus  can  receive  no  emphasis  in  the  gen- 
eric diagnosis,  inasmuch  as  this  is  elsewhere  evidently  a  specific 
rather  than  a  generic  character. 

13.  RICCARDIA  S.  F.  Gray,  [as  Riccardms^^  Nat.  Arr.  Brit.  PL 

683.  1821. 

Armira  Dumort.  Comm.  Bot.  115.  1822. 

Gametophore  a  pinnately,  palmately,  or  subdichotomously 
branched,  rarely  almost  simple,  somewhat  fleshy  thallus,  composed, 
for  the  most  part,  of  several  layers  of  cells,  with  the  interior  cells 
considerably  larger  and  more  elongated  than  those  of  the  surface, 
usually  undifferentiated  into  costa  and  lamina  though  sometimes 
provided  with  a  unistratose  border,  this,  in  its  best  development, 
giving  the  thallus,  especially  the  younger  branches,  the  appearance 
of  being  broadly  costate.  Root-hairs  few.  Monoicous,  dioicous, 
or  rarely  paroicous,  occasionally,  perhaps,  polyoicous.  Antheridia 
spherical,  immersed  singly  in  loculi,  commonly  arranged  in  two 
parallel  rows  on  a  small,  rounded,  or  most  often,  oblong  or  linear, 
lateral  branch.  Archegonia  several,  on  a  short  subovate  lateral 
branch,  this  by  subsequent  growth  of  the  main  thallus  sometimes 
appearing  to  be  attached  ventrally  ;  margin  of  the  9  branch  usu- 
ally furnished  with  numerous  narrow  laciniae,  these  often  produced 
into  single  rows  of  cells  ;  base  of  the  archegonium  confluent  with 
the  thallus,  the  latter  contributing  more  or  less  to  the  formation  of 
the  "calyptra."  Calyptra  tubular  or  pyriform,  large,  fleshy,  its 
walls  several  layers  of  cells  in  thickness,  the  surface  roughened 
near  the  apex  when  young  [exc.  in  Riccardia  pinnatifida  (?)]  by 
stout  papillae  or  short  trichomes.  Involucres  external  to  the 
calyptra  represented  only  by  the  fringed  margin  of  the  9  branch, 
this  becoming  more  or  less  coalesced  with  the  base  of  the  calyptra 
and  usually  quite  inconspicuous. 

Sporogonium  an  oval  or  oblong-ellipsoidal  capsule  on  a  some- 
what long  seta.  Capsule  before  dehiscence  with  a  short  internal  axile 
column  at  the  apex,  formed  as  if  by  continuation  of  the  inner  of 
the  two  layers  of  cells  composing  the  capsule-wall,  this  column 
separating,  on  dehiscence,  into  four  parts — the  **  elater-bearers  " — 


RiCCARDIA 


71 


one  portion  remaining  attached  to  the  apex  of  each  valve.  Elaters 
unispiral,  mostly  somewhat  attenuate  at  the  ends,  a  few  usually 
clinging  to  the  elater-bearers  "  and  forming  with  them  at  the  tip 
of  each  valve  a  pencil-like  tuft.  Inner  layer  of  the  capsule-wall 
with  annular  or  spiral  thickenings.     Spores  small. 

Key  to  the  Species. 

Main  axis  of  the  thallus  3-6  mm.  wide.  I,  R.  pinguis. 

Main  axis  or  principal  branches  less  than  2  mm.  wide. 
Ramification  pinnate. 

Unistratose  margin  of  branches  one  cell  in  width,  obsolete  in  older  parts. 

2.  R.  viajor. 

Unistratose  margin  of  branches  two  or  three  cells  in  width. 

3.  R.  nmltijida  affibrosioides- 

Ramification  palmate. 

Thallus  translucent  when  moist,  surface  cells  45-118  X  29-50//,  branches 
with  unistratose  margin  one  cell  in  width  ;  monoicous. 

4.  R.  latifrons. 

Thallus  opaque,  surface-cells  23-59  X  I5--9  branches  narrower,  usually 
without  trace  of  a  unistratose  margin,  almost  always  gemmiferous  at  apex  ; 
dioicous.  5.  R.  palm  at  a. 

I.  RiCCARDIA  PINGUIS  (L.)  S.  F.  Gray  (as  Riccardius)^  Nat.  Arr. 
Brit.  PI.  I  :  6S4.     1 82 1. 

Jungerinannia  pinguis  L.  Sp.  PI.  11 36.  1753. 

A neia^a  pinguis  Dumort.  Comm.  Bot.  115.  1822. 

Thallus  broad,  thick,  rigid  on  drying,  dull  green,  with  a  some- 
what greasy  lustre,  simply  pinnate  or  subpinnate,  the  main  axis 
3—6  mm.  wide,  apices  rounded  :  dioicous  :  androecia  suborbicular, 
becoming  short-oblong,  .76—1.7  mm.  x  .85  mm.,  often  geminate, 
margin  entire  or  crenulate  ;  antheridia  4-13,  irregularly  disposed  : 
9  branch  with  a  laciniate  margin,  archegonia  few  ;  calyptra  3.5—5 
mm.  high,  tuberculate  or  nearly  smooth  :  capsule  oval,  brown  ; 
elater-bearers  conspicuous,  j^—  the  length  of  the  valves  ;  elaters 
contorted,  mostly  145-300 /v.  x  9-12  a;  spores  brown,  minutely 
papillose,  1 8-24  u.. 

In  mountain  streams  and  very  wet  places.  Hay  Fork,  Trinity 
Co.  (no.  1 119);  Marin  Co.  (Underwood). 

The  larger  simpler  forms  of  this  species  may,  when  sterile,  be 
confused  with  certain  sterile  conditions  of  the  genus  Pellia.  Dis- 
tinguishing marks  are,  the  pinnate,  instead  of  dichotomous,  branch- 
ing, apices  rounded  rather  than  emarginate,  texture  more  rigid 
when  dry,  and  a  lustre  as  if  saturated  with  some  oleaginous  com- 
pound.   The  sporogonium  has  not  been  found  in  the  Californian 


72 


Metzgeriaceae 


specimens.  The  abov^e  description  has  been  completed  from  New 
York  plants  in  herb.  Underwood. 

2.  RiccARDiA  MAJOR  (Necs)  Lindb.  Muse.  Scand.  5.  1879. 
Aneiira  mnltifida  major  Nees,  Naturgesch.  Eur.  Leberm.  3  : 
450.  1838. 

Thallus  rather  broad  (main  axis  .66-1.7  "i^-  ^vide),  the  branch- 
ing sometimes  regular  and  compact  but  more  frequently  loose  and 
irregular,  always  pinnate  or  bipinnate  ;  ultimate  branchlets  usually 
slightly  biconvex  but  becoming  plane  or  even  concave  ventrally 
or  dorsally,  the  unistratose  margin  one  cell  in  width,  obsolete  in 
the  older  portions  :  monoicous  :  androecium  usually  distant  from 
9  branch,  rather  short,  oblong  or  subovate,  .23-6  x  .17-32  mm., 
furnished  with  an  erect,  translucent,  crenulate  margin  and  bearing 
1-6  (usually  3)  pairs  of  loculi  :  9  branch  broad,  its  margin  mul- 
tilaciniate  ;  calyptra  arcuate,  clavate-cylindrical,  2.5-4.5  mm.  high, 
.84-1.26  broad  above,  much  roughened  when  young  by  short 
tubular  trichomes,  nearly  smooth  at  maturity  :  capsule  oblong- 
ellipsoidal,  dark-brown  or  black ;  elater-bearers  about  |— 1-  the 
length  of  the  valves'^;  elaters  mostly  290-500  (i  long,  12  />«  in 
maximum  width,  the  spiral  band  disappearing  in  the  gradually  at- 
tenuate ends,  a  few  (from  near  the  elater-bearers?)  180  x  15 
obtuse;  spores  brown,  minutely  papillose,  14-18  //. 

Exsicc.  Hep.  Am.  26  (as  Aneiira  pinnatifida  Nees). 

Common  in  the  Coast  Range  Mountains  on  moist  banks  and  on 
logs  and  stones  about  streams.  Mill  Valley  (1185,  1 1 86),  Marin 
Co.;  Cazadero  (12 12)  and  Duncan's  Mills  (12 13),  Sonoma  Co.; 
Ukiah  (744),  Navarro  (Miss  Edith  S.  Byxbee),  and  Mendocino 
(550),  Mendocino  Co.;  Blue  Lake,  Humboldt  Co.;  Hay  Fork 
(i  1 18),  Trinity  Co. 

So  far  as  California  is  concerned,  this  plant  is  usually  very 
distinct  from  R.  mnltifida  ambrosioides  and  it  has  seemed  conduciv^e 
to  clearness  to  follow  Lindberg  in  treating  the  variety  of  Nees  as 
a  species.  Yet  it  must  be  admitted,  we  think,  that  transitional 
forms  do  occur  in  California,  though  rarely. 

Riccardia  pinnatifida  we  do  not  yet  recognize  from  California. 
Hep.  Am.  no.  26,  from  Marin  Co.  seems  to  go  better  with  Riccardia 
major.    Riccardia  pinnatifida  differs  from  the  latter  species  in  the 

-  In  measuring  the  "  elater-bearers  "  the  term  is  here  restricted  to  the  somewhat 
compact  coKimn  made  up  of  rigid  tubes  with  annular  or  spiral  thickenings  and  with 
the  fiee  extremities  rounded. 


RiCCARDIA 


73 


softer,  flatter  tballus,  without  indication  of  a  unistratose  margin,  in 
the  looser  texture,  and  according  to  authors,  in  being  dioicous  and 
in  having  a  smooth  calyptra.  All  specimens  of  Riccardia  pinnati- 
Jida  that  we  have  had  an  opportunity  of  studying  have,  unfortu- 
nately, been  sterile.  It  appears  to  be  an  aquatic  plant,  for  the 
most  part. 

3.  Riccardia  multifida  ambrosioides  (Xees)  Lindb.  Acta  Soc. 

Sci.  Fenn.  10  :  511.  1875. 

Aiiciira  inidtifida  ambrosioides  Nees,  Xaturgesch.  Eur.  Leberm- 
3:  450.  1838. 

Thallus  bi-  or  tri-pinnate,  often  expanding  to  form  radiating 
clusters  or  half-rosettes,  the  branches  crowded,  rather  narrow 
(.34-. 8 5  mm.  broad),  biconvex  with  a  unistratose  margin  two  or 
three  cells  in  width,  thus  presenting  a  quite  clearl}'  defined  sugges- 
tion of  a  costa  :  monoicous,  sometimes  paroicous  (rarely  dioicous  ?): 
androecium  usually  distant  from  9  branch,  oval,  oblong,  or  linear, 
•^5~93  X -^S-'SS  mm.,  with  a  hyaline  crenulate  margin,  and  bear- 
ing 3-7  pairs  of  loculi,  or  2  or  3  antheridia  sometimes  immersed  in 
base  of  the  9  branch  (paroicous)  :  margin  of  $  branch  with 
copious  laciniae,  each  commonly  ending  in  a  single  row  of  cells  ; 
calyptra  much  roughened,  3-4  mm.  high  :  capsule  brown,  oblong- 
■ellipsoidal  ;  elater-bearers  about  the  length  of  the  valves  ; 
■elaters  mostly  250-480  fi  long,  18  /v.  in  greatest  width,  attenuate 
at  the  ends  ;  spores  i  5  //,  nearly  smooth. 

On  decaying  wood  and  moist  banks.  Olema  (1187),  Mendo- 
cino (586,  666),  Eureka  (964)  and  Hay  Fork  (1188).  The  last- 
mentioned  specimen  seems  to  be  strictly  dioicous,  and  a  tendency 
to  restriction  of  the  two  kinds  of  organs  to  separate  individuals  is 
sometimes  to  be  observed  in  our  other  Californian  specimens.  The 
description  of  the  sporogonium  is  drawn  from  no.  944,  Jack,  Leiner 
and  Stizenberger,  Kryptogamen  Badens. 

4.  Riccardia  latifroxs  (Lindb.)  Lindb.  Acta  Soc.  Sci.  Fenn 

10  :  513.  1875. 
AneiLva  latifrons  Lindb.  Not.  pro  Fauna  et  Flora  Fenn.  13  : 
372.  1874. 

Thallus  light  green  or  darkening  on  drying,  translucent  when 
moistened,  palmately  divided  ;  the  ultimate  branches  oblong  or 
linear,  usually  2-4  mm.  long,  .83-1.32  mm.  wide,  commonly 
strongly  emarginate,  slightly  convex  dorsally,  plane  orsubconcave 


74 


Metzgeriaceae 


beneath,  without  suggestion  of  a  costa,  the  unistratose  margin  only 
one  cell  in  width,  very  rarely  gemmiferous  at  apex  ;  surface-cells 
pentagonal,  hexagonal  or  irregularly  oblong,  large,  pellucid,  45— 
118  X  29-50/7.:  monoicous  :  androecium  affixed  to  base  of  9 
branch  or  distant,  oblong,  .66-.85  x  .27— .33  mm.,  somewhat  nar- 
rowed at  base,  with  an  erect  laciniate  margin,  loculi  about  five 
pairs  :  9  branch  with  numerous  small  marginal  laciniae  mostly 
composed  of  a  single  row  of  cells;  calyptra  pyriform-clavate, 
slightly  verrucose  near  apex,  2.5—3.5  n^ni-  high,  .75—1  mm.  broad 
above  :  elaters  1 20-240  n  long,  i  5  //  in  maximum  width  ;  spores 
brownish-green,  minutely  papillate,  1 3-2 1  fjt. 

On  decaying  logs.  Redwood  Canon,  Marin  Co,  (1184); 
Navarro  (Miss  Edith  S.  Byxbee)  and  Mendocino  (592,  721,  676); 
Kneeland  Prairie,  Humboldt  Co.  (1023);  near  Hay  Fork,  Trinity 
Co.  (11 12,  1 1 29);  Sisson,  Siskiyou  Co.  (39).  The  species  was 
collected  in  California  by  Dr.  Bolander,  also,  the  exact  locality 
unknown. 

Lindberg's  original  description  seems  to  call  for  a  larger  plant 
than  ours,  yet  the  Californian  specimens  agree  very  closely  with 
G.  &  R.  Hep.  Eur.  no.  493  and  Sulliv.  Muse.  Allegh.  no.  279^ 
which  were  referred  to  R.  latifrons  by  Lindberg,  and  also  with 
specimens  collected  in  Sweden  by  Arnell. 

The  branching  may  be  subpinnate  below,  but  ultimately  is 
always  pronouncedly  palmate  or  subdichotomous. 

5.  RiccARDiA  PALMATA  (Hcdw.)  Carruth.  Seeman's  Jour.  Bot.  3  : 

302.  1865. 

JiDigcvDiannia  pabnata  Hedw.  Theor.  Gen.  Zj.pL  18.  f.  gj—g^, 
1784. 

Thallus  opaque,  compact,  dark  green  on  drying,  palmately 
branched  or  subdichotomous,  usually  without  trace  of  a  unistratose 
border,  almost  always  bearing  gemmae  at  apices  both  dorsally  and 
ventrally  ;  gemmae  of  two  cells,  formed  endogenously  from  cells 
of  the  surface  ;  surface-cells  small,  quadrate-rhombic  or  oblong- 
pentagonal,  23-59  ^  I5~29/^:  dioicous :  $  thallus  narrow,  bi- 
convex, sometimes  nearly  terete,  the  branches  .18-35  "^^n- 
width,  rounded  at  apex  or  slightly  notched  ;  9  plants  broader  and 
larger,  the  segments  .5-1  mm.  wide:  androecia  oblong  or  linear, 
often  in  pairs,  .5-1.1  x  .25-32  mm.,  with  entire  inflexed  mem- 
branous margins,  sometimes  proliferous  at  apex,  loculi  4-8  pairs 
at  maturity  :   9  branch  subentire,  or  with  a  few  short  and  incon- 


Pellia 


75 


spicuous  marginal  laciniae,  rarely  proliferating,  archegonia  5-18; 
calyptra  strongly  tuberculate  :  spores  small,  almost  smooth,  brown- 
ish-green. 

On  old  logs.  Russian  Gulch,  near  ]\Iendocino  (715)  and  at 
Sisson  (50 — associated  with  R.  latifrons). 

We  have  observed  no  calyptra  or  sporogonium  in  the  Califor- 
nian  plants  and  for  the  description  of  these  parts  have  depended 
upon  authors,  especially  Lindberg  and  Limpricht. 

In  R.  palniata,  nearly  the  entire  surface-layer  of  the  thallus 
sometimes  becomes  resolved  into  gemmae,  after  which  the  plant 
appears  more  translucent  and  as  if  composed  of  larger  cells. 

In  one  case  alone  out  of  many  preparations  from  the  specimens 
cited,  we  have  seen  what  may  have  been  a  monoicous  individual, 
but  here  we  were  unable  to  determine  with  certainty  whether  the 
union  was  really  organic  or  due  simply  to  adhesion. 

METZGERIA  Raddi,  Atti  Soc.  Ital.  Sci.  Mod.  18:  —(34).  1818. 
Mem.  Mat.  e  Fis.  Soc.  Ital.  Sci.  Mod.  18:  45.  1820. 
In  the  manuscript  on  the  Californian  Hepaticae  prepared  by  Dr. 
Watson  and  Mr.  Austin  (see  page  10),  there  is  to  be  found  a  short  de- 
scription oi  Metzgeria  conjugata  Lindb.,  followed  by  the  remark  :  "A 
few  fronds,  insufficient  for  proper  determination,  but  apparently  agreeing 
with  this  description,  were  found  among  the  specimens  of  FruUania 
Grayafia,  var.,  collected  by  Bolander."  A  Metzgeria  finrafa  Xees" 
is  included  among  the  seven  Hepaticae  enumerated  by  SuUivant  in  the 
Report  on  the  Botany  of  Whipple's  Expedition,^  but  whether  this  was 
collected  within  the  limits  of  California  or  not  is  a  matter  of  uncertainty. 
As  we  have  seen  no  California  specimens  of  Metzgeria,  we  will  not  at- 
tempt any  description  of  its  species.  It  is  hoped  that  the  brief  diagnosis 
given  in  the  key  to  the  genera  of  the  Metzgeriaceae  will  suffice  for  the 
recognition  of  the  genus.  At  least  two  species  of  Metzgeria  occur 
farther  to  the  northward  on  the  Pacific  Slope,  and  there  is  little  doubt 
that  the  genus  is  represented  in  California. 

14.  PELLIA  Raddi,  Atti  Soc.  Ital.  Sci.  Mod.  18  :  —(38).    18 18. 
Mem.  Mat.  e  Fis.  So:.  Ital.  Sci.  Mod.  18  :  49.  1820. 

Gametophore  a  thin  or  somewhat  fleshy,  irregularly  dichoto- 
mous,  undulate-margined  thallus  without  ventral  scales,  growing 
in  thick  masses  with  imbricate-ascending  lobes  or  scattered,  bright 


*  Pacific  R.  R.  Sun-ey,  4  :  191.  1857. 


76 


Metzgeriaceae 


green,  darkening  with  age  or  on  drying,  usually  soft  and  flaccid 
when  soaked  out,  furnished  with  a  broad,  for  the  most  part  indis- 
tinctly defined  costa  (9-16  cells  in  thickness),  this  passing  gradu- 
ally into  the  wide  unistratose  somewhat  sinuate-lobed  margin  ;  root- 
hairs  springing  from  the  costa,  brownish,  numerous.  Monoicous 
or  dioicous.  Antheridia  oval,  short-stalked,  immersed,  usually 
singly,  in  loculi  irregularly  distributed  along  the  dorsal  surface  of 
the  costa.  Archegonia  in  groups  of  4-18,  borne  in  a  pocket-like 
cavity  near  the  apex  of  the  costa,  becoming  dorsal  by  the  subse- 
quent growth  of  the  thallus,  covered  more  or  less  by  an  involucre 
opening  in  front.  Involucre  at  maturity  of  sporogonium  a  mere 
flap  or  scale  at  the  posterior  margin  of  the  cavity  or  becoming 
tubular  or  forming  a  short  and  narrow  ring,  the  margin  crenulate 
or  lacerate.  Calyptra  exserted  from  the  involucre  or  included,  its 
walls  composed  of  several  layers  of  cells. 

Sporogonium  consisting  of  a  spherical,  hght  brown  or  yellow- 
ish, 4-valved  capsule  on  a  very  long  delicate  seta ;  capsule-wall  bi- 
stratose,  cells  of  the  outer  layer  with  yellowish-brown  trigones  or 
nodular  thickenings,  those  of  the  inner  layer  with  semiannular  or 
imperfectly  spiral  thickenings  (wanting  or  rudimentary  in  P.  endivi- 
aefolid),  elater-bearers  "  and  elaters  both  present,  the  former  20- 
100  in  number,  persistently  attached  to  the  base  of  the  capsule, 
i-6-spiral,  differing  more  or  less  in  character  from  the  true  elaters  ; 
the  latter  2-4-spiral,  mostly  rather  obtuse,  often,  together  with  the 
spores,  remaining  for  a  time  clinging  to  the  elater-bearers  and  form- 
ing a  globular  entangled  mass  at  the  base  of  the  valves.  Spores 
very  large,  multicellular  at  the  time  of  dehiscence,  oval  or  ellipsoidal, 
punctulate. 

I.  Pellia  Neesiana  (Gottsche)  Limpr.;  Cohn,  Krypt.  Fl.  Schles. 

I  :  329.  1877. 
Pdlia  cpipJiylla,  forma  Neesiana  Gottsche,  Hedwigia,  6  :  69. 
1867. 

Dioicous  :  9  plants  somewhat  sparingly  dichotomous,  with 
oblong  or  linear  branches  ;  $  plants  forming  dense  mats  with  nu- 
merous broad  crenate  overlapping  ascending  segments  :  involucre 
forming  a  short  tube,  1-2  mm.  high,  with  a  lacerate  margin  ;  ca- 
lyptra exserted  1-3  mm.:  capsule  1.5  mm.  in  diameter  on  seta 
3-5  cm.  long  ;  cells  of  inner  layer  of  capsule-wall  with  often  im- 
perfect semiannular  thickenings  ;  contents  of  capsule  in  part  per- 
sisting for  some  time  after  dehiscence  in  an  intertangled  globular 
central  mass  ;  elater-bearers  about  30,  stout,  conical-  or  blunt- 
pointed,  120-270//.  X  18-36//.,  mostly  3- or  4-spiral,  sometimes 
bispiral  or  ar.nulatc  ;  elaters  strongly  contorted,  2-,  or  more  rarcl}', 


Blasia 


77 


3-spiral,  225-400/7.  x  9-12/7;  spores  84-105/^  x  51-69/7,4 
or  5  cells  in  length  and  2  or  3  in  greatest  width. 

On  banks  of  streams,  along  the  North  Fork  of  the  Little  River^ 
Mendocino  Co.  (605,  654,  657). 

Pellia  epipJiylla  (L.)  Corda  and  P.  endiviaefolia  (Dicks.)  Du- 
niort.*  may  both  occur  in  California.  P.  cpipJiylla  differs  from  P. 
Nccsiana  in  being  monoicous  and  in  the  reduction  of  the  involucre 
to  a  scale  or  flap  on  the  posterior  margin  of  the  cavity  from  which 
the  calyptra  arises.  P.  endiviaefolia  \P.  calycina  (Tayl.)  Nees]  is 
dioicous,  but  differs  from  both  the  others  in  the  greater  develop- 
ment of  the  involucre,  which  forms  a  tube  about  4  mm.  high,  from 
which  the  calyptra  scarcely  ever  protrudes,  and  is  markedly  differ- 
ent from  both  in  characters  of  elaters  and  elater-bearers,  the  latter 
being,  according  to  Jack,t  often  as  many  as  100,  much  longer  and 
more  slender  (600-800  a  x  5  fj)  and  bispiral,  while  the  true  elaters 
are  shorter  than  in  the  other  two  species  (150-200  /7),  less  con- 
torted, and  3-  or  4-spiral.  Semiannular  thickenings,  conspicuous 
on  the  inner  wall  of  the  capsule  of  P.  cpipJiylla  and  P.  Necsiaiia^ 
are  wanting  or  obscurely  developed  in  P.  endiviaefolia. 

A  sterile  Pellia  was  collected  in  1894  by  ^Messrs.  M.  S.  Baker 
and  F.  P.  Nutting  in  the  mountains  south  of  Dixey  Valley  in  Las- 
sen Co.,  but  the  species  is  scarcely  determinable.  Dr.  Bolander 
lists  P.  calycina  "  in  his  Catalogue  of  the  plants  growing  in  the 
Vicinity  of  San  Francisco,"  but  we  have  been  able  to  see  no  Pellia 
of  Dr.  Bolander's  collecting. 

15.  BLASL-\  L.  Sp.  PI.  1 1 38.     1753.    Ex  Mich.  Nov.  PI.  Gen. 

14.  1729. 

Gametophore  dark-  or  bluish-green,  becoming  yellowish,  usu- 
ally several  times  dichotomous,  forming  rosettes  or  irregularly 
tangled  tufts,  somewhat  transitional  in  character  between  the  thal- 
loid  and  foliose  types,  5-10  cells  in  thickness  along  the  narrow  or 
expanded  costa,  becoming  gradually  thinner  toward  the  unistratose 
border,  the  latter  sometimes  merely  crenulate  or  sinuate  but  more 
often  running  out  into  rounded,  distant,  or  close  and  incubous, 
leaf-like  lobes  ;  root-hairs  colorless,  springing  from  the  costa,  often 
numerous  toward  the  base  and  fastening  the  plant  closely  to  the 
substratum.     Underleaves  represented  by  small,  oblong,  ovate,  or 

■^Rec.  d'Obs.  Jung  27.  1835. 
t  Flora,  81  :  6,  7.  1895. 


78 


Metzgeriaceae 


heteromorphic,  dentate  scales,  in  an  irregular  row  on  each  side  of 
the  costa,  attached  by  the  posterior  margin  or  near  the  middle 
with  the  margins  free.  On  the  ventral  surface,  also,  are  small, 
hollow,  subspherical  organs,  the  "leaf-auricles,"  usually  two  near 
the  base  of  each  lateral  lobe,  their  cavities  commonly  soon  occu- 
pied by  colonies  of  A^ostoc."^  Marginal  cells  of  the  lateral  lobes 
20-30  n,  oblong,  quadrate,  or  oval,  each  often  with  a  salient  point, 
remaining  cells  of  lobe  30-60  ij.,  rhombic-hexagonal,  surface  cells 
of  the  axial  region  more  elongated.  Gemmae  of  two  kinds  ;  one 
borne  in  dorsal  flask -shaped  receptacles  formed  out  of  the  sub- 
stance of  the  apex  of  the  costa  and  appearing  single,  paired,  or 
in  threes,  according  to  the  development  of  the  branch-system,  fur- 
nished at  maturity  with  a  long  slender  neck  ( 1-2  mm.  x  .  1 7-.  3  mm), 
the  neck-wall  2-4  cells  in  thickness,  the  gemmae  flattened-oval, 
90-135  /^-, '3-5  cells  wide,  2  cells  thick  or  unistratose  at  margin, 
borne  on  slender  hyaline  pedicels  arising  from  the  base  and  sides 
of  the  receptacle,  the  pedicels  accompanied  by  short,  clavate,  uni- 
cellular trichomes  ;  second  form  of  gemmae  on  the  dorsal  surface 
of  the  younger  portions  of  the  plant,  scale-like,  stellate  or  coarsely 
dentate,  of  a  single  layer  of  cells  near  their  margin,  of  two  or  more 
layers  toward  the  middle.  Dioicous.  plants  smaller  ;  anthe- 
ridia  several,  immersed  singly  in  a  row  along  the  dorsal  surface  of 
the  costa.  Archegonia  numerous,  on  dorsal  surface  of  the  costa 
toward  the  apex,  the  single  fertilized  archegonium  becoming,  by 
the  arching  over  of  the  adjacent  parts,  completely  enclosed  within 
a  cylindrical  or  clavate  inflation  of  the  costa,  this  3-4  mm.  in 
length,  directed  forward,  finally  ruptured  at  the  anterior  end  by 
the  elongating  sporogonium,  and  functioning  as  an  involucre. 
Calyptra  light-colored,  membranous.  Capsule  oval,  dingy  brown, 
i-iyi  mm.  long,  dehiscing  by  4  (4-6)  valves,  inner  surface  with 
annular  or  spiral  thickenings,  the  outer  surface  showing  brownish 
nodules  in  the  radial  walls  ;  seta  rather  thick,  i  yl-2  cm.  long. 
Spores  ovoid  or  somewhat  angular,  38-50  fi,  slightly  granulate, 
long  remaining  with  the  elaters  in  a  central  globular  mass.  Elaters 
200-300  fix  10-14  ff.,  with  two  spiral  bands,  these  often  becoming 
paired  by  splitting. 

The  only  known  species  is 

I.  Blasia  pusilla  L.  /.  c. 
Jimgeruiaiuiia  Blasia  Hook.  Brit.  Jung.  //.  82-8^.     18 16. 

*  The  Nostoc  colonies  are  thought  to  live  in  symbiotic  relations  with  the  Blasia, 
perhaps  serving  as  reservoirs  of  moisture  (Goebel,  in  Schenk's  Handbuch  der  Botanik, 
2  :  360.  1882).  The  colony  is  surrounded  ventrally  by  a  single  layer  of  cells  and  is 
penetrated  l)y  a  branching  tube  which  springs  from  the  wall  of  the  cavity. 


FOSSOMBROXIA 


79 


Beside  a  pool,  Blue  Lake,  Humboldt  Co.  (1002).  Our  speci- 
men is  simply  gemmiferous  ;  the  above  description  of  the  sporo- 
gonium  is  drawn  from  a  Swedish  plant  collected  by  Dr.  Arnell. 

16.  FOSSOMBRONIA  Raddi,  Atti  Soc.  Ital.  Sci.  Mod.  18:— 
(29).     18 1 8.    Mem.  Mat.  e  Fis.  Soc.  Ital.  Sci. 
:\Iod.  18  :  40.  1820. 

Gametophore  consisting  of  a  creeping,  simple,  furcate,  or  inno- 
vating stem  with  2-ranked,  obliquely  inserted  succubous  leaves. 
Stem  flattened  above,  closely  attached  to  the  substratum  by  very 
long,  mostly  violet-  or  vinous-purple  root-hairs.  Leaves  assurgent 
or  subvertical,  undulate-crisped,  especially  on  drj-ing,  usually  im- 
bricate, subquadrate,  often  broader  than  long,  the  bases  extending 
far  onto  the  dorsal  side  of  the  stem  and  more  or  less  decurrent, 
margins  in  most  cases  irregularly  sinuate-lobed,  sometimes  toothed, 
rarely  subentire  ;  small  subulate  paraphyllia-like  appendages  often 
occurring,  especially  on  dorsal  surface  of  the  stem  ;  leaf-cells  large, 
thin-walled,  often  in  two  or  more  layers  near  the  base,  cuticle 
smooth.  Monoicous  (commonly  heteroicous,  /.  r.,  paroicous  and 
synoicous),  sometimes  dioicous.  Antheridia  spherical  or  oval,  on 
about  equally  long  stalks,  standing  singly  or  in  groups  of  2—4  on 
the  dorsal  side  of  the  stem  near  the  bases  of  the  leaves.  Arche- 
gonia  similarh'  situated  (the  fertile  one,  however,  always  near  the 
stem-apex),  single  or  in  groups  of  several,  sometimes  closely  asso- 
ciated with  the  antheridia.  The  fertilized  archegonium  becoming 
surrounded  by  a  large  campanulate  or  goblet-shaped  involucre, 
this  commonly  open  or  incised  to  the  base  in  front  (/.  toward 
the  stem-apex),  the  wide  mouth  undulate-lobed  or  dentate.  Invo- 
lucre often  surrounded  by  several  small  subulate  or  linear-lanceo- 
late scales,  these  in  most  cases  finally  adnate  to  the  involucre  and 
forming  slight  wing-like  ridges.     Calyptra  free,  thin. 

Sporogonium  a  spherical  capsule  on  a  short,  rarely  somewhat 
elongated  seta  arising  from  a  subglobose  "foot."  Capsule  brown 
or  when  young  and  liv^ing  nearly  black,  irregularly  dehiscent  or 
imperfectly  4-valved,  the  wall  bistatose,  cells  of  the  inner  layer 
with  often  incomplete  semiannular  thickenings.  Spores  large, 
several  times  broader  than  the  elaters,  round  tetrahedral,  somewhat 
flattened,  the  outer  face  echinate,  verrucose,  reticulate-alveolate,  or 
bearing  free  or  anastomosing  crests,  the  small  inner  faces  much 
less  strongly  roughened,  in  most  cases  simply  punctate,  verrucose- 
papillate,  or  marked  with  short,  low  ridges.  Elaters  usually 
2-  or  3-  (1-4-)  spiral. 


80 


Metzgekiaceae 


I.    FossoMBRONiA  LONGiSETA  Aust.  /.  /.    [as  synonym]  Proc. 
Acad.  Nat  Sci.  Philad.     1869:  228.  1869. 
AiidrocrypJiia  longiseta  Aust.  /.  c. 

Stems  mostly  6-15  mm.  long  and  once  dichotomous,  rather 
stout  (15-20  cells  in  thickness),  commonly  somewhat  tuberously 
thickened  at  apex  and  perennial  through  the  resumption  of  apical 
growth  on  termination  of  the  dry  season,  root-hairs  vinous-purple  : 
leaves  1.5-3  ^^ng,  subquadrate,  assurgent  or  nearly  horizontal, 
more  or  less  imbricate,  irregularly  lobed,  toothed,  or  subentire, 
often  of  2-5  layers  of  cells  toward  the  base  ;  leaf-cells  30-45  fiy^. 
40-60  u.  near  the  margin,  40-60 /.«x  60-1 50/-/  near  the  base  ;  mon- 
oicous  (polyoicous  ?)  :  involucre  usually  large,  1.5-3  high, 
campanulate,  open  to  the  base  on  the  side  toward  stem-apex  or 
often  connate  here  forming  a  complete  cup,  usually  with  several 
subulate  squamules  adnate  to  the  outer  surface,  these  mostly  short 
but  sometimes  reaching  nearly  to  the  lobate-dentate  mouth  :  seta 
finally  8  18  mm.  long;  capsule  1-1.2  mm.  in  diameter,  the  semi- 
annular  thickenings  of  the  inner  layer  of  the  wall  mostly  incom- 
plete :  spores  yellowish-brown,  distinctly  compressed,  38-50  fi  * 
in  maximum  diameter,  strongly  and  somewhat  remotely  cristate, 
the  crests  high,  projecting  3-6  ji  at  the  margin,  20-30  in  number 
in  the  basilar  circumference,  more  or  less  obliquely  ascending, 
slightly  flexuous,  thin,  acute,  usually  undulate-serrulate,  unequal 
in  length,  disappearing  or  sparingly  confluent  at  the  middle  of  the 
face,  forming  there  very  rarely  1—3  fully  closed  meshes  ;  crests 
sometimes  mostly  replaced  by  subacute  or  truncate  spines,  these 
often  numerous  and  crowded,  30-40  in  number  in  the  basilar  cir- 
cumference:  elaters  bispiral,  150—270  ft  in  length. 

On  banks  and  beside  paths.  Clarendon  Heights,  San  Fran- 
cisco ;  Mill  Valley,  Mt.  Tamalpais,  and  Lake  Lagunitas,  Marin 
Co.;  Duncan's  Mills,  Sonoma  Co.;  Ukiah  (743,  785),  Mendocino 
Co.;  Blue  Lake  (998),  Humboldt  Co.;  Lake  Pilarcitos,  San  Mateo 
Co.;  Reche.  Canon,  San  Bernardino  Co.  (Parish,  no.  3614);  Pasa- 
dena (A.  J.  McClatchie);  San  Diego  (Mrs.  Katharine  Brandegee). 
Collected  also  by  Dr.  Bolander ;  and  by  Prof  W.  H.  Brewer  at 
"  Egon  Pass"  (in  herb.  W.  H.  Pearson). 

Exsicc.    Hep.  Bor.-Am.  118  (/./.?). 
Hep.  Am.  i  57. 

The  "type  specimen"  of  Fossombronia  longiseta  appears  not 
to  exist  either  in  that  portion  of  the  Austin  collection  owned  by 


*  Measuring  to  the  extremities  of  the  crests. 


FOSSOMBRONIA 


81 


Mr.  Pearson  or  in  that  in  possession  of  the  Owens  College  at 
Manchester.  The  species  was  distributed  by  Mr.  Austin  a  few 
years  after  its  publication  as  no.  1 18  Hep.  Bor.-Am.,  but  we  have 
been  unable  to  examine  the  spores  in  this,  owing  to  the  imperfect 
and  fragmentary  condition  of  the  only  specimen  we  have  seen. 
Here,  as  after  the  original  description,  Texas  as  well  as  California 
was  cited  as  furnishing  the  species,  but  the  Texas  plant,  according 
to  Lindberg  (who  gave  it  the  name  F.  Texand),  would  appear  to 
be  quite  a  different  thing,  having  areolate  spores  and  being  thus 
allied  to  F.  angidosa.  A  specimen  of  F.  longiseta,  which  may  be 
a  portion  of  the  original  material  studied  by  Mr.  Austin,  is  to  be 
found  in  herb.  Underwood  marked  CaHfornia,  ex  herb.  C.  F. 
Austin,  comm.  O.  D.  Allen."  The  original  description,  it  should 
be  noted,  calls  the  species  dioicoiis,  while  our  specimens  can,  in 
most  cases,  be  shown  to  be  monoicoiis ;  yet  capsule-bearing  plants 
are  sometimes  found  which  exhibit  numerous  abortive  archegonia 
on  the  back  of  the  stem  without  traces  of  antheridia  in  any  part. 
The  two  kinds  of  organs  seem  never  to  be  closely  associated  in 
this  species.  Plants  collected  on  January  7,  1896,  near  Lake 
Lagunitas,  Marin  Co.,  which  we  believe  to  be  of  this  species, 
though  not  spore -bearing,  show  numerous  antheridia  along  the 
back  of  the  stem,  and,  from  the  examination  with  a  pocket  lens  at 
the  time  of  gathering,  the  specimens  were  thought  to  be  purely  $  , 
but  it  w^as  demonstrated  on  dissection  that  each  bore  also  a  number 
of  archegonia  near  the  stem-apex.  The  truth  in  the  matter  seems 
to  be  that  the  plants  are  perennial  in  most  cases  at  least  and  that 
each  plant  has  alternating  periods  of  archegonia-  and  antheridia- 
production  and  that  the  two  successive  periods  of  production  of 
either  organ  may  sometimes  be  quite  widely  separated. 

Fossouibronia  longiseta  is  allied  to  F.  pusilla  (L.)  Dumort.  and 
F.  Wondraczeki  (Corda)  Dumort.  {F.  cristata  Lindb.),  but  is  suf- 
ficiently distinct  from  both  in  the  longer  seta,  the  rather  larger 
leaves  and  leaf-cells,  the  arrangement  of  the  sexual  organs,  prob- 
ably in  the  perennial  habit,  and,  in  most  cases  at  least,  in  the  spore- 
markings.  The  crests  of  the  spores  are  usually  high,  as  in  F. 
pusUla,  but  are  commonly  more  numerous  and  often  show  a  ten- 
dency to  become  broken  up  into  spines,  thus  affording  a  transition 
to  the  purely  echinate  type  of  Fossombronia  spore.    The  spore- 


82 


JUNGERMANNIACEAE 


markings  are  extremely  variable,  and  it  is  possible  that  two  or 
more  species  should  be  recognized  in  the  Californian  specimens, 
but  we  have  been  unable  to  draw  separating  lines  in  any  satisfac- 
tory way.  The  difference  between  a  purely  and  rather  remotely 
cristate  spore  like  that  represented  in  our  figure  1 8  and  one  in  which 
the  crests  are  nearly  all  replaced  by  narrow  spines,  very  numerous 
and  densely  crowded  in  the  basilar  circumference,  like  that  shown 
in  fig.  20,  is  very  striking,  but  forms  that  seem  to  be  intermediate 
exist,  as  is  shown  by  our  other  figures.  It  is  worthy  of  remark 
that  the  spores  (fig.  17)  of  the  specimen  "  ex  herb.  C.  F.  Austin," 
alluded  to  above,  bear  more  spines  than  crests  ;  the  spines  are  here 
somewhat  shorter  than  in  specimens  from  the  southern  part  of  the 
state  [Pasadena  (McClatchie);  San  Bernardino  Co.  (Parish)] .  All 
the  specimens  that  we  have  seen  from  stations  north  of  San  Fran- 
cisco have  spores  of  the  purely  cristate  type,  having,  on  the  whole, 
more  in  common  with  the  spores  of  F.  pusilla  than  with  those  of 
F.  Wondraczeki. 

Plate  99. 

16-20.  Fossombronia  longiseta  Aust.    Spores,  X  3^5  • 

16.  From  Clarendon  Heights,  San  Francisco,  Mar.  28,  1896. 

17.  From  specimen  in  herb.  Underwood  marked  "  California,  ex  herb.  C.  F.  Aus- 
tin, comm.  O.  D.  Allen." 

18.  From  Mill  Valley,  Marin  Co.,  Apr.  4,  1896. 

19.  From  specimen  in  herb.  Underwood,  marked  San  Francisco,  Cal.,  coll.  T. 
S.  Brandegee." 

20.  From  Pasadena  (A.  J.  McClatchie). 

Family  IV.  JUNGERMANNIACEAE. 

Apex  of  the  gametophore  (or  of  some  of  its  branches)  at  length 
directly  transformed  into  an  archegonium.  Sporogonia  always 
terminal  on  the  main  axis  or  on  the  branches.  Gametophore  a 
leaf-bearing  stem.*  Leaves  in  two  lateral  rows,  in  addition  to 
which  there  is  often  a  ventral  row  of  "  underleaves  "  com- 
monly differing  from  the  lateral  in  size  and  form.  Antheridia 
borne  in  the  axils  of  more  or  less  modified  leaves  (''  $  bracts  "). 
Archegonia  usually  surrounded  by  a  special  involucre,  the  **peri- 

In  three  exotic  genera,  there  is  a  prominent  thalloid  or  filamentous  phase,  but 
this,  in  two  of  the  cases,  at  least,  is  doubtless  to  be  homologized  with  the  protonema 
rather  than  with  the  gametophore.  In  all,  the  branches  bearing  the  sexual  organs  are 
foliose. 


JCNGERMANNIACEAE 


83 


anth,"  which,  in  many  cases,  at  least,  represents  a  union  of  the 
leaves  of  the  adjacent  cycle.  The  leaves  9  bracts  ")  and  under- 
leaf     bracteole  ")  subtending  the  perianth  often  somewhat  modified. 

Capsule  quadrivalvate  (the  valves  very  rarely  again  cleft). 
Elaters  always  developed  in  association  with  the  spores. 

More  than  four  fifths  of  all  the  species  of  Hepaticae  of  the 
world,  according  to  Schiffner,  belong  to  the  present  family. 

Kejr  to  the  Genera  of  Junsrermanniaceae.^ 

Underleaves  present  throughout. 

Leaves  complicate-bilobed,  the  ventral  lobe  much  the  smaller. 

Ventral  lobe  usually  inflated,  sac-like,  helmet-shaped,  or  cbovoid-clavate, 

rarely  explanate  ;  underleaves  bifid.  35.  Frullania. 

Ventral  lobe  plane  or  with  revolute  margins,  ovate  to  linear-lanceolate,  some- 
times nearly  separate  from  the  dorsal  ;  underleaves  entire  or  dentate. 

34.  Par  ell  a. 

Leaves  with  2-5  lobes  or  teeth,  or  divided  to  the  base  into  capillary  segments, 
never  acutely  complicate. 
Leaves  bifid  or  bidentate. 

Underleaves  much  smaller  than  the  leaves  and  usually  different  in  form. 
9  branch  a  fleshy  almost  leafless  subterranean  sac  depending  from 
the  ventral  surface  of  the  stem  ;  the  appressed  underleaves  bifid  to 
the  middle  or  below,  with  rather  narrow  sinus  and  nearly  parallel 
lobes,  otherwise  entire.  25.  Geocalyx. 

9  stem  or  branch  comparatively  little  modified,  conspicuously  foliose. 
Root-hairs  in  tufts  at  the  base  of  the  underleaves,  the  latter 
deeply  bifid,  the  somewhat  spreading  segments  again  cleft  or 
externally  unidentate  near  the  base  ;  perianth  more  or  less 
triangular-prismatic,  the  third  angle  always  dorsal. 

23.  Lophocolea. 

Root-hairs  irregularly  scattered;  perianth  triangular-prismatic,  at 
least  when  young,  the  third  angle  always  ventral. 

26.  Cephalozia.-\ 
Root-hairs  irregularly  scattered ;  perianth  ovoid,  obovoid,  or 
cylindrical,  sometimes  slightly  compressed  laterally,  terminal 
on  main  stem  or  leading  branches,  now  and  then  falsely 
dorsal .  2 1 .  Z oph ozia . 

Underleaves  similar  to  the  leaves  both  in  form  and  size. 

30.  Anthelia. 

Leaves  with  3-5  principal  lobes  or  teeth,  these  more  than  one  cell  wide  at 
base. 

Leaves  3-  or  4-lobed  to  about  the  middle  (in  our  species)  ;  the  lobes 
entire,  somewhat  acute,  incurved  ;  underleaves  a  little  smaller  than  the 
leaves,  otherwise  similar ;  perianth  on  short  branch  arising  from  the 
axil  of  an  underleaf.  28.  Lepidozia. 

*  The  present  key  is  based,  for  the  most  part,  upon  the  known  Californian  repre- 
sentatives of  the  various  genera,  and  may  not  always  apply  to  extra-limital  species. 

f  Underleaves  are  rather  uncommon  in  this  genus  outside  of  the  9  branches,  where 
they  are  always  represented. 


84 


J  UNG  ER  M  ANN  I  ACE  A  E 


Leaves  3-  or  4-cleft  to  below  the  middle  ;  the  segments  filiform-acumin- 
ate, entire  or  more  or  less  ciliate-fringed  ;  antheridia  on  the  main  stem 
or  lateral  branches  ;  perianth  really  acrogenous  (/.  e.,  terminal  on  main 
stem  or  principal  branches),  but  finally,  through  innovation,  on  a  short, 
lalsely  lateral  branch,  31.  PtilidiitJii. 

Leaf-lobes  3-5,  obtuse,  acute,  apiculate,  or  spinescently  pointed,  the 
sinuses  not  extending  to  the  middle  ;  underleaves  very  different  from 
the  leaves,  usually  deeply  bifid  and  often  ciliate-fringed ;  perianth 
acrogenous;  plants  usually  large.  21.  Lophozia  Barbatac,  not  yet 
found  in  California). 
Leaves  divided  to  the  base  into  2-4  capillary  segments  ;  underleaves  similar. 

29.  Blcpha  rost,  ma. 
Leaves  entire  or  merely  retuse  at  apex  (sometimes  bidentulate  in  Kaniia'). 

Leaves  incubous  (/.  c,  the  anterior  margin  covering  the  posterior  margin  of 
the  next  leaf  in  front)  ;  underleaves  suborbicular  to  ovate,  emarginate  or 
bifid  ;  9  branch  a  fleshy  almost  leafless  subterranean  sac  depending  from 
the  axil  of  an  underleaf.  27.  Kaiitia. 

Leaves  succubous  (/.  e.,  the  anterior  margin  covered  by  the  posterior  margin 
of  the  next  leaf  in  front). 
Plants  with  creeping,  nearly  or  quite  leafless,  rhizomatous  primary  stems 
and  ascending  foliose  branches  ;  leaf-margin  usually  more  or  less  den- 
tate ;  underleaves  minute,  subulate  and  entire  or  2-  or  3-cleft, 

22.  PlagiocJiila. 

Plants  without  rhizomatous  primary  stems  ;  leaves  without  marginal  teeth 
(but  often  retuse  or  bilobed  at  apex  in  Lophocolea  heterophytla). 
Root-hairs,  when  present,  in  tufts  at  the  base  of  the  underleaves  ; 
leaves  immarginate  ;  capsules  dehiscing  by  straight  valves. 
9  branch  very  short ;  the  bracts  a  single  pair,  with  sometimes 
rudiments  of  a  second,  very  much  smaller  than  the  leaves  ; 
androecium  on  the  main  stem.  24.  Chiloscyphiis. 

9  branch  longer,  bracts  (  9  and  $  )  3-5  pairs  ;  paroicous. 

23.  Lophocolea  (Z.  hcterophylla'). 
Root-hairs  long  and  numerous,  springing  from  nearly  all  parts  of  the 
often  reddish-purple  ventral  surface  of  the  stem  ;  leaves  mostly 
linguiform,  margined  by  a  row  of  enlarged  cells,  trigones  conspic- 
uous ;  capsule  dehiscing  spirally,  the  valves  very  long  and  slender. 

18.  Gyi'othyra. 

Underleaves  wanting  throughout  or  present  only  in  association  with  the  9  bracts.* 
Leaves  complicate-bilobed. 

Ventral  lobe  much  the  smaller ;  root-hairs  springing  from  a  mammilliform 

outpocketing  of  the  ventral  lobe  near  its  base.  33.  Radula, 

Ventral  lobe  the  larger  (the  two  lobes  rarely  equal)  ;  root-hairs  springing  from 
the  stem. 

Perianth  strongly  compressed  dorso-ventrally  (parallel  with  the  plane  of 
the  "frond")  ;  fold  of  the  leaf  usually  acute,  often  alate ;  root-hairs 
commonly  scanty  on  the  foliose  branches.  32.  Scapania. 

*  Chiloscyphns polyanthos  riviilaris  may  be  sought  here,  inasmuch  as  the  underleaves 
are  commonly  destroyed  or  wanting  except  in  the  youngest  parts,  where,  however,  they 
may  always  be  demonstrated. 


Marsupella 


85 


Perianth  not  compressed  dorso-ventrally,  deeply  plicate  at  mouth  ;  leaf 
folded  at  about  a  right  angle,  the  fold  rounded  ;  lobes  acuminate,  ob- 
scurely crenulate  toward  apex  ;  root-hairs  long  and  numerous. 

21.  Lophozia  (Z.  ovata). 
Leaves  with  two  nearly  equal  lobes  or  teeth,  never  acutely  complicate. 

Archegonia  terminal  on  the  main  stem  ;  uppermost  pair  of  9  bracts  more  or 
less  highly  connate,  enclosing  the  delicate  perianth,  to  the  lower  part  of 
which  the  bracts  are  adnate  ;  leaves  typically  transverse. 

1 7 .  Ala  rsti pella . 

Archegonia  usually  terminal  on  short,  ventral  branches  ;  perianth  free  from 
the  bracts,  much  exserted,  triangular-prismatic,  at  least  when  young,  the 
third  angle  always  ventral ;  branches,  typically,  all  of  ventral  origin 

26.  Cephalozia. 

Archegonia  terminal  on  the  main  stem  or  principal  branches  ;  9  t)i'acts  free 
from  each  other  and  from  the  perianth  ;  perianth  oval  to  cylindrical-obovoid, 
never  triangular-prismatic,  commonly  plicate  toward  the  mouth  ;  leaves  suc- 
cubous.  21.  Lophozia. 

Leaves  with  3-5  lobes.  21.  Lophozia. 

I>eaves  entire  (or  in  Plagiochila  often  dentate-margined),  never  bilobed 

Plants  with  creeping,  nearly  or  quite  leafless,  rhizomatous  primary  stems  and 
ascending  foliose  branches  ;  leaf-margins  usually  somewhat  dentate. 

22.  Plagiochila  * 

Plants  without  rhizomatous  primary  stems ;  leaf-margins  entire  or  merely 
repand. 

9  bracts  more  or  less  adnate  to  base  of  perianth.      19.  A'ardia. 

9  bracts  wholly  free  from  perianth.  20.  Jtingerniannia. 

17.  MARSUPELLA  Dumort. Comm.  Bot.  114.  1822. 

SarcoscypJws  Corda  ;  Opiz,  Beitr.  652.  1829.! 
Nardiiis  S.  F.  Gray,  pro  parte  minima,  Nat.  Arr.  Brit.   PI.  I  : 
694.     1 82 1. 

Plants  from  large  and  stout  to  very  small  and  slender,  usually 
densely  caespitose.  Stems  erect  or  ascending,  very  rarely  pros- 
trate, subsimple  or  dichotomously  branched,  commonly  sending  out 
stolons  or  nearly  leafless  flagella  from  near  the  base,  root-hairs  in 
most  species  infrequent  except  at  base  and  on  the  stolons.  Leaves 
patent  or  erecto-patent,  always  bilobed  ;  in  the  typical  species 
transversely  inserted,  disposed  in  opposite  ranks,  canaKculate-con- 
cave  or  subcomplicate-carinate,  the  plant  thus  appearing  com- 
pressed dorso-ventrally ;  in  M.  Bolandeid  often  obliquely  inserted 
and  distinctly  succubous.  Underleaves  wanting.  Usually  dioicous 
or  paroicous,  rarely  autoicous  or  synoicous.    9  bracts  commonly 

*  Structures  representing  underleaves  are  commonly  present  in  the  North  American 
Plagiochilae  but  they  are  often  so  minute  and  fugacious  that  they  may  sometimes  escape 
observation. 

I  See  footnote,  p.  33. 


86 


JUXGERMANNIACEAE 


2—4  pairs,  larger  and  less  deeply  lobed  than  the  cauline  leaves,  the 
inmost  pair  more  or  less  highly  connate  at  base  and  fused  with  the 
lower  part  of  the  perianth  and  the  hollowed-out  upper  part  of  the 
stem  to  form  a  somewhat  goblet-shaped  involucre.  Perianth  deli- 
cate' included  in  the  involucre,  irregularly  4-6-lobed  after  the  ex- 
sertion  of  the  capsule.  Calyptra  free  or,  toward  the  base,  adnate  to 
the  perigynial  tube.  Capsule  globose  or  slightly  elongated,  the 
valves  of  2  or  3  layers  of  cells,  the  walls  of  which  exhibit  numerous 
brown  nodular  or  imperfectly  semiannular  thickenings.  Elaters 
bispiral. 

Key  to  tlie  Species. 

Stems  2-6  mm.  long  ;  leaves  in  sterile  and  lower  part  of  J  plants  somewhat  obliquely 
inserted  and  succubous  ;  median  leaf-cells  16-36  /z ;  antheridia  single  or  in  pairs  ;  9 
bracts  nearly  always  margined  with  radially  elongated  cells.         i.  M.  Bolandei'i. 

Stems  5-80  mm.  long  ;  leaves  always  transversely  inserted  and  canaliculate-concave  or 
subcomplicate  ;  median  leaf-cells  - 16-28  u  ;  antheridia  in  groups  of  2-6;  9  bracts 
ver}'  rarely  submarginate.  2.  M.  eniarginata. 

I.  Marsupella  BoLANDERi  (Aust.)  Underw.  Zoe,  i:  365.  1890. 

SarcoscypJms  Bolanderi  Aust.  Bull.  Torr.  Bot.  Club,  3  :  9. 
1872. 

Nardia  Bolanderi  Aust.;  Underw.  Bull.  111.  State  Lab.  Nat. 
Hist.  2  :  113.  1884. 

Densely  and  intricately  caespitose,  lurid  green  or  dusky  red, 
often  blackening  :  stems  prostrate  or  ascending  at  apex,  rarely  sub- 
erect,  2-6  mm.  long,  .1-22  mm.  in  diameter,  simple,  sparingly 
branched  below,  or  innovating  from  the  axils  of  the  9  bracts,  fur- 
nished to  the  apex  in  most  cases  (especially  in  the  9  plants)  with 
somewhat  copious  colorless  root-hairs  :  leaves  close  or  distant, 
variable  in  size  and  position,  ovate-orbicular,  usually  broader  than 
long,  .14-.36  mm.  in  length,  .22-55  width,  subtransverse 

and  erecto-patent  in  ^  plants  and  toward  the  perianth  in  the 
9 ,  otherwise  mostly  patulous-subvertical  (turned  toward  the  dor- 
sal aspect  of  the  stem)  and  succubous,  sometimes  obscurely 
marginate,  concave,  occasionally  subcomplicate,  rarely  almost 
plane,  emarginate-bilobed  y^—Yi  their  length,  the  sinus  acute 
or  somewhat  obtuse,  the  lobes  triangular-ovate,  mostly  obtuse  ; 
leaf-cells  rather  thin-walled,  with  small  or  obsolete  trigones,  the 
marginal  subquadrate,  16-25  ft^  sometimes  becoming  a  little 
elongated  radially,  the  median  mostly  rounded-hexagonal,  16- 
36  /i,  cytoplasm  in  exposed  parts  containing  a  diffused  reddish- 
brown  pigment  :  dioicous  :  antheridia  single  or  more  commonly 
in  pairs,  on  stalks  a  little  shorter  than  their  long  diameter,  the  $ 
bracts  often  distinctly  bordered  with  subquadrate  cells  a  little  elon- 


Marsupella 


87 


gated  radially,  the  dorsal  margins  usually  subrevolute  :  9  bracts 
3  or  4  pairs,  rather  abruptly  2-6  times  larger  than  the  leaves, 
nearly  always  distinctly  marginate  with  radially  elongated  often 
more  highly  colored  cells  25-45  //  in  greatest  diameter,  median 
cells  oval -elliptical,  25-50/7.;  inmost  pair  of  bracts  finally  the 
largest,  more  or  less  highly  connate,  forming  an  exserted  peri- 
anth-like, cylindrical  or  goblet-shaped  involucre  from  a  some- 
what obconical  base,  emarginate-bilobed  (rarely  trilobed) 
their  length  ;  outer  bracts  transversely  inserted,  erecto-patent, 
bilobed  ^5-^4  their  length,  otherwise  entire  or  slightly  repand,  the 
lobes  obtuse  :  perianth  subpyriform,  a  little  shorter  than  the  invo- 
lucre, free  in  the  upper  third  or  half,  with  4—6  connivent  lobes  at 
the  mouth  :  calyptra  mostly  of  two  layers  of  cells  :  capsule  dark 
brown,  globose,  .4-45  in  diameter,  the  walls  of  the  cells  with  very 
numerous  nodular  thickenings  ;  seta  2-4  mm.  long. 

On  or  about  rocks  or  on  exposed  banks,  sometimes  under 
chaparral  ;  hills  and  ridges  of  the  Coast  Range  ]Mountains.  San 
Francisco  :  ^Mission  Dolores  (Bolander),  Clarendon  Heights 
(Howe);  Mt.  Tamalpais  and  near  Lake  Lagunitas,  Alarin  Co.;  Pieta, 
Mendocino  Co.  (804) ;  near  Lake  San  Andreas,  San  Mateo  Co. 
Often  associated  with  CcpJialozia  divaricata,  to  which,  sometimes, 
the  more  slender  forms  bear  a  superficial  resemblance.  Xear  Lake 
San  Andreas,  it  grows  in  company  with  Scapania  nemorosa,  Xardia 
crejiidata,  CepJialozia  Tiirneri,  and  Marsupella  cmarginata.  It  is 
always  readily  distinguishable  from  the  smaller  conditions  of  the 
last-named  species  by  its  darker  color,  the  often  distinctly  succu- 
bous  leaves,  the  larger  and  thinner-walled  leaf-cells,  the  nearly  al- 
ways marginate  $  bracts,  the  more  exserted  perianth-like  involucre 
formed  by  the  inmost  pair  of  bracts,  the  fewer  antheridia  to  each 
perigonial  leaf,  the  smaller,  darker,  and  more  nearly  spherical  cap- 
sule, etc. 

All  of  the  minute  European  Marstipcllac  described  more  re- 
cently by  Limpricht  and  by  Spruce  seem  to  be  abundantly  distinct 
from  Marsupella  Bolanderi.  The  larger  and  otherwise  different 
Marsupella  sphacelata  erythrorhiza  (Limpr.)  [under  Sarcoscyphus 
in  Cohn,  Krypt.-Fl.  Schles.  i:  248,  431.  1877]  is  possibly  its 
nearest  ally,  judging  from  specimens  kindly  communicated  by 
Professor  Limpricht. 

The  often  obliquely  inserted,  subv'ertical,  and  distinctly  succu- 
bous  character  of  the  leaves  in  Marsupella  Bolanderi  serves  to 


88 


JUNGERMAXXIACEAE 


bridge  over  one  of  the  more  important  gaps  between  the  genera 
Marsiipella  and  Nardia  as  defined  by  Spruce  and  accepted  by  most 
of  the  modern  writers  on  hepaticology. 

Plate  ioi.    ]Marsupella  Bolaxderi. 

1.  Plants,  natural  size. 

2,  3.  Apical  portions  of  plants,  showing  innovations  from  axils  of  9  bracts,  X  12. 
4.  Apical  portion  of  a  sterile  plant,  lateral  view,  showing  succubous  leaves,  X  23. 
5-7.  Dorsal,  lateral,  and  ventral  views  of  apex  of  different  9  plants,  showing 

bracts,  etc.,  X  23. 

8.  $  plant,  X  23. 

9.  $  bract  and  antheridia,  X  23. 

10.  A  sterile  (9?)  plant,  X  23. 

11.  Cauline  leaf,  X  23. 

12.  Cauline  leaves,  outer  (convex)  face,  X  23. 

13.  Cauline  leaves,  inner  (concave)  face,  X  23. 

14.  Involucre,  opened  to  show  perianth  and  sterile  archegonia,  calyptra  removed, 
X23- 

15.  Margin  of  cauline  leaf,  X  225. 

16.  Part  of  9  bract,  from  one  of  the  pair  next  subtending  the  perianth-like  in- 
volucre, showing  the  radially  elongated  marginal  cells,  X  225. 

Figures  10  and  15  from  the  original  specimen  ex  herb.  Austin  ;  2,  8,  and  9,  Mt. 
Tamalpais ;  3,  12,  13,  near  Lake  San  Andreas,  San  Mateo  Co.;  4-7,  ii,  14,  and  16, 
near  Lake  Lagunitas,  Marin  Co. 

2.  Marsupella  EMARGiXATA  (Ehrh.)  Dumort.  Comm.  Bot.  114. 

1822. 

Jtingcnnannia  cinargi)iataY\\x\\.^€\'ix.       80.     1788.  Hook. 

Brit.  Jung.  pi.  2j.    18  16. 

Sarcoscj'phos  Ehrharti  Cor dd.\  Opiz,  Beitr.  652.  1829."^ 
NardiiLS  cinarginatus  S.  F.  Gray,  Nat.  Arr.  Brit.  PI.  I  :  694. 

1821. 

Variable  in  size  and  habit,  usually  stout,  loosely  or  somewhat 
densely  caespitose,  green  or  often  yellowish-  or  reddish-brown, 
sometimes  dark  purple  :  stems  commonly  erect,  .5-8  cm.  (mostly 
1-4  cm.)  high,  simple  or  dichotomously  branched,  usually  desti- 
tute of  root-hairs  except  near  base  and  on  the  numerous  basal  sto- 
lons :  leaves  rather  firm  and  rigid,  transversely  inserted,  canalicu- 
late-concave or  subcomplicate,  patent  or  erecto-patent,  mostly 
approximate,  often  imbricate  toward  the  apex,  quadrate-orbicular, 
.3-.9  mm.  in  diameter,  from  a  broad  base  embracing  one  half  the 
stem,  emarginate  or  bilobed,  the  obtuse  sinus  rarely  descending  be- 
low one  fourth  the  length  of  the  leaf,  the  lobes  broadly  ovate,  ob- 


*  See  footnote,  p.  33. 


Gyrothyra 


89 


tuse  or  rarely  acute  ;  leaf-cells  oval  or  round-hexagonal,  the 
median  16-28  //.,  the  marginal  usually  a  little  smaller,  all  with 
large  trigones  :  dioicous  :  antheridia  in  groups  of  2-6,  on  stalks 
■equaling  their  vertical  diameter  or  sometimes  twice  as  long,  the 
bracts  often  with  slightly  revolute  margins  :  9  bracts  2-4  pairs, 
gradually  2  or  3  times  larger  than  the  leaves,  very  rarely  sub- 
marginate,  the  inmost  pair  highly  connate,  exserted  or  immersed, 
one  or  two  of  the  exterior  pairs  sometimes  connate  at  base,  lobes 
similar  to  those  of  leaves,  median  cells  25-34  n  :  perianth  some- 
what shorter  than  bracts,  4-6-lobed  :  capsule  yellowish-brown, 
oblong-globose,  .54-./  mm.  in  length  ;  seta  2-10  mm.  long. 

On  a  moist  roadside  bank  near  Lake  San  Andreas,  San  Mateo 
Co.,  Apr.  20,  1895 — in  company  with  the  foregoing  species. 

18.  GYROTHYRA  M.  A.  Howe,  Bull.  Torr.  Bot.  Club,  24:  201. 

1897. 

Stem  creeping,  foliose,  subsimple  or  somewhat  sparingly 
branching,  radiculose.  Leaves  succubous,  entire,  alternate  ;  under- 
leaves  present,  free,  bifid,  segments  narrowly  lanceolate  or  subu- 
late ;  walls  of  the  leaf-cells  with  triangular  thickenings  at  the 
angles.  Antheridia  short-stalked,  in  the  axils  of  smaller  saccate 
leaves,  forming  short  median  or,  at  first,  terminal  spikes.  C  bracts 
2—4  pairs  (commonly  3  pairs).  Perianth  terminal,  confluent  for 
half  its  length  or  more  with  the  bases  of  the  bracts,  the  greater  part 
of  the  calyptra,  and  the  tissues  of  the  stem,  to  form  a  thick -walled 
tube  (perigynium),  with  a  small  bulbous  or  saccate  base  ;  peri- 
gynium  erect  or  ascending,  making,  at  maturity,  nearly  a  right 
angle  with  the  stem.  Capsule  cylindrical,  long-exserted,  dehiscing 
spirally  by  four  very  long  and  slender  valves  ;  capsule-valves  of 
two  layers  of  cells,  the  walls  wholly  destitute  of  spiral,  semian- 
nular,  or  other  local  thickenings.  Elaters  free,  bispiral,  acute  or 
bluntly  pointed  ;  spores  minutely  papillate.  Livolucellum  "  of 
the  sporogonium  foot  well  developed. 

I.  Gyrothyra  Uxderwoodiaxa  M.  A.  Howe,  /.  c.  202.  pi.  J02, 

Plants  rather  large,  1-2  cm.  long,  2-4  mm.  wide,  mostly  in 
compact  light  green  tufts  :  stems  creeping,  thick,  often  shghtly 
flattened  dorso-ventrally,  .5-65  mm.  in  diameter,  about  15  cells 
wide  in  cross  section,  very  densely  radiculose,  slightly  ascending  at 
apex,  subsimple  or  with  a  few  irregularly  disposed  lateral  branches, 
in  female  plant  innovating  from  near  base  of  perigynium  ;  root-hairs 
long,  nearly  colorless  or  of  a  dilute  yellowish-brown  hue,  some- 


90 


JUNGERMANNIACEAE 


times  tinged  proximally  with  purple,  springing  in  older  parts  of 
the  stem  from  oblong  or  linear  dark -purple  callosities,  made  up 
of  the  closely  coherent  root-hair  bases  and  of  other  ventrally 
elongated  cells  ;  leaves  obliquely  inserted,  linguiform  or  oval, 
succubous,  rather  close,  translucent,  scarcely  decurrent  dorsally, 
often  crowded  and  suberect  at  stem-apex,  marginate,  1.7-4  mm.  x 
1.4-2  mm.,  commonly  concave  below,  apex  decurved  ;  cells  of 
the  margin  quadrate  or  oblong,  equaling  in  size  the  adjacent  or 
twice  as  large ;  other  leaf-cells  mostly  quite  regularly  pentagonal 
or  hexagonal,  25-70/^  in  diameter,  oblong  and  larger  towards  the 
base ;  all  with  conspicuous  trigones  :  underleaves  free,  often  wine- 
colored,  .6—1  mm.  long,  bifid  their  length  into  narrowly 
lanceolate  or  subulate  segments,  usually  running  out  into  a  single 
series  of  cells  at  apex,  concealed  by  the  dense  mat  of  root-hairs 
except  in  the  younger  portions  of  the  stem :  perigynium  tubular, 
I- 1  y2  mm.  in  diameter,  and,  with  the  free  portion  of  the  perianth, 
3-4  mm.  long,  erect  or  ascending,  nearly  at  right  angles  with  the 
stem,  tinged  with  purple  ventrally,  bulbous  or  saccate  at  base  ;  wall 
of  perigynium-tube  5-20  cells  in  thickness :  $  bracts  2-4  pairs 
(commonly  3  pairs),  entire  or  repand,  similar  in  form  to  the  cauline 
leaves,  margins  approximate  at  base  dorsally,  distant  ventrally  ; 
uppermost  pair  inserted  at  about  middle  of  perianth-tube  or,  more 
rarely,  at  two  thirds  its  height,  erect,  apex  and  dorsal  margins 
narrowly  reflexed  and  exposing  the  perianth,  or  closely  appressed 
and  wholly  concealing  it ;  next  lower  pair  usually  inserted  at  about 
one  third  height  of  perianth-tube,  more  broadly  reflexed  ;  the  one 
or  two  basal  pairs  but  slightly  attached  to  perigynium ;  bracteoles 
inconspicuous,  sometimes  subentire  and  slightly  adherent  to 
base  of  bracts  ;  bulbus  of  perigynium  without  radicles,  but  a 
dense  tuft  of  root-hairs  springs  from  the  stem  just  back  of  the 
bulbus  and  long  root-hairs  come  from  the  cells  of  the  involucral 
leaves  near  their  bases  :  perianth  free  for  ^3-^  its  length,  free 
portion  nearly  echlorophyllose,  subtubular,  somewhat  inflated  be- 
low, contracted  and  lax  above,  crenulate  at  mouth,  3-5  cells  thick 
at  juncture  with  perigynium-tube,  2  cells  thick  at  mouth  :  calyptra 
fleshy,  upper  third  or  fourth  free  at  maturity,  3-6  cells  in  thick- 
ness :  dioicous  :  archegonia  several,  the  unfertilized  raised  on  the 
base  of  the  free  portion  of  calyptra  :  male  plants  more  slender ; 
antheridia  (1-6)  in  the  axils  of  smaller  saccate  leaves,  forming 
spikes  of  3-6  pairs  of  leaves  decreasing  in  size  upwards,  appressed, 
apices  patent  or  recurved,  or,  in  uppermost  pair,  erect ;  antheridia 
ellipsoidal  or  pyriform,  .15X.24  mm.,  on  pedicels  as  long; 
slender  stems  (male  ?)  occasionally  gemmiferous  at  apex,  gemmae 
unicellular,  10-24  //  in  diameter:  capsule  long-cylindrical  ;  valves 
very  slender,  3.3-6  mm.  x.13-.  17  mm.,  widely  spreading  when 


Gyrothyra 


91 


dry,  attached  spiro-radially  to  a  basal  disc  composed  of  large 
hyaline  cells,  flexuous,  contorted,  or  spiral,  when  moistened — 
always  with  a  spiral  twist  at  the  apex  ;  foot  of  sporogonium  form- 
ing a  more  or  less  goblet-shaped  "  involucellum  "  ;  seta  iyi-2  cm. 
long ;  elaters  bispiral,  very  rarely  trispiral,  acute  or  subobtuse, 
210-420  fxx  12-15  ti]  spores  about  12  a,  minutely  papillate. 
Exsicc.  Hep.  Am.  184. 

Near  Eureka,  Humboldt  Co.,  June,  1896  (no.  1026) ;  collected 
also  by  Prof.  John  Macoun  (herb.  Underwood),  on  earth  in  a 
brook,  Burrard  Inlet,  British  Columbia,  April  6,  1889,  and  on  rocks, 
British  Columbia,  April  29,  1889;  and  by  A.  A.  and  E.  Gertrude 
Heller,  near  Montesano,  Chehalis  Co.,  Washington  (June  7,  1898, 
no.  4289). 

The  leaves  of  the  British  Columbia  and  Washington  plants 
stand  with  their  margins  more  often  erect  than  in  the  Californian 
specimens,  upon  which  our  description  and  figures  are  based.  In 
the  sterile  condition,  Gyrothyra  somewhat  resembles  the  larger 
forms  of  Nardia  scalaris — also  collected  by  Macoun  on  Vancouver 
Island  (Can.  Hep.  80) — but  can  readily  be  distinguished  by  the 
margined,  linguiform,  more  translucent  leaves,  and  by  the  bifid 
underleaves. 

The  9  bracts,  though  more  or  less  apparently  paired,  are  in  a 
strict  sense  alternate  like  the  cauline  leaves,  and  a  single  unpaired 
leaf  is  sometimes  found  to  occur  inside  the  pair  we  have  described 
as  the  uppermost. 

It  should  be  remarked  that  the  actual  dehiscence  of  the  capsules 
has  not  been  observed,  but  the  extremely  long  valves,  which  on 
being  soaked  out  take  easily  a  position  strongly  suggestive  of  the 
paring  of  an  apple,  the  spiro-radial  attachment  to  the  basal  disc, 
the  never  failing  spiral  twist  of  the  valve-apex,  and  the  spiral  lines 
feadily  discernible  on  the  surface  of  the  embryo  capsule  (fig.  10) 
make,  in  our  judgment,  the  induction  that  the  dehiscence  is  spiral 
so  safe  and  certain  that  v/e  have  felt  no  hesitation  in  so  describing 
it  and  basing  the  generic  name  upon  this  character.  The  absence 
of  thickenings  in  the  walls  of  the  cells  of  the  capsule  valves  is 
noteworthy.  Schiffner  states'^  of  all  the  Jungcrinaiiniaceac  akrogy- 
nae :  "  Die  reife  Kapsel  besitzt  eine  aus  2  bis  mehr  Zellschichten 


*  Engler  and  Prantl,  Nat.  Pflanzenfam.       :  71.     1 893. 


92 


JUXGERMAXNIACEAE 


bestehende  Wand,  deren  Innenschicht  in  ihren  Zellen  stets  Ver- 
dickungsleisten  enthalt."  In  Gyi'otJiyra,  the  transverse  walls  of 
these  cells  usually  appear  a  trifle  thicker  than  the  longitudinal,  but 
the  walls  are  otherwise  wholly  without  traces  of  local  thickening. 

In  respect  to  structure  of  the  sporogonial  envelopes,  GyrotJiyra 
is  one  of  several  interesting  transitions  from  the  ordinary  Junger- 
mannia  type  to  the  various  pouch -bearing  genera.  Considered 
from  this  point  of  view  and  from  certain  other  gametophytic  char- 
acters, its  nearest  affinities  are  undoubtedly  to  be  found  in  that 
section  of  Nardia  represented  by  Nardia  Jiaematosticta  (Nees) 
Lindb.,  of  Europe.  In  manner  of  dehiscence  of  capsule  it  recalls 
the  marsupiiferous  genus  Kantia  ;  but  the  valves  of  GyrotJiyra  are 
much  loncrer  and  their  cell -walls  lack  the  local  thickening,  while, 
of  course,  no  generic  comparison  of  the  two  is  necessary  so  far 
as  the  gametophore  is  concerned. 

Plates  102  and  103. 

G  Y  R  OTH  Y  R  A  UnDERWOODI  AN  A. 

1.  Entire  9  plant,  X  5- 

2.  Cauline  leaves,  X  iS. 

3.  Marginal  and  adjacent  leaf- cells,  X  225. 

4.  Transverse  section  through  marginal  portion  of  leaf,  X  216. 

5.  Underl eaves,  X  24. 

6.  Transverse  sections  of  stem,  X  22,  showing  ventral  callosity  from  which  the 
root-hairs  arise. 

7.  Antheridium,  X  40- 

8.  Median  sagittal  section  of  perigynium  and  adjacent  portions  of  stem,  showing 
embryo  sporogonium  with  capsule,  seta,  foot,  and  "  involucellum,"  also  unfertilized 
archegonia,  perianth,  insertion  of  9  bracts,  the  root-hair  callosity,  etc.,  X  23  (slightly 
schematized).  The  free  part  of  the  perianth  as  drawn  here  and  in  the  next  is  propor- 
tionally rather  too  short  and  not  sufficiently  inflated  below. 

9.  Sagittal  section  of  mature  perigynium  from  which  the  seta  has  been  detached, 
showing  fully  developed  calyptra  and  the  unfertilized  archegonia  raised  upon  the  base 
of  its  free  portion,  X  20  (slightly  schematized). 

10.  Surface  view  of  embryo  capsule,  exhibiting  the  spiral  lines,  which  presumably 
bound  the  valves,  X  SO- 
IL Valves  of  capsules,  showing  position  taken  by  them  when  moistened,  X  ^2. 

12.  Apex  of  a  single  valve,  X  ^2. 

13.  Base  of  dehisced  capsule  from  above,  showing  spiro-radial  insertion  of  valves, 

X  36.  • 

14.  Cells  of  inner  surface  of  capsule  valve,  X  ^S*^- 

15.  Elaters  and  spores,  X  137- 


Nardia 


93 


19.  NARDIA  S.  F.  Gray  [as  Nardius}  p.  p.  Xat.  Arr.  Brit.  PL 

I  :  694.     1 82 1. 

Mesophylla  Dumort.  Comm.  Bot.  112.  1822. 

Aliailaria  Corda  ;  Opiz,  Beitr.  652.  1829."^ 

Plants  medium-sized  or  small.  Stems  creeping  or  ascending, 
rarely  suberect,  simple  or  with  a  few  latero-ventral  branches  :  root- 
hairs  usually  long  and  numerous,  often  reddish.  Leaves  alternate, 
obliquely  inserted  and  succubous  or  nearly  transverse  toward  the 
perianth,  often  vertical-connivent,  subreniform  to  orbicular-ovate, 
entire  or  merely  repand,  rarely  retuse  at  apex,  sometimes  margin- 
ate.  Underleaves  present,  lanceolate  or  subulate,  or  (in  our  species) 
wanting.  Dioicous  or  paroicous.  Androecium  in  dioicous  species 
terminal  or  median,  $  bracts  saccate,  otherwise  similar  to  the 
leaves,  antheridia  mostly  in  groups  of  2—4.  Archegonia  terminal 
on  the  main  stem.  5  bracts  2-5  pairs  :  in  the  subgenus  Etniardia, 
the  superior  opposite,  connate  in  pairs,  fused  with  the  base  of  the 
perianth  and  the  hollowed-out  stem-apex  to  form  a  perigynial  tube, 
this  sometimes  ventrally  bulbous-thickened,  perianth  small,  rather 
delicate,  included  ;  in  the  subgenus  Eucalyx,  perianth  firm,  much 
exserted,  ovoid  to  obovoid-prismatic,  4-5-carinate  and  abruptly 
contracted  to  a  short  subtubular  mucro  at  the  mouth  or  carinae 
wanting  and  the  apex  conical  and  plicate,  the  superior  bracts  more 
or  less  adnate  to  perianth  (or — more  accurately,  perhaps — borne  on 
the  excavated  stem-apex).  Calyptra  free,  surrounded  at  base  by 
the  unfertilized  archegonia.  Capsule  globose-oval,  dehiscing  by 
straight  rigid  valves  ;  cells  bounding  the  inner  surface  of  the  valves 
with  numerous  semiannular  thickenings,  those  of  the  outer  surface 
with  columnar  or  nodular  thickenings  ;  seta  rather  short  or  moder- 
ately long.     Elaters  bispiral. 

Both  of  the  species  described  below  belong  to  the  subgenus 
Encalyx,  which  passes  gradually  into  the  genus  Jiingerinannia. 
Nardia  crenulata,  in  which  the  9  bracts  are  often  only  very  slightly 
adherent  to  the  base  of  the  perianth,  stands  on  the  border  line  be- 
tween Nardia  and  Jungennannia  and  would  be  about  equally  well 
at  home  in  either  group.  Nardia  scalaris  (Schrad.)  S.  F.  Gray,  of 
the  subgenus  Eiinardia,  has  been  collected  by  Professor  Macoun 
on  Vancouver  Island  and  may  be  expected  to  occur  in  California. 
This  may  be  readily  distinguished  from  either  of  the  following 
species  by  the  presence  of  lanceolate  or  subulate  underleaves  and 
by  the  conspicuous  trigones  of  the  leaf-cells  ;  the  perianth  is  in- 
cluded within  the  involucre  formed  by  the  bracts  and  bracteole. 


*  See  footnote,  p.  33. 


94 


JUXGERMANNIACEAE 


Key  to  the  Species. 

Stems  3-8  mm.  long,  .15-. 35  mm.  in  diameter,  root-hairs  colorless  or  yellowish  ;  leaves 
.4-1.3  mm.  long,  usually  broader  than  long,  margined  by  enlarged  cells  or  immar- 
ginate,  slightly  or  not  at  all  decurrent,  median  and  superior  leaf-cells  25-40  fi  ;  9 
bracts  more  or  less  adnate  to  perianth  at  its  extreme  base.  I.  N'.  crenulata. 

Stems  10-20  mm.  long,  .27-. 64  mm.  in  diameter,  root-hairs  reddish-purple  (occasionally 
decolorate ) ;  leaves  .9-1.7  mm.  long,  usually  longer  than  broad,  mostly  patent-hori- 
zontal and  but  slightly  overlapping,  obscurely  if  at  all  margined,  decurrent,  median 
and  superior  leaf-cells  32-60  ,u  ;  the  inner  9  bracts  adnate  to  perianth  for  two  thirds 
its  length.  2.  A^.  obovata. 

I.  Nardia  crenulata  (Sm.)  Lindb.  Act.  Soc.  Sci.  Fenn.  10  :  529. 

1875. 

Jimgerinannia  cre}iuIataSm,Y.r\g.^ot.  pi.  i^6j.  1805.  Hook. 
Brit.  Jung.       jy.     18 16. 

JiLiigermannia  gracillima  Sm.  Eng.  Bot.  //.  22j8.     181 1. 

Jiingermannia  crenulata  gracillima  Hook.  Brit.  Jung.  pi.  jy. 
1816. 

Jiingermannia  Genthiana  Hiiben.  Hepaticol.  Germ.  107.  1834. 

Nardia  gracillima  Lindb.  Act.  Soc.  Sci.  Fenn.  10  :  530.  1875. 

Nardia  crenulata  gracillima  Lindb.  Muse.  Scand.  8.  1879. 

Jungermannia  rubra  Gottsche ;  Undervv.  Bot.  Gaz.  13:  113. 
//.  4.  1888.  Bolander,  Cal.  Med.  Gaz.  1870:  184  (40)  (name 
only).  1870. 

More  or  less  red  or  brownish-red,  very  rarely  green  through- 
out, gregarious  or  forming  thin  compact  mats  :  stems  rather  slen- 
der (.15-35  mm.  in  diameter),  3-8  mm.  long,  prostrate,  ascend- 
ing at  apex,  simple  or  often  sending  out  flagelliform  latero-ventral 
branches ;  root-hairs  numerous,  moderately  long,  colorless  or 
slightly  yellow  :  leaves  subcartilaginous,  obliquely  attached,  slightly 
or  not  at  all  decurrent ;  those  of  the  fertile  stems  usually  crowded, 
subvertical,  concave  or  nearly  plane,  broadly  rounded-ovate  or 
semiorbicular,  .4-1.3  mm.  long,  .54-1.4  wide,  entire  or  repand,  in 
most  cases  distinctly  margined  by  a  single  series  of  enlarged  quad- 
rate thick-walled  cells,  leaves  of  the  sterile  stems  and  flagella  com- 
monly distant,  smaller,  longer  than  broad,  oval  or  elliptical,  patent, 
often  not  at  all  or  obscurely  marginate  ;  leaf-cells  with  walls  weakly 
thickened  at  the  angles,  the  superior  and  median  25-40//,  the  mar- 
ginal often  2-3  times  as  large  (in  surface  area)  as  the  next  adja- 
cent ;  cuticle  irregular  and  minutely  hyaline-punctate  :  underleaves 
wanting  or  very  rarely  present  in  association  with  the  9  bracts  : 
dioicous  :  androecium  terminal,  $  bracts  3-10  pairs,  concave,  mar- 
gined, suberect,  imbricate  ;  antheridia  usually  in  pairs,  globose, 


Nardia 


95 


,16- 2  mm.  in  diameter,  short-stalked,  accompanied  by  a  few  short 
paraphyses  :  9  bracts  gradually  larger  than  the  leaves,  more  re- 
pand  and  commonly  more  conspicuously  marginate,  crowded  and 
suberect  or  rather  loosely  disposed  and  spreading,  the  uppermost 
one  or  two  more  or  less  adnate  to  the  base  of  the  perianth  or  to 
the  perigynial  tube  formed  by  the  hoUowed-out  stem-apex,  the 
very  rarely  occurring  bracteole  small,  obovate,  semiorbicular,  or 
linguiform  :  perianth  usually  tinged  with  red  or  purple,  nearly  im- 
mersed or  exserted  ^-J^  its  length,  subcomplanate-ovoid  to  nar- 
rowly obovoid  or  prismatic,  1.5-2  mm.  long,  .8-1.3  mm.  in  greatest 
width,  nearly  always,  especially  when  young,  distinctly  quad- 
rangular (rarely  pentagonal),  the  angles  often  incrassate  (of  two 
layers  of  cells),  the  perianth  wall  otherwise  unistratose  except  in 
the  basal  third  or  fourth,  the  mouth  at  first  abruptly  contracted 
into  a  very  short  subtubulose  mucro,  rather  obscurely  ciliolate- 
■denticulate,  at  length  lacerate  :  calyptra  often  reddish  or  purple, 
unistratose  toward  the  apex,  otherwise  bistratose :  capsule  dark 
brown,  ovoid,  .7-. 8  mm.  in  greatest  diameter,  the  wall  consisting 
of  two  layers  of  cells;  seta  4-10  mm.  long;  spores  brown,  13- 
16  a,  very  minutely  granulate;  elaters  brown,  bispiral,  attenuate 
at  the  extremities,  contorted,  80-120 /y.  long,  8-13/^  in  maximum 
width. 

Exsicc.  Hep.  Am.  200. 

On  moist,  usually  exposed  banks  in  the  Coast  Range  Moun- 
tains, especially  near  the  sea.  Mendocino  (Dr.  H.  N.  Bolander)  ; 
Santa  Cruz  (W.  G.  Farlow,  May,  1885),  Santa  Cruz  Mountains 
(L.  Underwood,  August,  1888);  near  Lake  San  Andreas,  San 
Mateo  Co.;  Bolinas,  Marin  Co.  (W.  A.  Setchell) ;  near  Cazadero, 
Sonoma  Co.;  IMendocino  (582,  591,  620,  682) ;  Eureka  (902,  924). 

The  Californian  forms  mostly  agree  with  Gottsche's  Jiuiger- 
inannia  rubra,  founded  on  a  specimen  collected  on  "  metamorphic 
sandstone,  quite  near  the  coast"  at  Mendocino  by  Dr.  Bolander, 
but  Jiingcrmannia  rubra  seems  to  us  not  to  differ  in  any  essential 
structural  characters  from  the  European  conditions  of  Nardia  cren- 
ulata  which  have  at  times  been  known  as  Jtingermajinia  gracilliina 
and  J.  Gcnthiana.  Leaves  conspicuously  margined  by  cells  2  or  3 
times  the  size  of  the  adjacent  occur  in  the  Californian  specimens, 
though  rarely.  In  general,  the  leaves  here  are  only  submarginate. 
But  both  Californian  and  European  specimens  show  such  extremes 
of  variation  in  this  particular  in  different  parts  of  a  single  tuft  or 
even  in  different  parts  of  a  single  plant  that  this  character  cannot 


96 


JUNGERMANNIACEAE 


be  given  much  weight.  The  leaves  are  sometimes  shghtly  more 
decurrent  than  we  have  observed  them  to  be  in  European  forms. 
The  wall  of  the  perianth  is  often  two  cells  thick  at  the  angles  but 
we  have  never  found  the  angles  externally  papillose-serrate. 

2.  Nardia  obovata  (Nees)  Lindb.  Bot.  Notis.  1872  :  167.  1872. 
Jungermannia  obovata  Nees,  Naturgesch.  Eur.  Leberm.  i  :  332. 

1833- 

SoutJibya  obovata  Lindb.  Hartm.  Skand.  Fl.  2:  130.  1871. 
[ed.  10.] 

Bright  green  to  brownish-red,  loosely  caespitose  :  stems  rather 
stout  (.27— .64  mm.  in  diameter),  1—2  cm.  long,  prostrate  or  as- 
cending, simple  or  innovating  above  in  fertile  plants,  very  rarely 
otherwise  branched  ;  root-hairs  abundant,  long  and  fasciculate,, 
reddish-purple  or  occasionally  decolorate :  leaves  rather  firm  in 
texture,  distant  or  slightly  overlapping,  the  lower  obliquely  at- 
tached (at  an  angle  of  about  45°),  patent-horizontal,  the  upper 
more  transversely  inserted,  divergent  and  subsquarrose  or  some- 
what erect,  all  orbicular-ovate  from  a  narrowed,  concave,  sheath- 
ing base,  plane,  slightly  concave  or  convex  in  the  upper  half,  .9— 
1.7  mm  X  .7—1-5  mm,  dorsally  decurrent,  entire  or  subrepand, 
the  apex  very  obtusely  or  subtruncately  rounded,  rarely  retuse  ; 
leaf-cells  thin-walled,  usually  with  small  trigones,  the  sparse  or 
moderately  abundant  chlorophyll-bodies  finally  disposed  near  the 
vertical  (lateral)  walls,  median  and  superior  cells  32-60/;?,  the 
marginal  often  subquadrate  or  a  little  elongated  radially  and  form- 
ing an  obscure  border,  cuticle  delicately  striatulate-papillate  : 
underleaves  none  :  paroicous  :  antheridia  in  pairs,  short-stalked, 
accompanied  by  a  few  short  paraphyses,  borne  in  the  saccate  bases 
of  the  lower  9  bracts  and  adjacent  leaves  :  9  bracts  4-6,  larger 
than  the  leaves,  obovate,  erecto-patent,  commonly  reflexed  at  the 
apices,  the  superior  2-4  very  highly  adnate  to  the  perianth,  but 
not  connate  :  perianth  elongate-obovoid,  free  from  the  bracts  only 
in  the  terminal  third,  the  free  portion  scarcely  projecting  beyond 
the  uppermost  bracts  or  included,  thin,  composed  of  oblong, 
nearly  hyaline  cells,  plicate  toward  the  subentire  or  lobate  mouth. 

On  the  banks  of  a  shaded  slough.  Blue  Lake,  Humboldt  Co., 
June  3,  1896  (1002). 

Nardia  obovata  has  not  before  been  recorded,  we  believe,  for 
America,  and  as  the  above  specimen  is  wholly  sterile,  its  reference 
to  this  species  is  open  to  some  question.  With  the  exception  of 
the  color  of  the  root-hairs — which  show  only  very  rarely  a  tinge 


JUNGERMANNIA 


97 


of  reddish-purple — the  agreement  with  European  specimens  of 
Nardia  obovata  is,  however,  so  close  that  at  least  until  more  com-^ 
plete  material  is  obtained  we  can  do  no  better  than  to  identify  it 
with  this  species.  In  European  specimens  the  root-hairs  are  some- 
times colorless  in  certain  parts  of  the  stem,  though  w^e  have  never 
observed  in  these  any  such  widely  extended  decoloration  as  in  our 
H  umboldt  County  plant.  In  regard  to  characters  of  leaves  and  leaf- 
cells,  the  correspondence  between  the  Californian  specimen  and  no. 
Zjoa  of  Jack,  Leiner,  and  Stizenberger's  Kryptogamen  Badens  is 
especially  striking.  The  larger,  more  ovate,  more  distant,  more 
patent-horizontal,  and  occasionally  subsquarrose  leaves  stand  in 
the  w^ay  of  referring  the  plant  to  Nardia  liyalina  (Lyell)  Carr., 
while  the  firmer,  subtruncately  rounded,  occasionally  emarginate- 
retuse,  decurrent  leaves,  and  the  more  distinct  trigones  seem  to 
preclude  the  possibility  of  its  being  any  condition  of  Jiuigermannia 
idparia  Tayl. 

20.    JUNGERMANNIA*  L././.  (?)Sp.  PI.  1 13  I.     1753.  Ex 
Rupp.  Fl.  Jen.  345.     171 8.    Dumort. /. max. 
Rec.  d'  Obs.  Jung.  16.  1835. 

Jungcrinaiinia  §  Aplasia  Dumort.  /.  /.  7nax.    Syll.  Jung.  47. 

1831. 

Aplozia  Dumort././.  max.  Hep.  Eur.  55.  1874. 
LiocJdaena  Nees,  G.  L.  &  N.  Syn.  Hep.  150.  1845. 
Jiingermannia,  subgenus  Eiijiuigcrmannia  §1,  Spruce,  Trans, 
and  Proc.  Bot.  Soc.  Edinb.  15:  508.  1885. 

*  Professor  Schiffner  (Eng.  &  Prantl,  Nat.  Pflanzenf.  :  82.  1893)  has  discarded 
Jtmgermannia  altogether  as  a  generic  name  inasmuch  as  the  Jungerjuannia  of  modern 
authors  has  come  to  bear  Httle  or  no  resemblance  to  the  group  recognized  under  this 
name  by  Linnaeus.  Jungerniatinia  lanceolata  [Lioc/ilacna  lanceolata  Nees)  is  the 
only  representative  of  the  Linnaean  Jungcrmannia  species  of  1753  to  be  found  in  any 
recent  conception  of  the  genus  and  this  same  Jungermannia  lanceolata  of  Linnaeus — if 
we  are  to  interpret  his  species  by  the  synonymy  alone — was,  according  to  Lindberg,  a 
mixture  of  several  species  none  of  which  was  the  Jiingcrviannia  lanceolata  as  known 
today,  even  though  one  or  two  of  them  did  belong  to  the  group  Aplozia.''''  Dumortier 
in  1831  divided  what  was  left  of  Jungermannia,  after  several  excisions  had  been  made, 
into  nine  sections,  two  of  which  were  Aplozia  and  Lophozia.  In  1835  he  elevated 
Lophozia  to  generic  rank,  retaining  the  name  Jungermannia  for  his  former  section 
Aplozia.  None  of  the  Linnaean  specific  names  appeared  under  his  Lophozia  and  none 
of  the  species  recognized  in  the  first  edition  of  the  Species  Plantarum  would  have  nat- 
urally found  a  place  in  this  group.    The  Linnaean  name  Jtingermamtia  lanceolata. 


98 


JUXGERMANNIACEAE 


Plants  mostly  of  medium  size.  Stems  creeping  or  ascending, 
less  commonly  erect,  simple  or  bearing  a  few  latero-ventral 
branches,  sometimes  with  one  or  two  innovations  from  below  the 
perianth ;  root-hairs  usually  long  and  moderately  abundant. 
Leaves  alternate,  obliquely,  or  almost  longitudinally  inserted  and 
succubous,  or  transverse  throughout,  mostly  patent-horizontal, 
rarely  semivertical,  oblong-elliptical,  oblong-ovate,  orbicular,  or  sub- 
reniform,  never  lobed,  the  margins  very  entire.  Underleaves  mostly 
wanting  or  small.  Paroicous,  dioicous,  or  autoicous.  Androecium 
terminal  or  median,  $  bracts  ventricose,  otherwise  similar  to  the 
leaves,  antheridia  mostly  1-3,  short-stalked,  with  or  without 
paraphyses.  Archegonia  terminal  on  the  main  stem.  9  bracts 
scarcely  different  from  the  leaves,  distinct  from  each  other  and  free 
from  the  perianth.  Perianth  ovoid,  cylindrical,  or  clavate,  plicate 
in  the  conical  upper  part  or  very  abruptly  contracted  to  a  short 
tubular  mouth,  now  and  then  somewhat  compressed  either  laterally 
or  dorso-ventrally.  Calyptra  free,  surrounded  at  base  by  the  sterile 
archegonia.  Capsule  oval  or  globose,  dehiscing  by  straight  valves  ; 
cells  bounding  the  inner  surface  of  the  valves  provided  with  semi- 
annular  thickenings  ;  seta  long  or  somewhat  short.    Elaters  bispiral. 

Key  to  the  Species. 

Leaves  decurrent. 

Leaves  very  obliquely  or  almost  longitudinally  attached,  mostly  longer  than  broad. 

I.  J.  Bo/anden' . 

Leaves  transversely  attached,  clasping,  mostly  broader  than  long.  3.  J.  Danicola. 
Leaves  not  decurrent,  obliquely  attached,  elliptical-oblong.  2.  J.  pui/iila. 

which  has  now  come  to  have  a  definite  meaning,  however  much  it  may  have  covered  at 
the  start,  was  given  a  place  under  Dumortier's  section  Aplozia,  which  four  years  later 
became  his  genus  Jungerfuaiinia.  It  therefore  seems  to  us  that,  in  the  spirit  at  least  of 
the  unwritten  '*  law  of  residues,"  the  Aplozia  group  is  the  logical  heir  to  the  generic  name 
Jungerniannia.  In  1874,  however,  Dumortier  made  an  unfortunate  and  (according  to 
our  notions)  an  unjustifiable  transposition  of  names  by  which  Jungcrtnannia  was  applied 
to  his  LopJwzia  of  1835  while  his  Jioigcrniaunia  of  1835  was  rechristened  Aplozia  ;  in 
this  he  has  been  recently  followed  by  Massalongo,  Evans,  Heeg,  Loitlesberger,  and  pos- 
sibly others. 

Relying  upon  a  statement  by  Professor  J.  B  Wilbrand  (Flora,  9'^:  518.  1826) 
to  the  effect  that  "Ludovicus  Jungerman"  never  wrote  his  name  Jungerma;;;/  but  always 
Jungermaw,  Lindberg,  Spruce,  Arnell,  Underwood,  Schiffner,  Pearson,  and  many 
others  have  amended  the  original  spelling  of  the  generic  name  by  omitting  one  of  the 
n's.  But  M.  Emile  Bescherelle  states  (Journal  de  Botanique,  7  :  191.  1893)  that  he 
has  been  informed  by  other  botanists,  among  them  Herr  Stephani,  that  Jungermann  wrote 
his  name  with  a  single  ;/  or  with  the  two  ;;'s.  Under  these  circumstances,  we  agree 
with  M.  Bescherelle  that  it  is  better  not  to  interfere  with  the  spelling  adopted  by  Rup- 
pius,  Micheli,  and  Linnaeus. 


JUNGERMANNIA 


99 


I.  JuxGERMAXXiA  BoLAXDERi  Gottsclic  ;  Uiiderw.  Cot.  Gaz.  13: 

113. //.J.  1888. 

Pallescent,  caespitose :  stems  2-30111.  long,  .3-.  3  5  mm.  in 
diameter,  prostrate  or  ascending  (?),  simple  or  with  infrequent  lat- 
eral branches,  sometimes  innovating  below  the  first  pair  of  9 
bracts  ;  root-hairs  nearly  obsolete,  colorless  or  very  dilutely  yel- 
low :  leaves  hyaline,  rather  soft,  often  flexuous  or  undulate,  ap- 
proximate or  slightly  succubous-imbricate  or  on  the  sterile  shoots 
somewhat  distant,  very  obliquely  or  almost  longitudinally  affixed 
to  the  stem,  patent-horizontal,  orbicular-ovate,  .8-1.5  t^^^^-  long, 
,7-1.8  mm.  wide,  usually  longer  than  broad,  entire  or  lightly  re- 
pand,  somewhat  concave  dorsally  with  the  dorsal  margin  more  or 
less  inflexed,  those  of  the  sterile  stems  nearly  plane,  all,  especially 
the  latter,  strongly  decurrent ;  superior  leaf-cells  25-40 the 
marginal  not  differentiated,  the  basal  45-96  fi,  the  thin  walls 
scarcely  or  not  at  all  thickened  at  the  angles,  cuticle  smooth  or 
slightly  roughened  :  dioicous  (?)  :  9  bracts  one  or  two  pairs, 
wholly  free  from  the  perianth,  the  uppermost  twice  as  large  as  the 
leaves,  more  concave,  and  more  transv^ersely  inserted,  otherwise 
similar :  perianth  cylindrical-obovoid,  abruptly  contracted  when 
young  to  a  small  subentire  or  slightly  denticulate  subrostellate 
mouth,  exhibiting  finally  3-5  obtuse  irregular  folds  in  the  superior 
half :  antheridia  and  sporogonia  unknown. 

In  ditches,  Mt.  Dana,  altitude  about  3100  meters  (Dr.  H.  N. 
Bolander,  September,  1866). 

Jiingcrinannia  Bolandcri  is  evidently  allied  to  Jiuigennaniiia 
riparia  Tayl.  and  J.  pumila  With.,  yet  is  sufficiently  distinct  from 
either  in  the  nearly  rootless  stems,  the  softer,  strongly  decurrent, 
almost  longitudinally  affixed  leaves,  and  in  the  larger  leaf-cells,  of 
which  the  marginal  show  no  (or  extremely  slight)  tendency  to  be 
smaller  than  their  neighbors  and  quadrate.  The  leaves  are  usually 
broadest  below  the  middle,  while  in  Jungermannia  pinnila  they  are 
usually  broadest  at  just  about  the  middle.  Of  perianths  we  have 
seen  but  two  and  these  were  apparently  enclosing  unfertilized 
archegonia  and  were  thus  probably  not  fully  and  perfectly  devel- 
oped. The  larger  of  these  was  1.4  mm.  long  and  .8  mm.  in 
greatest  width,  which  was  about  the  size  of  those  seen  by  Dr. 
Gottsche,  judging  from  his  figures  and  his  scale  of  magnification. 
The  mouth  in  this  was  smaller,  more  nearly  entire,  and  more  ab- 
ruptly contracted  than  in  either  Jungermannia  riparia  or  /.  pmnila 
— was,  in  fact,  almost  liochlaenoid.    Our  description  of  the  peri- 


100 


JUNGERMAXNIACEAE 


anth  has  been  drawn  in  part  from  Gottsche's  figures.  Sterile 
plants  bear  some  resemblance  to  Nardia  obovata  but  differ  in  the 
almost  entire  absence  of  root-hairs,  the  softer,  more  decurrent, 
and  much  more  longitudinally  affixed  leaves, 

A  type  duplicate  of  Jiingcrnianiiia  Bolandcri  from  the  hand  of 
Dr.  Gottsche,  together  with  his  original  pencil  sketches  of  the 
species,  is  preserved  in  the  herbarium  of  Harvard  University, 
which  is  true  also  of  Dr.  Gottsche's  Jiingerinannia  Daiiicola,  J. 
Mulleri  Danaciisis,  and  J.  rubra.  We  have  made  an  effort  to  see 
the  remainder  of  Dr.  Gottsche's  material  of  this  and  of  his  other 
Californian  species,  but  without  success.  Dr.  Gottsche's  herbarium 
is  in  the  possession  of  the  Royal  Botanical  Museum  at  Berlin,  but 
Dr.  Hennings,  in  response  to  our  inquiries,  wTites  that  the  Gottsche 
collection  is  only  partly  arranged  and  that  the  Californian  speci- 
mens desired  cannot  thus  far  be  found. 

2.  JuNGERMANNiA  PUMiLA  With.  An*.  Brit.  PL  3  :  866.  //.  i8.  f.  /. 

1796.  [ed.  3.]* 

Dark  green,  sometimes  brownish  or  blackening,  forming  thin, 
loosely  or  closely  interwoven  mats  :  stems  creeping,  commonly 
ascending  at  apex,  simple  or  sparingly  branched,  rhizomatous  from 
the  ventral  surface  or  now  and  then  innovating  from  the  axils  of 
the  bracts  ;  root-hairs  usually  long  and  numerous,  colorless  or 
yellowish-brown  :  the  lower  leaves  loosely  disposed,  scarcely  con- 
tiguous, obliquely  affixed  (line  of  attachment  making  an  angle  of 
about  45°  with  stem),  patent-horizontal,  less  commonly  semivertical, 
the  upper  closer,  more  transversely  attached  and  more  erect,  all 
elliptical-oblong,  .36-9  x  .27-.8  mm.,  obtuse,  somewhat  concave 
or  nearly  plane,  entire  or  slightly  repand,  not  decurrent  ;  leaf-cells 
transparent,  thin-walled,  scarcely  or  not  at  all  thickened  at  the 
angles,  the  median  1 8-3  5  fi,  only  a  little  enlarged  toward  the  base, 
the  marginal  slightly  smaller  and  subquadrate  ;  cuticle  finely  striatu- 
late  or  punctulate  :  underleaves  none  :  paroicous :  antheridia 
spherical,  short-stalked,  occurring  singly  in  the  axils  of  2-4  pairs 
of  saccate,  clasping,  suberect,  more  or  less  enlarged  leaves,  these 
functioning  also  as  $  bracts  :  perianth  narrowly  cylindrical-obo- 
void,  1.2-2.5  X  .36-1  mm.,  somewhat  plicate  in  the  terminal  half 
or  third  and  rather  gradually  narrowed  to  a  small  slightly  hyaline- 
denticulate  mouth. 

On  moist  rocks,  stones  in  brooks,  etc. 


*  We  have  been  unable  to  see  the  third  edition  of  Withering's  work. 


JUXGERMAXNIA 


101 


The  occurrence  in  California  of  this  widely  distributed  species 
cannot  be  considered  as  demonstrated  beyond  the  possibility  of  a 
doubt.  It  is  inserted  hereon  the  strength  of  the  existence  in  Pro- 
fessor Underwood's  collection  of  a  specimen  from  the  Roy  Her- 
barium labeled  Bolander,  Sept.,  1866,"  without  indication  of 
locality.  From  such  an  inscription  it  is  natural  to  infer  that  Bo- 
lander  was  the  collector.  Dr.  Bolander  was  upon  "Mt.  Dana  early 
in  September,  1866,  and  according  to  his  journal,  for  an  extract 
from  which  we  are  indebted  to  Dr.  W.  L.  Jepson,  seems  to  have 
spent  the  remainder  of  the  month  in  San  Francisco.  Another 
specimen,  however,  from  the  Roy  Herbarium,  so  identical  with  the 
former  even  in  its  algal  associates  as  to  make  it  practically  certain 
that  the  two  grew  in  the  same  tuft,  is  labeled  Near  McLeod's 
Lake,  B.  C,  J.  Macoun,  26  June.  1875."  A  similar  plant,  though 
apparently  dioicous  and  having  larger,  more  ovate  leaves,  has  been 
collected  since  that  date  by  Professor  Macoun  in  several  localities 
in  British  Columbia. 

3.  JuxGERMAXXiA  Daxicola  Gottsche ;  Underw.  Bot.  Gaz.  13 : 

113.  //.J.  1888. 

Fuscous-brown,  caespitose  :  stems  prostrate-entangled,  6-10 
mm.  long,  .18-.36  mrri.  (in  attenuate  conditions  only  .08-.  i  mm.) 
in  diameter,  simple,  sparingly  dichotomous,  or  emitting  a  few  lateral 
branches,  sometimes  innovating  from  near  the  base  of  perianth  ; 
root-hairs  very  long,  colorless  or  tawny,  commonly  wanting  to- 
ward the  stem-apex  and  on  the  more  slender  stems  and  innovations, 
otherwise  moderately  abundant  :  leaves  somewhat  distant  or  con- 
tiguous, slightly  overlapping  and  a  little  larger  toward  apex  of  stem, 
patent  or  patent-horizontal,  transversely  inserted,  concave,  embrac- 
ing more  than  one  half  the  stem,  decurrent,  entire,  those  of  the 
fertile  stems  broadly  orbicular  to  subreniform,  .5—9  mm.  long, 
.65-1.2  mm.  wide,  those  of  the  attenuate  sterile  stems  smaller,  or- 
bicular or  broadly  ovate  ;  superior  and  median  leaf-cells  24-50  ri, 
the  walls  with  small  trigonal  thickenings  at  the  angles,  chloroplasts 
and  oil  bodies  rather  few,  the  marginal  cells  slightly  smaller  but 
scarcely  differentiated  in  form  or  color,  cuticle  nearly  smooth  or 
sometimes  very  minutely  hyaline-papillate  :  underl eaves  none  : 
monoicous  (paroicous  ?):  antheridia  in  pairs,  short-stalked,  in  the 
axils  of  hardly  modified  stem-leaves  somewhat  removed  from  the 
9  bracts,  which  are  similar  to  the  leaves  though  more  erect :  peri- 
anth finally  obovoid,  exserted,  irregularly  plicate  toward  the  some- 


102 


JUNGERMANXIACEAE 


what  abruptly  contracted,  at  first  crenulate-denticulate  mouth  : 
sporogonium  unknown. 

In  ditches,  Mt.  Dana,  alt.  about  3100  m.  (Dr.  Henry  N.  Bolan- 
der,  September,  1866). 

Jungerniannia  Danicola  is  very  closely  allied  to  the  Jiingcr- 
mannia  ainplexicaiilis  Dumort.  (y.  tersa  Nees)  of  Europe.  It  dif- 
fers, however,  in  several  more  or  less  significant  particulars  and  had 
best  be  considered  distinct,  at  least  until  better  known.  We  have 
been  unable  to  find  either  perianth  or  antheridia  in  that  portion  of 
the  type  material  preserved  in  the  Gray  Herbarium,  and  have 
drawn  our  brief  description  of  these  parts  from  Dr.  Gottsche's 
figures.  What  we  have  seen  of  the  original  material  consists 
mostly  of  slender  and  distant-leaved  sterile  shoots  ;  these,  judging 
from  the  stouter  stems  which  sometimes  occur  and  which  Dr. 
Gottsche  sketched,  probably  do  not  fairly  represent  the  species. 
From  Jungerinannia  ainplexicaiilis,  the  Mt.  Dana  plant  seems  to 
differ  in  the  dark  brown  color,  the  smaller  size,  the  apparently 
prostrate  (possibly  ascending)  stems,  the  smaller  rather  more  dis- 
tant, more  translucent  leaves,  and  the  less  differentiated  marginal 
cells.  The  two  antheridia  figured  by  Dr.  Gottsche  occupy  the  axil 
of  the  fourth  leaf  below  the  perianth,  counting  on  one  side  only. 
Jungerinannia  aniplexicaiilis  [J.  tersa)  is  described  by  Nees,  Lim- 
pricht,  and  Stephani  as  dioicous,  but  Lindberg  (Kongl.  Sv.  Vet. 
Akad.  Handl.  23'^:  39.  1889)  states  that  it  is  paroicous.  As  rep- 
resented in  Jack,  Leiner,  and  Stizenberger,  Kryptogamen  Badens, 
no.  873  ^  and  c,  the  species  is  certainly  paroicous,  it  being  not 
difficult  to  demonstrate  collapsed  antheridia  with  their  still  per- 
sistent stalks,  intermingled  with  a  few  paraphyses  of  various  forms, 
in  the  slightly  saccate  bases  of  the  9  bracts.  In  Gottsche  and 
Rabenhorst,  Hep.  Eur.  nos.  359  and  511,  we  find  paraphyses  in 
the  axils  of  the  9  bracts,  but  detect  no  unquestionable  antheridial 
remains.  The  occurrence  of  antheridia  in  the  position  figured  by 
Gottsche  in  Jungerniannia  Danicola  suggests  the  possibility  that 
they  may  occur  also  nearer  the  archegonia. 

The  perianth  of  Jungerniannia  aniplexicaulis  is  when  young 
rather  abruptly  contracted  at  the  apex  to  form  a  small  subtubulose 
mouth.  Dr.  Gottsche's  figure  of  a  very  young  perianth  of  /.  Dani- 
cola shows  no  appreciable  contraction  but  it  would  appear  that  this 


LOPHOZIA 


103 


surrounds  immature  archegonia,  when  the  contraction  would  natur- 
ally be  less  than  at  a  little  later  period  in  the  development  of  the 
perianth  ;  however,  we  suspect  that  there  may  be  some  difference 
in  this  regard  between  Jiuigcnnannia  Danicola  and  J.  ample xicaidis. 

From  Jungcnnannia  sphacrocarpa  Hook,  (including  J.  liirida 
Dumort.  and  /.  nana  Nees),  Jungcrniannia  Danicola  appears  to  be 
distinct  in  the  transversely  inserted,  proportionally  broader,  more 
concave,  more  clasping,  and  more  spreading  leaves. 

21.  LOPHOZIil  Dumort.       /.  max.  Rec.   d'Obs.  Jung.  17. 
1835.    Schiffn.;  Engler  &  Prantl,  Nat.  Pflanzenfam.  i'^ : 
84-  1893. 
Gymnocolca  Dumort.  p.  p.,  I.  c. 
DiplopJiylliun  Dumort.  /.      /.  c,  15. 

Jungennannia  %  LopJiozia  Dumort././.  max.    Syll.  Jung.  53. 

1831. 

Jiingcrmannia  Dumort.  Hep.  Eur.  68.  1874.  And  of  nearly 
all  authors. 

Plants  small  to  very  robust.  Stems  creeping  or  ascending, 
simple,  dichotomous,  or  with  a  few  latero-ventral  branches,  often 
innovating  from  near  the  base  of  the  perianth  ;  root-hairs  usually 
numerous.  Leaves  alternate,  succubous,  or  transversely  inserted, 
with  2-5  (commonly  acute)  teeth  or  lobes,  plane  or  more  often 
dorsally  concave,  sometimes  complicate-bilobed  but  never  acutely 
carinate  ;  walls  of  the  leaf-cells  thin  or  with  trigones  at  the  angles, 
never  pitted  or  nodulose.  Underleaves  wanting  or  when  present 
mostly  small,  lanceolate-subulate  and  entire,  or  bifid,  or  ciliate- 
fringed.  Androecium  terminal  or  median  ;  $  bracts  sometimes 
with  an  additional  tooth  or  lobe  on  the  dorsal  margin  near  the 
ventricose  base,  antheridia  1-9  (commonly  1-3),  mostly  short- 
stalked,  with  or  without  paraphyses.  ilrchegonia  terminal  on  the 
main  stem.  9  bracts  distinct,  free,  usually  somewhat  different 
from  the  leaves,  often  with  more  lobes,  and  sometimes  dentate- 
margined.  Perianth  oval,  cylindrical,  or  cylindric-obovoid,  pli- 
cate above  the  middle  or  sometimes  only  obscurely  at  the  very 
mouth,  the  apex  obtuse,  or  more  commonly  conical,  or  in  some 
species  abruptly  contracted  to  a  small  tubulose  mucro  at  the  mouth. 
Calyptra  free.  Capsule  subglobose  to  elongate-ovoid,  dehiscing 
by  four  straight  rigid  valves,  these  usually  composed  of  three  lay- 
ers of  cells,  the  inner  with  semiannular  thickenings.  Elaters  bi- 
spiral. 


104 


JUNGERMANNIACEAE 


Key  to  the  Species. 

Underleaves  present  (usually  inconspicuous).  3.  Z.  heterocolpa. 

Underleaves  wanting  (unless  in  association  with  9  bracts). 

Leaves  bilobed  to  below  the  middle,  complicate,  transversely  inserted  ;  the  dorsal 
lobe  lanceolate,  suberect  or  ascending  ;  the  ventral  lobe  ovate  or  oblong-ovate, 
patent-horizontal.  5.  L.  ovata. 

Lobing  or  cleavage  of  the  leaves  not  extending  below  the  middle. 

Leaves  unequally  2-5-lobed  '^-'X  their  length,  the  lobes  acutely  and  in  most 
cases  spinescently  pointed,  the  margins  more  or  less  spinulose-dentate  ; 
median  leaf-cells  30-60 /i.  i.  L.  incisa. 

Leaves  2-  (rarely  3-)  lobed  i-f  their  length,  close  or  approximate,  the  lobes 
acute,  occasionally  apiculate,  rarely  subobtuse,  entire,  the  sinus  broad  ;  me- 
dian leaf-cells  24-32 11.  2.  L.  z-cntricosa. 
Leaves  2-lobed  \-\  their  length,  usually  distant  or  merely  approximate,  the 
lobes  obtuse,  entire,  the  sinus  rather  narrow.                   4.  L.  i)ifiata. 

I.   LoPHOZiA  INCISA  (Schrad.)  Dumort.  Rec.  d'Obs.  Jung.  17. 

1835- 

Jitngcnnannia  incisa  Schrad.  S}'st.  Samml.  Krypt.  Gew.  2  :  5. 
1797.  Schrad.  Jour.  Bot.  1801  :  67.  1803.  Hook.  Brit.  Jung. 
pi.  10.     1 8 16. 

Light  green  or  darkening,  densely  and  intricately  caespitose  or 
widely  depressed-gregarious  :  stems  very  short  and  thick  (4-6  mm. 
X  .27-1.1  mm.),  ascending,  simple, 'once  furcate,  or  sending  out  one 
or  two  lateral  branches  toward  the  apex  ;  root-hair.3  numerous,  long, 
colorless  :  leaves  usually  close,  commonly  crowded  at  stem-apex, 
plicate-crispate,  succulent,  almost  transversely  inserted  or  some- 
what oblique  and  succubous,  subcomplicate,  erecto-patent,  sub- 
quadrate  or  ovate-oblong,  .7-1.8  mm.  x  .5-1.7  mm.,  deeply  {Yi  — 
Yz)  and  unequally  2-5-lobed,  the  lobes  triangular-ovate, acutely  and 
in  most  cases  spinescently  pointed,  their  margins  usually  reflexed 
and,  especially  in  the  larger  ventral  lobes,  more  or  less  spinulose- 
dentate,  sinuses  acute  or  rounded  ;  leaf-cells  quadrate-hexagonal, 
becoming  oblong  toward  the  base,  the  median  30-60//,  thin-walled, 
with  rather  small  though  distinct  trigones,  the  abundant  chloro- 
plasts  and  small  oil-bodies  usually  aggregated  near  the  middle  of 
the  cell,  cuticle  smooth  ;  gemmae  frequent,  mostly  in  clusters  at 
the  tips  of  the  leaf-lobes,  irregularly  tetrahedral  or  cubico-sphae- 
roidal,  uniseptate  :  lanceolate  or  subulate  underleaves  sometimes 
present  in  association  with  the  9  bracts,  otherwise  wholl}-  wanting  : 
dioicous  :  $  bracts  closely  crowded  together  toward  stem-apex, 
scarcely  different  from  the  ordinary  leaves  ;  antheridia  single  or  in 
pairs,  subglobose,  .18-.  2  mm.  in  diameter,  on  very  short  pedicels, 
paraphyses  none  :  9  bracts  a  little  larger  than  the  leaves,  broader 
than  long,  deepl}'  3-5 -cleft,  more  abundantly  dentate  and  more 


LOPHOZIA 


105 


strongly  crisped:  perianth  oval  to  elongate-obovoid,  2-3.5  mm. 
X  .9-1.5  mm.,  bistratose  at  base,  the  pHcate  mouth  cihate  dentate  : 
seta  5—12  mm.  long;  capsule  subglobose,  reddish-brown,  .7—1 
mm.  in  diameter,  the  valves  thick  and  rigid,  of  three  layers  of  cells, 
the  walls  of  the  cells  of  the  inner  layers  provided  with  very  nu- 
merous annular  or  semiannular  thickenings,  those  of  the  external 
layer  with  similar  but  much  less  perfect  fibers  ;  spores  cinnamon- 
brown,  13-15/^-,  minutely  granulate-papillate;  elaters  fusiform, 
80-145  n  long,  8-10/^  in  greatest  width,  somewhat  geniculate. 

On  logs  or  decaying  wood  in  moist  places.  Sisson,  Siskiyou 
Co.  (40,  41)  ;  near  Mendocino  (672)  ;  near  Eureka  (962). 

All  our  Californian  specimens  with  the  exception  of  no.  41  are 
sterile  and  gemmiferous,  with  the  leaves  mostly  2-,  less  commonly 
3-lobed,  otherwise  subentire,  the  lobes,  with  rare  exceptions, 
merely  acute.  Our  no.  40,  with  leaves  of  this  character,  was  once 
(Erythea,  4  :  49.  1896)  erroneously  referred  to  Jiuigcnnannia  ven- 
tricosa  Dicks.,  from  which  it  may  be  readily  distinguished  by  the 
more  deeply  and  unequally  lobed  and  more  complicate  leaves,  the 
smaller  dorsal  lobe  being  often  inclined  forward  somewhat  parallel 
to  the  stem,  and  by  the  larger,  thinner-walled  leaf-cells. 

Our  description  of  the  sporogonium  has  been  drawn  from  a 
Montana  specimen  collected  by  Mr.  R.  S.  Williams  and  from 
European  material. 

2.  LoPHOZiA  VEXTRicosA  (Dicks.)  Dumort.  Rec.  d'Obs.  Jung.  17. 

1835- 

JnngenjiaiiJiia  vcntricosaYy'icks.  PI.  Crypt.  2  :  14.  1790.  Hook. 
Brit.  Jung.  //.  28.     18 16. 

Jungcrinannia  porpJiyrolciica  Nees,  Naturgesch.  Eur.  Eeberm. 
2:  78.  1836. 

Yellowish-green,  sometimes  reddish-brown  or  subfuscous, 
densely  caespitose  :  stems  .5-1.5  cm.  long,  prostrate  or  ascending, 
sparingly  branched,  often  reddish  underneath,  the  numerous  root- 
hairs  mostly  colorless  and  rather  short  :  leaves  close  or  approxi- 
mate, obliquely  inserted  or  almost  transverse  toward  the  apex, 
semivertical  or  spreading,  slightly  concave,  now  and  then  (espe- 
cially the  superior)  canaliculate  or  obtusely  subcomplicate,  often 
soft  and  somewhat  flaccid,  ovate-quadrate,  2-  (rarely  3-)  lobed,  the 
sinus  broad,  mostly  rounded-obtuse,  occasionally  gibbous,  descend- 
ing i-2  the  length  of  the  leaf,  lobes  acute,  occasionally  apiculate, 
rarely  subobtuse,  often  erose  through  the  formation  of  gemmae. 


106 


JUXGERMANNIACEAE 


these  marginal  or  more  commonly  in  clusters  at  the  apices  of  the 
lobes,  irregular,  somewhat  tetrahedral  or  subcubical,  mostly  uni- 
septate  ;  median  leaf-cells  rounded-hexagonal,  24-32  //.,  trigones 
distinct  but  variable  in  size,  the  cuticle  smooth  or  slightly  rough- 
ened :  subulate  or  broader  and  bifid  underleaves  usually  present  in 
association  with  the  ?  bracts  and  sometimes  on  young  "subfloral  " 
innovations,  otherwise  wholly  wanting  :  dioicous  :  androecia  term- 
inal, oval ;  ^  bracts  imbricate,  transversely  inserted,  obtusely  sub- 
complicate,  ventricose  ;  antheridia  in  pairs  or  single,  oval,  .12-18 
mm.  in  greatest  diameter,  on  short  stalks  composed  of  a  single 
row  of  cells,  accompanied  by  a  few  short  paraphyses  :  9  bracts 
somewhat  larger  than  the  leaves,  the  inmost  deeply  and  unequally 
2-5-  (mostly  3-  or  4-)  lobed,  often  slightly  plicate,  the  lobes 
acute  or  subobtuse  :  perianth  cylindrical-obovoid,  2—3  mm.  long, 
1-1.5  mm.  in  greatest  width,  unistratose  except  at  extreme  base, 
the  plicate  mouth  ciliolate-denticulate  :  calyptra  mostly  bistratose  : 
seta  10-15  mm.  long  ;  capsule-valves  purplish-  or  yellowish-brown, 
.9-1.1  mm.  long,  of  three  layers  of  cells,  the  inner  with  close  an- 
nular or  semiannular  fibers,  the  outer  with  nodular  or  imperfect 
semiannular  thickenings  ;  spores  yellowish-  or  reddish-brown, 
finely  granulate-papillate,  12-14  u.;  elaters  fusiform,  subobtuse, 
80-1 30  u.  X  8-10 

On  soil  in  moist  places  and  on  decaying  wood.  ]\It.  Dana  at 
about  3100  m.  alt.,  associated  with  Lophozia  Jieterocolpa,  Junger- 
maiuiia  Danicola,  and  J.  Bolandcri  (Dr.  H.  N.  Bolander,  Septem- 
ber, 1866);  between  Farewell  Gap  and  Mineral  King,  Tulare  Co., 
alt.  3500  m.  (Coville  and  Funston,  Death  Valley  Expedition,  no- 
1573,  Aug.  10,  1 891);  beside  "  Horse  Camp  Spring,"  above  timber- 
line,  Mt.  Shasta,  alt.  (est.)  about  3100  m.  (Howe,  August,  1894). 

These  three  high-altitude  specimens,  the  second  $  and  the* 
others  sterile,  seem  clearly  to  fall  in  the  form -cycle  of  LopJiozia 
ventricosa,  agreeing  essentially  with  the  soft-leaved  terricolous  con- 
ditions of  this  species  from  other  parts  of  the  world,  as  represented, 
for  example,  in  Carr.  and  Pears.  Brit.  Hep.  no.  171.  The  plants 
grow  in  densely  compacted  tufts,  and  the  walls  of  the  leaf-cells,  in 
the  last  two  specimens  especially,  are  but  slightly  thickened  at  the 
angles  save  in  the  more  exposed  parts.  Lindberg  and  Arnell 
(Kongl.  Sv.  Vet.  Akad.  Handl.  23':  50.  1889)  would  restrict  the 
application  of  the  specific  name  voitricosa  to  forms  with  poorly 
developed  trigones,  but  Lindenberg  (Syn.  Hep.  Eur.  86.  1829), 
who,  it  would  appear,  had  examined  original  Dicksonian  speci- 


LOPHOZIA 


107 


mens,  affirms  that  these  agree  with  his  form  confcrta  characterized 
partly  by  a  ''textiira  firma,''  instead  of  the  ''contextit  laxiore''  of  the 
form  laxa,  which  possibly  indicates  that  the  form  with  well  devel- 
oped trigones  should  be  considered  typical. 

The  leaves  of  the  Mt.  Shasta  plant  have  a  rather  unusually 
pronounced  tendency  toward  being  complicate,  but  this  feature  is 
observable  now  and  then  in  specimens  from  other  localities.  The 
form  or  variety  inhabiting  chiefly  decaying  logs  and  stumps  (y. 
porphyroleitca  Nees)  is  to  be  expected  to  occur  in  California,  par- 
ticularly, perhaps,  in  the  northern  coast  counties. 

We  have  observed  in  North  American  specimens  of  Lophozia 
ve7itricosa  the  two  kinds  of  coloration  of  the  capsule  noted  by 
Limpricht  (Cohn,  Krypt.-Fl.  Schles.  i:  280,  281.  1877),  but 
cannot  determine  that  this  character  stands  in  any  constant  relation 
to  other  characters  which  have  been  supposed  by  some  to  distin- 
guish Jungermamiia  porphyroleuca  from  J.  ventricosa.  In  Austin's 
Hep.  Bor.-Am.  no.  36,  for  instance,  the  purplish-brown  or  ' 'violet- 
red-brown"  capsules  occur  on  gametophytes  which  exhibit  leaf- 
cells  with  the  strongly  developed  trigones  and  roughened  cuticle 
supposedly  characteristic  of  J.  porphyroleuca. 

It  is  extremely  doubtful  if  the  original  of  Jimgermannia  ven- 
tricosa exists  in  Dickson's  Herbarium  in  the  British  Museum.  Two 
scraps  are,  however,  to  be  found  there,  under  which  is  written,  in 
pencil,  in  what  is  said  to  be  Dickson's  hand, 

"  bid««  With.  ed.  3 
ventricosa ' ' 

But  these  belong  to  the  genus  LopJwcolea.  The  leaves  of  these, 
with  the  exception  of  the  younger,  are  considerably  less  deeply 
cleft  than  those  in  the  figures  of  Micheh  and  Dillenius  which  Dick- 
son cites,  and  in  allusion  to  which  he  remarks  :  Folia  in  nostra 
profundius  fissa,  quam  in  figuris  Michelii  et  Dillenii  depinguntur." 
A  young  trigonous-prismatic  perianth,  twice  as  long  as  broad — 
which  Dickson  probably  could  not  have  described  by  ''vagina 
sphaeroidea  " — is  present  on  one  of  the  scraps.  The  reference  to 
the  third  edition  of  Withering  would  indicate  that  the  inscription 
under  these  specimens  was  written  at  least  six  years  after  the  orig- 
inal publication  of  Jiingenna?inia  ventricosa. 


108 


JUNGERMAXXIACEAE 


3.  LoPHOziA  HETEROCOLPA  (Thed.) 

JiLngcrinannia  lictcrocolpa  Thed.  Kongl.  Sv.  Vet.  Acad.  Handl. 
1838  :  52,  72.  //.  7.     1839.     [As /.  heterocolpos.'] 

Jungermannia  Miilkri  heterocolpos  G.  L.  &  X.  S)'n.  Hep.  99. 
1844. 

Jungermannia  Wattiana  Aust.  Bull.  Torr.  Bot.  Club,  3:  11. 
1872. 

Jiuigerniannia  Mi'illeri  Danaensis  Gottsche  ;  Underw.  Bot.  Gaz. 
13  :  1 14.  //.  6.  1888. 

Yellowish-  or  lurid -green,  densely  caespitose  or  creeping 
among  mosses:  stems  6-15  mm.  long,  .12-3  mm.  thick,  pros- 
trate or  ascending,  sparingly  ramose,  the  9  commonly  continued 
through  the  development  of  an  innovation  from  underneath  the 
perianth,  the  median  innovation  sometimes  accompanied  by  one  or 
two  lateral ;  root-hairs  numerous,  long,  colorless  :  leaves  close  or 
approximate,  obliquely  inserted,  patent  or  sometimes  subvertical 
toward  the  stem-apex,  usually  firm  in  texture,  rarely  somewhat 
soft  and  flaccid,  not  decurrent,  orbicular-ovate  or  subquadrate- 
elliptical,  .36-1  mm.  x. 35-8  mm.,  the  superior  usually  somewhat 
concave  dorsally,  the  inferior  more  or  less  arcuate-conx^ex,  all 
emarginate-bilobed  \-\  their  length,  the  acute  or  obtuse  sinus 
commonly  gibbous-recurved,  the  lobes  obtuse,  rarely  subtri- 
angular-acutate,  those  of  the  inferior  leaves  often  slightly  arc- 
uate-recurved;  marginal  and  superior  leaf-cells  16-32  ft  (mostly 
20-25  fi),  trigones  well  developed  (6—1 1  /i),  the  cuticle  with 
very  obscure  hyaline  roughenings  :  underleaves  small,  lanceo- 
late-subulate and  subentire  or  bifid  and  sometimes  sparingly 
dentate  at  base  :  gemmae  frequent,  brown,  subglobose-ellipsoidal, 
uniseptate,  24-30  //.  x  16-24  borne  on  the  apices  of  much  modi- 
fied, larger-celled,  3 -ranked  leaves,  these  erect-imbricate  in  a  more 
or  less  elongated  terminal  spike  :  dioicous  :  $  bracts  unidentate 
on  dorsal  margin  ;  antheridia  paired,  on  very  short  stalks  :  9  bracts 
gradually  a  little  larger  than  the  leaves  and  sometimes  with  repand 
margins,  otherwise  similar :  perianth  ovoid  to  obovoid-oblong, 
1.5-2  mm.  X  .9-1.2  mm.,  often  apparently  dorsal  owing  to  the 
early  development  of  a  strong  hypogynous  innovation,  somewhat 
laterally  compressed  in  the  upper  half  when  young,  the  small 
mouth  minutely  denticulate. 

In  ditches,  Mt.  Dana,  alt.  about  3100  m.  (Dr.  H.  N.  Bolandcr, 
September,  1866);  Fish  Creek,  Mariposa  Co.  (Miss  Edith  S. 
Byxbec,  July,  1895). 

In  the  description  oi  Jungerniawiia  Miilleri  Danaensis  (/.  r.),  it 


LOPHOZIA 


109 


is  stated  that  amphigastria  "  are  wanting,  but  an  examination  of 
the  specimen  sent  by  Dr.  Gottsche  to  the  Gray  Herbarium  dis- 
closes underleaves  of  the  /.  Miillcri-Jictcrocolpa  t}'pe  more  or  less 
concealed  by  the  root-hairs,  and  Dr.  Gottsche  had  evidently  in- 
tended to  represent  these  in  his  camera-lucida  tracings,  especially 
in  the  figures  published  by  Professor  Underwood  (/.  c^)  under 
Nos.  v.,  IX.,  X.,  XII.,  and  XV.  The  leaf-cells  of  the  Mt  Dana 
material  are  mostly  less  strongly  collenchymatous  than  those  of 
Scandinavian  specimens  of  Lopliozia  lictcrocolpa  that  we  have  ex- 
amined, yet  portions  of  this  same  material  show  all  the  transitions 
to  trigones  of  the  normal  size.  The  plant  collected  by  Miss  Byx- 
bee  has  larger  leaf-cells,  together  with  larger  trigones,  and  in  this 
respect  makes  an  approach  to  Jiingcrniaunia  Mullcri  Xees,  but 
differs  from  that  species  in  the  usually  obtuse  leaf-lobes,  in  the 
9  bracts  being  entire  (except  for  the  median  cleft)  or  mereh*  re- 
pand,  in  the  ovoid  or  obovoid-oblong  perianth,  which  is  not  at  all 
rostellate,  and  in  the  presence  of  transformed  spicate  gemmiparous 
leaves.  The  perianths  in  this  specimen  are  nearly  all  falsely  dor- 
sal. In  the  original  diagnosis  of  Jungerniannia  Jieterocolpa^  The- 
denius  writes  '\friictii  tcruiinali  (raruis  psetidoalariy  and  his  figure 
shows  a  terminal  perianth.  We  have  seen  but  one  European  peri- 
anth-bearing specimen  of  J.  Jietcrocolpa,  which  we  owe  to  the  kind- 
ness of  Dr.  H.  \V.  Arnell ;  in  this  about  one  fourth  of  the  peri- 
anths have  become  falsely  dorsal  or  lateral.  In  this,  too,  the  peri- 
anths have  only  about  two  thirds  of  the  length  assigned  to  the 
perianths  of  the  species  of  Lindberg  (Kongl.  Sv.  Vet.  Akad. 
Handl.  23'^:  43.  1889),  agreeing  closely  in  this  respect  with  the 
American  forms. 

Inasmuch  as  Limpricht  (Cohn,  Kr}'pt.-Fl.  Schles.  i  :  276. 
■  1877)  has  remarked  that  an  apparent  original  of  Jungcnnannia 
Bantriensis  Hook,  (found  in  Herb.  v.  Flotow,  with  the  legend, 
J.  bantriensis  X.  v.  E.  Irland,  Herb.  Hook.l  dd.  X.  ab.  E.")  is  the 
gemmiferous  ^  plant  of  Jiingermannia  Mulleri  and  as  Lindberg 
(Kongl.  Sv.  Vet.  Akad.  Handl.  23-^:  43.  1889)  has  observed 
that  the  gemmiparous  forms  of  J.  Mullcri  of  authors  ought  prob- 
ably to  be  referred  to  J.  heterocolpa,  we  have  made  an  effort  to 
examine  the  Hookerian  material  of  J.  Bantriensis.  In  response  to 
an  inquiry  addressed  to  Sir  W.  T.  Thiseiton-Dyer,  as  to  whether 


110 


JUXGERMANXIACEAE 


the  original  specimen  of  Hooker's  Jungerjumuiia  Bantricnsis,  col- 
lected by  Miss  Hutchins  at  Bantry,  Ireland,  was  still  preserved  at 
Kew,  we  received  the  following  statement  :  "  There  is  no  speci- 
men of  this  at  Kew  marked,  'Bantry,  Miss  Hutchins.'  But  on  a 
specimen  not  localized  is  written  (I  believe  in  Wilson's  handwrit- 
ing) :  '  This  is  probably  the  specimen  mentioned  in  Brit.  Jung'*^ 
See  note  to  description  of  J.  stipidacca'  The  note  referred  to 
contains  the  diagnosis  of  J.  Bantriensis.  A  portion  of  this  speci- 
men is  enclosed."  This  specimen  proves  to  be  HarpaiitJius  Floio- 
viaiius  Nees.  It  is  readily  distinguishable  from  any  of  the  Lophozia 
Mi'illeri-Jieterocolpa  alliance  by  the  decurrent  leaves,  the  wholly 
smooth  cuticle,  the  larger  underleaves,  and  the  short  ventral  9 
branches,  which  here  bear  unfertilized  archegonia  ;  some  of  the 
stems  are  gemmiparous  at  the  apex,  but  the  gemmiparous  leaves 
are  not  transformed  in  any  such  way  as  in  LopJwzia  lictcrocolpa. 
A  comparison  of  this  specimen  with  Hooker's  figure  and  descrip- 
tion precludes  the  possibility  of  its  being  the  desired  original. 

To  the  friendly  generosity  of  Mr.  \V.  H.  Pearson,  we  owe  the 
privilege  of  examining  a  specimen  marked  "  Jung.  Bantriensis 
Hook.  Bantry.  Coll.  Miss  Hutchins,  Nov.  19,  1812.  Original^ 
This  specimen,  though  sterile,  we  can  easily  believe  to  have  been 
a  part  of  the  original  material  studied  by  Hooker.  It  agrees  es- 
sentially with  JiDigcrniannia  HornschucJiiana  Nees  as  represented 
in  Rab.  Hep.  Eur.  128,  differing  chiefly  in  the  slightly  smaller 
leaf-cells  and  the  more  prominently  verruculose-striolate  or  gran- 
ulate-papillate cuticle.  From  Lopliozia  Jietcrocolpa  it  differs  in  the 
larger  (about  1.5  mm.  wide)  flaccid  decurrent  leaves,  which  are  bi- 
dentate  rather  than  bilobed,  the  usually  acute  teeth  being  sepa- 
rated by  a  broad,  shallow  sinus,  in  the  larger  leaf-cells  (marginal 
and  superior  28-50  /i),  in  the  very  distinctly  roughened  cuticle, 
and  in  the  absence  of  gemmae. 

4.  Lophozia  inflata  (Huds.) 
Jimgerina)ima  i)iflata  Huds.  Fl.  Angl.  511.     1778  [ed.  2]. 
Gymnocolca  iiifla4a  Dumert.  Rec.  d'Obs.  Jung.  17.  1835. 
Green,  yellowish-brown,  or  sometimes  blackening,  with  (espe- 
cially in  the  darker  forms)  a  somewhat  oleaginous  lustre,  usually 
forming  soft  closely  interwoven  mats  :  stems  prostrate,  ascending, 
or  occasionally  erect,  6-20  mm.  long,  .09-25  mm.  in  diameter, 


LOPHOZIA 


111 


mostly  weak  and  slender,  sparingly  subdichotomous,  or  with  a  few 
lateral  or  latero-ventral  branches,  often  innovating  from  near  the 
base  of  the  perianth  ;  root-hairs  short,  scanty,  colorless  or  yellow- 
ish :  leaves  usually  distant  or  contiguous,  obliquely  inserted, 
patent-horizontal  to  erecto-patent,  now  and  then  reflexed,  orbicular- 
ovate  to  subquadrate-oblong,  .36-9  mm.  x  .27-8  mm.,  some- 
what decurrent  dorsally,  concave  or  nearly  plane,  bilobed  y^—Yi, 
their  length,  the  sinus  rather  narrow,  more  or  less  obtuse,  the 
margin  otherwise  entire  or  slightly  repand,  lobes  obtuse,  nearly 
parallel,  rarely  inclined  toward  each  other  or  subdivergent,  the 
dorsal  a  little  the  smaller  ;  leaf-cells  rather  thick-walled,  with  in- 
distinct trigones,  the  superior  and  median  24—40/7.,  the  basal 
scarcely  larger,  cuticle  smooth  or  obscurely  striolate  :  underleaves 
wanting  unless  in  association  with  the  9  bracts  :  dioicous  :  androe- 
cium  terminal  or  median,  ^  bracts  imbricate,  concave,  mostly 
broader  than  long,  almost  transversely  inserted  ;  antheridia  ovoid, 
.2  mm.  in  greatest  diameter,  on  very  short  stalks,  unaccompanied 
by  paraphyses  :  9  bracts  often  a  little  smaller  than  the  leaves, 
otherwise  similar  :  perianth  much  exserted,  sometimes  exposed  to 
the  extreme  base  or  even  appearing  stipitate,  now  and  then  falsely 
dorsal  through  the  development  of  a  "  subfloral  "  innovation,  pro- 
late-ellipsoidal or  elongate-pyriform,  2-2.7  mm.  x  .85-1.2  mm., 
inflated,  smooth  throughout  or  obscurely  plicate  only  at  the  obtuse 
apex,  the  wall  bistratose  in  the  basal  third  or  fourth,  the  mouth 
with  a  few  short  connivent  teeth  :  capsule  elongate-ovoid,  its  wall 
of  three  layers  of  cells,  the  external  layer  with  nodular  or  colum- 
nar thickenings,  the  others  with  semiannular  ;  spores  14-18 
granulate -papillate,  elaters  contorted,  slightly  attenuate  at  extremi- 
ties, 120— 1 80/7-  long,  8-10  «  in  maximum  width. 
In  swamps,  on  wet  rocks,  etc. 
Mt.  Dana,  Cal.,  legit  H.  N.  Bolander  "  in  herb.  Underwood. 
The  specimen  is  wholly  sterile,  but  we  feel  no  doubt  as  to  its 
identity  with  the  above  species. 

5.  LoPHOZiA  ovATA  (Dicks.) 
Jiuigcrmajinia  ovata  Dicks.  PI.  Crypt.  Brit.  3:   11.  pi.  8.f.  6. 
1793. 

Jiingermannia  Dicksoni  Hook.  Brit.  Jung.       ^8.     18 16. 
DiplopJiylhim  Dicksoni  T>\imox\..  Rec.  d'Obs.  Jung.  16.  1835. 
DiplopJiylleia  Dicksoni  Trevis.  Mem.  r.  1st.  Lomb.  III.  4  :  420. 

1877- 

Bright  green  above,  darker  below,  caespitose,  often  in  compact 
extended  mats  ;  stems  8-20  mm.  long,  simple  or  dichotomously 


112 


JUNGERMANNIACEAE 


branched,  prostrate-entangled  or  slightly  ascending,  rather  rigid, 
beset  with  long  root-hairs  almost  to  the  apex  :  leaves  transversely 
inserted,  of  nearly  uniform  size  throughout,  .5-9  mm.  long,  not 
decurrent,  bilobed  to  below  the  middle  (f- f )  with  an  acute  sinus, 
complicate,  the  fold  nearly  a  right  angle  but  rounded  ;  lobes  of 
aboui  equal  length,  acuminate,  obscurely  crenulate  towards  the 
apex,  the  dorsal  narrower,  lanceolate,  suberect  or  ascending,  some- 
what parallel  to  the  stem,  the  ventral  ovate  or  oblong-ovate, 
patent-horizontal  (75^-80°),  submarginate  toward  the  base;  leaf- 
cells  opaque,  firm-walled,  with  small  trigones,  hexagonal-quadrate, 
1 7-24  //,  more  elongated  at  base,  cuticle  minutely  roughened  : 
underleaves  none  :  antheridia  i  or  2,  subglobose,  without  paraphy- 
ses.  on  short  pedicels  in  the  axils  of  closer  somewhat  ventricose 
leaves  with  appressed  imbricate  dorsal  lobes  :  9  bracts  larger  than 
the  leaves,  long-acuminate,  sparingly  dentate-;  perianth  oval  or  ob- 
long, 2-3  times  length  of  the  involucral  leaves,  deeply  plicate,  the 
mouth  ciliate-lacinulate. 

Exsicc.  Hep.  Am.  193  (as  Diplophylleia  Dicksoni). 

On  the  face  of  a  shaded  cliff  near  Cazadero,  Sonoma  Co.,  on  the 
road  to  Fort  Ross  (Mar.  14,  1896).  Collected  also  by  Dr.  Bolan- 
der  at  Mendocino  City.  These  are  the  only  American  stations 
thus  far  known  for  this  hepatic.  Our  specimens  are  sterile  or  bear 
antheridia  alone,  so  that  the  species  is  presumably  dioicous.  The 
description  of  involucre  and  perianth  is  drawn  from  Husnot's  Hep. 
Gall.  no.  29.  The  leaves  are  rarely  three-lobed  in  the  Californian 
plants  as  also  in  the  European. 

The  identification  of  Dickson's  Jiingermaniiia  ovata  'V  spccwiiiic 
auctoris"  was  first  made  by  Lindberg  (Muse.  Scand.  7.  1879). 
Mr.  A.  Gepp  of  the  British  Museum  has  kindly  at  our  request  ex- 
amined the  specimen  of  J.  ovata  Dicks,  in  the  Dickson  herbarium 
and  confirms  the  determination  of  Lindberg.  Mr.  Gepp  writes  : 
"Our  specimen  of  J.  ovata  Dicks,  is  labeled  ovata  by  Dickson,  and, 
so  far  as  the  arrangement  of  the  specimens  goes,  it  is  the  un- 
doubted type  of  the  species." 

The  rounded,  instead  of  acutely  carinate,  fold  of  the  leaf  suffi- 
ciently removes  the  species  from  the  genus  Diplophylleia  and  places 
it  in  the  SpJienolohts  section  of  LopJwzia. 

22.  PLAGIOCHILA  Dumort.  Rec.  d'Obs.  Jung.  14.  1835. 
Martinellius  S.  F.  Gray  /.  /.  Nat.  Arr.  Brit.  PI.  i  :  690.  1S21. 
Radula  Dumort././.  Comm.  Bot.  112.  1822. 


Plagiochila 


113 


Radula  §  Plagiochila  Dumort. /. /.  Syll.  Jung.  42.     183  I. 

Plants  usually  large.  The  primary  stems  creeping,  radiculose, 
nearly  leafless,  rhizome-like  ;  secondary  stems  foliose,  commonly 
ascending  or  suberect,  stout,  rufous  or  dark-brown,  rarely  pallid, 
mostly  destitute  of  root- hairs,  simple  or  dichotomously  branched, 
less  commonly  pinnate,  often  subdendroid.  Leaves  large,  succu- 
bous,  in  most  species  alternate,  piano-distichous  or  deflexed,  some- 
times secund,  decurrent  dorsally  and  more  or  less  ventrally,  un- 
symmetrical  ;  the  apex  mostly  rounded  or  truncate,  dentate  or 
spinose,  rarely  entire  ;  the  dorsal  margin  nearly  straight  or  slightly 
concave,  recurved,  especially  toward  the  base,  in  most  cases  sub- 
entire  ;  ventral  margin  arcuate,  plane  or  decurved  at  the  base, 
dentate  or  spinose,  very  rarely  entire  ;  leaf-cells  generally  medium- 
sized  or  rather  large,  with  distinct  trigones.  Underleaves  wanting 
or  occurring  only  toward  the  stem-apex  or  sometimes  present 
throughout,  small  or  minute,  entire  or  irregularly  bi-multi-fid. 
Androecium  spicate,  terminal  or  median  ;  $  bracts  smaller  than 
the  leaves,  transversely  inserted,  closely  imbricate,  their  bases  erect, 
ventricose-saccate,  the  basal  margins  highly  adnate  to  the  stem  ; 
antheridia  i-io.  Archegonia  very  numerous,  terminal  on  the 
main  (secondary)  stem  or  the  branches.  ^  bracts  distinct,  free, 
usually  differing  but  little  from  the  leaves,  somewhat  larger  and 
more  strongly  toothed.  Perianth  exserted,  strongly  compressed 
laterally  (at  least  when  young),  with  a  more  or  less  winged  suture 
dorsally  and  sometimes  also  ventrally,  campanulate  or  obconic- 
cylindrical,  the  wide  mouth  usually  obliquely  truncate,  often  bi- 
labiate, the  lips  dentate-spinose  or  ciliate-fringed.  Calyptra  free, 
included.  Capsule  globose-oval,  dehiscing  to  the  base  by  four 
straight  sometimes  bifid  valves,  these  composed  of  several  layers 
of  cells,  cells  bounding  the  inner  surface  provided  with  numerous 
semiannular  bands  ;  seta  rarely  more  than  twice  the  length  of  the 
perianth.    Elaters  bispiral. 

I.  Plagiochila  asplenioides  (L.)  Dumort.  Rec.  d'Obs.  Jung.  14. 
1835.  Lindenb.  Spec.  Hep.  {Plagiochila)^  no.  pi.  2j.  1839  (^)- 

Jungcrinannia  asplenioides  L.  Sp.  PI.  1131.  1753-  Hook. 
Brit.  Jung.  //.  /j.     18 16. 

Jungenuaimia  porelloidcs  Torrey  ;  Xees,  Naturgesch.  Eur. 
Leberm.  i  :  169.  1833. 

Plagiochila  porelloidcs  Lindenb.  Spec.  Hep.  {Plagiochila),  61. 
pL  12.     1839  (?). 

Densely  caespitose  or  scattered  among  mosses,  pale  green  or 
darkening  :  secondary  stems  ascending  or  erect,  mostly  I- 10  cm. 


114 


JUXGERMAXXIACEAE 


long,  simple,  dichotomous  or  with  a  few  lateral  branches,  light 
brown,  nearly  or  quite  devoid  of  root-hairs  :  leaves  alternate,  sub- 
imbricate,  patent-horizontal  to  erecto-patent,  deflexed,  orbicular- 
obovate  to  obovate-oblong,  1.5-4  mm.  long,  1.2-3  mm.  in  greatest 
width,  the  dorsal  margin  strongly  reflexed,  the  ventral  margin  de- 
curved  and,  like  the  rounded  or  broadly  obtuse  apex,  dentate  or 
subciliate,  or  sometimes  entire  ;  superior  and  median  leaf-cells  24— 
48  a,  all  with  rather  small,  though  distinct,  trigones  :  underleaves 
usually  present,  minute,  .15-.  3  mm.  long,  lanceolate-subulate  and 
entire  or  often  reduced  to  a  small  cluster  of  capillary  filaments  : 
dioicous  :  androecium  terminal  or  median  ;  $  bracts  4—8  pairs, 
their  apices  spreading  or  squarrose  ;  antheridia  1-3,  on  stalks 
about  equaling  their  long  diameter  :  9  bracts  scarcely  different  from 
the  leaves:  perianth  obconic-oblong,  mostly  4-6  mm.  long,  1.6— 
2.2  in  greatest  width,  sometimes  falsely  lateral  through  the  devel- 
opment of  an  innovation,  compressed  and  somewhat  twisted  toward 
the  bilabiate,  ciliate-dentate  or  denticulate  mouth  :  seta  15-30  mm. 
long;  elaters  140-200  ti  long,  somewhat  tapering  at  the  ends,  9- 
14  a  in  maximum  width  ;  spores  16-20  n,  minutely  punctulate. 

On  rocks,  soil,  and  logs,  in  moist  woods.  Blue  Lake  (1005,  $) 
and  Deer  Creek  Canon,  Russ  &  Graham's  Ranch  (1064,  c.  pcr\ 
Humboldt  Co.  Measurements  of  spores  and  elaters  are  drawn 
from  G.  &  R.  Hep.  Eur.  620. 

23.  LOPHOCOLEA  Dumort.  Rec.  d'Obs.  Jung.  17.  1835. 

JiingcvDiannia  §  LopJiocolea  Dumort.  Syll.  Jung.  59.    183  i. 

Plants  usually  moderately  large,  rarely  small,  soft  and  flaccid, 
mostly  yellowish-green  or  pallescent,  forming  rather  thin  effuse 
mats  or  repent-spreading.  Stems  creeping,  subsimple  or  irregu- 
larly ramose,  the  branches  lateral  or  latero-ventral,  those  bearing 
the  sexual  organs  sometimes  ventral  ;  root-hairs  in  tufts  at  the 
base  of  the  underleaves,  colorless,  commonly  long.  Leaves  alter- 
nate or  (in  exotic  species)  often  opposite,  succubous,  decurrent 
dorsally,  for  the  most  part  obliquely  oblong-ovate,  often  somewhat 
triangular,  occasionally  subquadrate,  truncate  or  more  commonly 
bidentate  or  bifid  at  apex,  with  broad  or  narrow  sinus,  the  margins 
otherwise  very  entire  or  (in  extra-limital  species)  now  and  then 
serrulate,  spinose,  or  ciliate-fringed  ;  leaf-cells  usually  rather  large 
or  medium-sized,  thin-walled,  occasionally  with  small  trigones  at 
the  angles.  Underleaves  present  throughout,  cuneiform,  ovate,  or 
subquadrate  in  general  outline,  in  our  species  deeply  bi-  or  quadri- 
fid  with  subulate,  mostly  diverging  segments.  Dioicous,  autoicous, 
or  paroicous.    Androecium  occupying  the  end  or  middle  of  a 


LOPHOCOLEA 


115 


branch  ;  S  bracts  several  pairs,  commonly  smaller  than  the  leaves, 
more  erect  and  imbricate,  with  an  incurved  lobule  at  the  saccate 
base,  antheridia  large,  usually  solitary.  Archegonia  numerous, 
terminal  on  the  main  stem  or  on  somewhat  elongated  branches. 
Q  bracts  usually  somewhat  larger  than  the  leaves,  often  more 
deeply  bifid  or  more  dentate.  Perianth  exserted,  triangular-pris- 
matic, at  least  when  young,  with  the  third  angle  always  dorsal, 
mostly  2-4  times  longer  than  broad,  3-lobed  at  the  wide  mouth, 
the  lobes  rarely  somewhat  truncate  and  entire,  usually  bifid  and 
more  or  less  ciliate-dentate  or  spinose.  Calyptra  free,  thin,  shorter 
than  the  perianth.  Capsule  ellipsoid-oval,  dehiscing  to  the  base 
by  straight  rigid  valves  ;  capsule  wall  of  several  layers  of  cells, 
those  of  the  external  layer  thick,  with  columnar  or  imperfectly 
semiannular  thickenings,  those  of  the  inner  layers  thin,  the  inmost 
with  numerous  semiannular  bands  ;  seta  long.     Elaters  bispiral. 

Key  to  the  Species. 

Leaves  all  bidentate  or  bifid,  with  long  subulate-acuminate  teeth  or  segments  ;  autoicous. 

I.  Z.  cuspidata. 

Upper  leaves  merely  retuse,  or  entire  and  truncate  ;  paroicous.        2.  L.  heterophylla. 

I.  LoPHOCOLEA  cuspiDATA  (Xces)  Limpr.;  Cohn,  Kiypt.-Fl. 
Schles.  I  :  303.  1877. 

LopJiocolea  bidoitata  ciispidata  Xees,  Xaturgesch.  Eur.  Leberm. 
2:  327.  1836. 

Pallid-green,  closely  caespitose  or  repent-spreading :  stems 
6-20  mm.  long,  .15-27  mm.  in  diameter,  bearing  (especially  the 
fertile)  frequent  diverging  lateral  branches,  occasionally  dichoto- 
mous  ;  root-hairs  few  or  moderately  abundant :  leaves  ovate  to  ob- 
long-ovate, .63-1.6  mm.  X  .45-1.3  mm.,  almost  equally  narrowed 
to  the  apex  on  both  sides,  bidentate  with  a  broad  crescentic  sinus 
or  on  the  more  slender  branches  sometimes  bifid  their  length 
with  a  rather  narrow  subobtuse  sinus,  segments  (or  teeth)  long, 
subulate-acuminate,  subequal  and  mostly  parallel ;  leaf-cells  usu- 
ally pellucid,  somewhat  incrassate,  the  superior  and  median  24— 
48  a,  thin-walled,  trigones  minute  or  wanting,  cuticle  smooth  :  un- 
derleaves  patent,  often  only  bifid  and  extrorsely  unidentate  on 
either  side,  or  the  subulate,  commonly  divergent,  principal  seg- 
ments again  bifid  :  autoicous  ;  antheridia  and  archegonia  borne  in 
most  cases  on  more  or  less  elongated  ventral  or  latero -ventral 
branches  :  androecium  terminal,  rarely  median  ;  $  bracts  some- 
what similar  to  the  leaves  or  smaller,  less  acutely  lobed,  and  more 
imbricate,  saccate  dorsally  at  the  base,  the  inflexed  margin  com- 
monly provided  with  one  or  two  teeth  or  lobes  ;  antheridia  mostly 


116 


JUXGERMAXXIACEAE 


single,  SLibglobose,  .2  5-.  3  mm.  in  diameter,  the  ver>'  short  stalk 
of  a  single  row  of  cells  :  9  branches  2-10  mm.  long  (archegonia 
sometimes  apparently  terrninal  on  the  main  stem),  lea\'es  increas- 
ing in  size  upward,  becoming  more  erect  and  crowded  and,  like  the 
9  bracts,  externally  canaliculate-concave  ;  9  bracts  bifid  ^3-^ 
their  length,  otherwise  usually  entire,  bracteole  similar,  almost 
equally  large  :  perianth  longer  than  the  bracts,  triangular-pris- 
matic, 2-3.5  X  -8-1.3  nim.,  3-lobed  %-y2  its  length,  the 
lobes  usually  bifid  and  often  sparingly  dentate  :  capsule  dark- 
brown,  1-1.2  mm.  X  .65-.72  mm.;  spores  16-24  minutely 
punctulate  ;  elaters  contorted,  somewhat  attenuate  at  the  extrem- 
ities, 160-220  a  long,  9-12  //.  in  maximum  width. 

On  decaying  sticks,  logs,  and  stumps,  in  moist  woods,  rarely 
on  moss-cov^ered  trunks  of  living  trees.  Not  uncommon  in  the 
Coast  Range  Mountains  from  Marin  County  northward.  Often 
with  perianths,  yet  rarely  with  capsules.  Olema  (27),  Marin  Co.; 
North  Fork  of  the  Little  River  (631,  635,  653)  and  Mendocino 
(697),  Mendocino  Co.;  Eureka  (901)  and  Blue  Lake  (988),  Hum- 
boldt Co. 

LopJiocolca  Lcibcrgii  Underwood  and  Cook,  Hep.  Am.  no.  70 
(without  diagnosis)  cannot,  we  think,  be  safely  distinguished  from 
the  above  species. 

2.   LoPHOCOLEA  HETEROPHYLLA  (Schrad.)  Dumort.  Rec.  d'  Obs. 

Jung.  17.  1835. 

Juiigcniiaiinia  hctcrophyila  Schrad.;  Schrad.  Jour.  Bot.  1801  : 
66.     1803.     Hook.  Brit.  Jung. //.  J/.  1816. 

Yellowish-green,  caespitose  or  repent-spreading :  stems  5— 
20  mm.  long,  .15—27  mm.  in  diameter,  closely  creeping,  usually 
with  frequent  lateral  branches,  root-hairs  numerous  :  leaves  ovate 
to  subquadrate-oblong,  .54-1.3  mm.  x.4-1  mm.,  very  obliquely 
or  almost  longitudinally  inserted,  semivertical,  dimorphous,  the 
lower  obtusely,  or  sometimes  acutely,  bidentate  or  bilobed,  with  a 
rounded-obtuse  sinus,  the  upper  retuse,  or  entire  and  truncate  ; 
superior  and  median  leaf-cells  24-48  n,  the  thin  walls  with  distinct 
trigones  at  the  angles,  cuticle  nearly  smooth  :  underleaves  usually 
appressed,  bifid  to  below  the  middle,  the  subulate-acuminate  seg- 
ments each  commonly  furnished  externally  with  a  single  tooth  or 
short  cilium  :  paroicous,  rarely  heteroicous  :  $  bracts  3-5  pairs 
immediately  below  the  archegonia  or  perianth,  the  uppermost 
functioning  also  as  9  bracts,  almost  transv^ersely  inserted,  close, 
erect-appressed  with  commonly  squarrose  apices,  truncate  or  re- 
tuse, dorsally  ventricose  at  base  with  one  or  two  inflexed  lobules 


Chiloscyphus 


or  teeth,  androecia  rarely  occurring  also  at  a  distance  from  the 
archegonia  either  on  the  same  axis  or  on  special  branches  ; 
antheridia  globose,  about  .27  mm.  in  diameter,  the  short  stalk  of 
a  single  series  of  cells  :  9  bracts  a  little  larger  than  the  subjacent 
perigonial  leaves,  otherwise  similar ;  bracteole  deeply  bifid  with 
lanceolate-subulate  segments  :  perianth  terminal  on  a  more  or  less 
elongated  late ro -ventral  branch,  longer  than  the  bracts,  cylindri- 
cal or  somewhat  goblet-shaped,  1.3-3.2  mm.  x  .63-1.3  mm.,  3- 
angled  toward  the  mouth  and  3-lobed  1— i  its  length,  the  lobes 
subtruncate,  rarely  acute,  repand-dentate  :  capsule  brown,  i  — 
1.2  mm.  X  .75—1  mm.;  spores  12—16/^,  minutely  punctulate ; 
elaters  contorted,  slightly  attenuate  at  the  extremities,  1 10-2 10 /y. 
long,  8—1 1  fJi  in  greatest  width. 

On  moldering  wood  (logs,  stumps,  etc.)  in  moist  caiions,  oc- 
curing  rarely  on  moist  soil  in  which  there  is  considerable  vege- 
table matter.  "  Redwood  Canon"  (near  Mill  Valley)  and  Olema, 
Marin  Co.;  Turner's  Canon  (between  Cazadero  and  Fort  Ross), 
Sonoma  Co.;  Big  River  Boom  (near  Mendocino,  684),  North 
Fork  of  the  Little  River  (695),  and  near  Half-Way  House" 
(726),  Mendocino  Co.;  Eureka  (919,  950,  952,  1220),  Hum- 
boldt Co. 

No.  695  was  found  in  a  rivulet,  apparently  of  only  periodic 
duration,  in  "The  Plains,"  North  Fork  of  the  Little  River,  the 
substratum  being  sand  mixed  with  vegetable  remains.  The  plants, 
in  this  case,  are  wholly  sterile,  yet  they  seem  to  belong  with 
Lophocolea  heterophylla,  though  their  habit  of  growth  is  more 
densely  caespitose  and  the  leaves  are  often  more  acutely  bidentate 
than  is  usual.  All  our  other  Californian  specimens  occur  upon 
dead  wood. 

24.  CHILOSCYPHUS  Corda   [as   Chcilocyphos'];  Opiz,  Beitr. 

651.  1829.* 

Vegetative  characters  of  the  genus  as  a  whole  agreeing  very 
closely  with  those  of  Lophocolea.  Leaves  more  often  entire. 
Archegonia  terminal  on  a  very  short  ventral  or  latero-ventral  branch. 
9  bracts  i  or  2  pairs,  the  outer  pair  (if  present)  minute,  the  inner  al- 
ways considerably  smaller  than  the  leaves.  Perianth  usually  cam- 
panulate  or  obconical,  3 -angled  only  toward  the  3-lobed  mouth. 
Calyptra  carnose,  especially  in  the  lower  part,  shorter  than  the 
perianth  or  often  much  exserted.  Sporogonium  essentially  as  in 
Lophocolea. 


*  See  footnote,  p.  33. 


118 


JUXGERMANXIACEAE 


I.  Chiloscyphus  polyaxthos  (L.)  Corda  ;  Opiz,  Beitr.  651.  1829.'^ 
Jungcrinaiiiiia  polyanthos  L.  Sp.  PL   1131.     1753-  Hook. 

Brit.  Jung.  //.  62.      18 16. 

Jungcrniannia  pallescens  Schrad.  Syst.  Samml.  Krypt.  Ge\Y.  2  : 

7-  1797- 

Chiloscyphus  pallescens  Dumort.  Syll.  Jung.  67.     183  i. 
Chiloscyphus  lophocolcoides  Nees,  Naturgesch.  Eur.  Leberm. 
2:365.  1836. 

Green  or  pallescent,  closely  or  laxly  caespitose  :  stems  1-3 
cm.  long,  .18-45  n^'^-  ii"^  diameter,  prostrate  or  ascending,  spar- 
ingly branched,  root-hairs  conspicuous  or  nearly  obsolete  :  leaves 
close-imbricate  or  approximate,  patent-horizontal,  spreading  in  the 
plane  of  the  stem  or,  toward  the  stem-apex,  ascending  or  semiver- 
tical,  quadrate-orbicular  to  oblong,  .7—1.6  mm.  x  .54-1.2  mm., 
more  or  less  concave  ventrally,  the  apex  rounded,  subtruncate,  or 
retuse  ;  leaf-cells  translucent,  the  superior  and  median  32—65  a; 
the  walls  somewhat  thickened,  yet  exhibiting  obscure  trigones  or 
none  at  all,  cuticle  slightly  granulate  or  nearly  smooth  :  under- 
leaves  oblong-ovate,  bifid  to  the  middle  or  below,  the  segments 
often  hnear- subulate,  occasionally  bearing  a  single  tooth  or  short 
cilium  externally  ;  autoicous  ;  androecium  terminal  or  median  on 
the  main  stem,  $  bracts  with  small  dorsal  pcuches,  otherwise  sim- 
ilar to  the  leaves,  the  margin  of  the  pouch  usually  with  several 
short  inflexed  cilia  ;  antheridia  single  or  less  commonly  in  pairs, 
subglobose,  .14-2  mm.  in  diameter,  the  very  short  stalk  composed 
of  a  single  row  of  cells;  axis  of  9  branch  .3-.6  mm.  long;  9 
bracts  a  single  pair,  with  sometimes  rudiments  of  a  second  (outer) 
pair,  very  much  smaller  than  the  leaves,  retuse  or  bifid,  bracteole 
minute,  bifid  ;  perianth  goblet-shaped  or  obovoid,  2—2.7  mm.  long, 
.9-1.6  mm.  in  greatest  width,  deeply  3-lobed,  the  lobes  subtrun- 
cate and  nearly  entire,  rarely  spinose-dentate  :  calyptra  exserted  or 
included:  capsule  brown,  globose-oval,  1.2-1.5  mm.  x  1.1-1.4 
mm.,  the  valves  rigid,  of  4  or  5  layers  of  cells,  those  of  the  exter- 
nal layer  thick,  with  columnar  or  imperfectly  semiannular  thick- 
enings, those  of  the  inmost  layer  with  very  numerous  semiannular 
bands;  spores  14-20 minutely  granulate;  elaters  1 20-200 /i  x 

8-  1 1  a. 

On  old  logs  and  on  moist  ground  among  mosses.  "  Paper 
Mill  Creek"  [Marin  Co.?]  (Dr.  H.  N.  Bolander,  Sept.  9,  1863); 
Sisson,  Siskiyou  Co.  (Howe,  37);  near  Hay  Fork,  Trinity  Co. 
(i  125) ;  North  Fork  of  the  Little  River  (636)  and  near  Mendocino 


*  See  footnote  under  Ricciocarpiis,  page  33. 


Chiloscyphus 


119 


(683),  Mendocino  Co.;  Big  Valley  ]\Its.  and  White  Horse  Lake, 
Modoc  Co.  (:\L  S.  Baker  and  F.  P.  Nutting). 

In  our  only  specimens  with  sporogonia  (Big  Valley  ]\Its., 
Baker  and  Nutting),  the  calyptra  is  included  within  the  perianth  or 
but  slightly  exserted.  The  9  branches  in  this  originate  a  little  to 
one  side  of  the  middle  line  of  the  ventral  surface  of  the  stem  and 
are  no  more  strictly  ventral  than  in  LopJiocolea  aispidata.  When 
sterile,  there  is  a  possibility  of  confusing  the  species  with  LopJio- 
colea JieteropJiylla,  but  the  more  entire  leaves  with  larger  leaf-cells 
and  usual  absence  of  trigones  and  the  simpler  underleaves  are 
commonly  distinctive.  Antheridia  can  be  found  in  most  specimens 
and  their  distribution  along  the  back  of  the  main  stem  in  smaller, 
more  dorsal  pouches  of  otherwise  scarcely  modified  leaves  is  also 
characteristic. 

\b.  Chiloscyphus  polyanthos  rivularis  (Schrad.)  Nees,  Natur- 
gesch.  Eur.  Leberm.  2:  374.  1836. 

Jungcnnannia  pallescciis  rivularis  Schrad.  Syst.  Samml.  Krypt. 
Gew.  2  :  7.  1797. 

Jungcrjjiannia  aqiiatica  Schrank  (?),  Bai.  Fl.  2:  496.  1789. 

Aquatic,  usually  larger  in  all  parts,  dark  green,  with  a  some- 
what fatty  lustre  :  stems  1.5-10  cm.  long,  more  ramose:  leaves 
sometimes  broader  than  long  (now  and  then  2—2.5  nim.  broad), 
mostly  rounded  at  apex  ;  leaf-cells  opaque,  the  superior  and  me- 
dian 24-50  «:  underleaves  destroyed  or  wanting  except  in  the 
youngest  parts  :  always  (?)  sterile. 

Exsicc.  Hep.  Am.  92. 

In  running  water  or  on  the  banks  of  streams.  "California" 
(Bolander);  Cazadero,  Sonoma  Co.;  Navarro  (Miss  Byxbee),  North 
Fork  of  the  Little  River  (613),  Mendocino  Co.  ;  Eureka  (956)  and 
Russ  &  Graham's  Ranch  (1085),  Humboldt  Co.  ;  near  Douglas 
City  (i  143),  Trinity  Co.  ;  Sisson  (35),  Siskiyou  Co.  ;  Dixey  Val- 
ley and  Bridge  Creek,  Lassen  Co.  (Baker  &  Nutting)  ;  Lake 
Tahoe  (W.  C.  Blasdale,  1897);  region  of  the  Yosemite  (C.  M. 
Cooke,  Jr.,  1896). 

The  variety  rivularis  is  often  very  different  from  the  pale,  large-  . 
and  translucent-celled,  wood-inhabiting  condition  of  the  species, 
yet  the  two  appear  to  be  connected  by  riparian  forms.    Our  Sisson 
specimen,  distributed  in  Hep.  Am.  as  no.  92,  differs  considerably 


120 


JUXGERMAXXIACEAE 


in  general  aspect  from  the  usual  form  of  the  \'ar.  riviilaris  owing  to 
the  proportionally  broader  leaves  which,  in  drying,  are  commonly 
semivertical  or  somewhat  wrapped  around  the  stem.  This  grew 
in  very  cold  swiftly  running  water. 

Jiingcrniannia  aqiiatica  Schrank  is  sometimes  quoted  as  a 
s\nion}'m  of  Cliiloscyplius  polyanthos  rivularis.  If  a  true  synonym, 
the  name  aqiiatica  would  replace  rivularis,  according  to  the  pro- 
visions of  the  Rochester  Code.  But,  personally,  we  are  not  fully 
convinced  that  Jiuigcruianuia  aqiiatica  Schrank  is  really  such  a 
synonym.  Schrank's  description  would,  perhaps,  on  the  whole, 
apply  equally  well  to  Jungcrriiannia  cordifolia  Hook. 

25.  GEOCALYX  Xees,  Xaturgesch.  Eur.  Leberm.  i  :  97,  102. 
1833.    Et  op.  cit.  2:  395.  1836. 

Calypogia  Dumort.  Syll.  Jung.  73.  183 1.  Xot  Calypogeja 
Raddi,  Atti  Soc.  Ital.  Sci.  Mod.  18  :— (33).     18 18. 

Saccogyna  Lindb. /. /.  Xot.  Sallsk.  Fauna  et  Flora  Fenn.  For- 
handl.  13:  377.  1874.  Act.  Soc.  Sci.  f^enn.  10  :  509.  1875. 
Schiffn. /. /.  Eng.  &  Prantl,  Xat.  Pflanzenfam.  i'^  :  93.  1893. 

Plants  resembling  in  habit  Lophocolea  and  Cliiloscyplius.  Stems 
creeping,  simple,  or  with  a  few  branches,  these  springing  from  the 
axils  of  the  underleaves  or  latero -ventral ;  root-hairs  usually  few, 
mostly  from  near  the  bases  of  the  underleaves.  Lea\-es  alternate, 
succubous,  bidentate  or  bilobed.  slightly  decurrent  dorsally.  Un- 
derleaves present  throughout,  much  smaller  than  the  leaves,  bifid 
to  the  middle  or  below,  free  or  connate  with  the  leaf-base  on  one 
side  by  a  very  narrow  isthmus.  Androecium  on  a  short  branch 
from  the  axil  of  an  underleaf,  $  bracts  considerably  smaller  than 
the  leaves,  unequally  and  for  the  most  part  acuminately  bilobed, 
with  an  additional  tooth  or  lobule  on  the  dorsal  margin  ;  antheridia 
single  or  less  commonly  in  pairs,  short-stalked,  without  paraphyses. 
Archegonia  few,  terminal  on  a  very  short  branch  springing  from 
the  axil  of  an  underleaf,  the  branch  in  its  further  development  be- 
coming a  fleshy  pendulous  subterranean  sac  (perigynium)  enclos- 
ing the  young  sporogonium  and  bearing  about  its  mouth  a  few  in- 
conspicuous scale-like  9  bracts,  these  often  destroyed  on  the 
exsertion  of  the  capsule.     Perianth  wanting.     Calyptra  shorter 

*The  differences  in  vegetative  characters  between  Geocalyx  gravcolens  and  Sacco- 
gyna -i  iticulosa  are  so  striking  that  we  feel  unwilling  to  follow  Lindberg,  Carrington, 
Schiffner,  Bescherelle,  and  Kaalaas  in  reducing  Geocalyx  to  Saccogyna — at  least  until 
we  have  had  an  opportunity  of  examining  the  exotic  species  which  are  said  to  hold  an 
intermediate  position. 


Geocalyx 


121 


than  the  perigynial  tube,  with  which  it  is  coalesced  for  two  thirds 
its  length,  the  unfertilized  archegonia  (i,  2,  or  o)  situated  near  the 
base  of  the  free  portion.  Capsule  nearly  cylindrical,  dehiscing  to 
the  base  by  straight  valves,  these  composed  of  two  layers  of  cells, 
the  cells  of  the  outer  layer  with  inconspicuous  columnar  or  nodu- 
lar thickenings,  those  of  the  inner  layer  with  semiannular  bands  ; 
seta  moderately  long  ;  involucellum  short,  goblet-shaped,  finally 
surrounding  the  base  of  the  seta.     Elaters  bispiral. 

I.  Geocalyx  graveolexs    (Schrad.)    Nees,   Naturgesch.  Eur. 
Leberm.  i  :  98.     1833.     Et  op.  cit.  2:  397.  1836. 
Jungcnnannia  graveolcns  Schrad.  S}^st.  Samml.  Kr}^pt.  Gew. 
2:  6.  1797. 

Saccogyna  gravcolcns  Lindb.  x\ct.  Soc.  Sci.  Fenn.  10 :  509. 
1875. 

Bright  or  olive-green,  caespitose  :  stems  6-20  mm.  long,  .22- 
,3  mm.  in  diameter:  leaves  piano-distichous,  less  commonly  semi- 
v^ertical,  imbricate  or  approximate,  rather  rigid,  subquadrate-ovate, 
.8-1.4  ""irn-  X  -55-1  mm.,  bidentate  or  bilobed  (rarely  i)  their 
length,  the  lobes  acute  or  now  and  then  somewhat  obtuse,  parallel 
or  sometimes  slightly  connivent,  the  sinuses  rounded  or  subacute  ; 
leaf-cells  24-48  scarcely  enlarged  toward  the  base,  thin-walled 
with  small  trigones,  the  numerous  chloroplasts  commonly  ar- 
ranged in  a  ring  leaving  a  clear  area  in  the  middle  and  often  a 
hyaline  peripheral  border ;  cuticle  finely  granulate-papillate  : 
underleaves  appressed,  bifid  to  below  the  middle,  the  lobes  lanceo- 
late, entire,  nearly  parallel  :  autoicous  :  $  branch  .3—1.2  mm. 
long  :  9  bracts  4  or  5  :  perigynium  cylindrical,  about  2.5-3  x 
I  mm.,  its  entire  outer  surface  sparingly  radiculose  :  capsule  i  — 
1.6  mm.  X  •5—63  mm.;  seta  8-20  mm.  long;  spores  11-16//, 
slightly  granulate-papillate;  elaters  1 10-200 /v.  x  9-14/^,  somewhat 
obtuse. 

On  a  charred  stump  of  Sequoia  sempcrvirois,  Eureka,  May  31, 
1896  (no.  938).  The  species  occurs  also  on  moist  soil  and  rocks. 
Our  single  Californian  specimen  bears  a  few  perigynia  and  dehisced 
capsules,  but  we  have  not  observed  $  branches  upon  it  with  any 
certaint}'. 

26.  CEPHALOZIA  Dumort.  /.  /.  max.  Rec.  d'Obs.  Jung.  18. 

1835- 

Jungcnnannia  §  CcpJialozia  Dumort.  p.  p.  max.  Syll.  Jung.  60. 
1831. 


122 


JUNGERMAXNIACEAE 


Noivdlia  Mitt.;  Godman,  Nat.   Hist.   Acores,   321.  1870. 
Schiffn.;  Eng.  &  Prantl,  Nat.  Pflanzenfam.  i"^ :  97.  1895. 
Eiiccplialozia  Schiffn.  /.  c. 

Prinolobiis  Schiffn.  /.  Eng.  &  Prantl,  Nat.  Pflanzenfam.  i'^ :  98- 
1895. 

CcpJialoziclla  Schiffn.  /.  c. 

Green,  whitish,  brown,  olivaceous,  or  sometimes  tinged  with 
rose-color,  small,  medium-sized,  or  minute,  depressed-caespitose 
or  creeping  among  mosses.  Stems  leafy  throughout  or  rhizo- 
matous  at  base,  or,  in  some  species,  flagelliferous,  subramose,  the 
branches,  with  rare  exceptions,  arising  ventrally,  root-hairs  nu- 
merous or  scanty.  Gemmae  occurring  in  a  few  species,  borne 
on  stem-apex  or  leaf-margins.  Leaves  alternate,  succubous 
and  sometimes  almost  horizontal,  more  rarely  nearly  transv^erse, 
rather  broad,  more  or  less  concave,  bifid  from  one  third  to  more 
than  one  half  their  length,  often  subcomplicate,  the  lobes  tri- 
angular-ovate, lanceolate,  or  subulate,  sometimes  connivent,  the 
margins  plane  or  slightly  incurved,  never  recurved,  entire  in  most, 
in  a  very  few  denticulate  or  sharply  serrate-dentate  ;  leaf-cells  from 
rather  large  and  pellucid  to  minute  and  subopaque,  the  walls  thin 
in  most  of  the  species,  thickened  in  a  few,  especially  toward  the 
angles,  cuticle  smooth  or  somewhat  roughened.  Underleaves 
much  smaller  than  the  lateral  and  often  entire,  wholly  wanting  in 
most  species  except  in  the  9  involucre  where  they  are  always 
represented  and  often  nearly  or  quite  equal  the  bracts  in  size. 
Dioicous,  autoicous,  or  polyoicous,  very  rarely  paroicous.  An- 
droecium  subamentiform,  median  or  more  rarely  terminal,  some- 
times occupying  an  entire  branch  ;  perigonial  leaves  often  a  little 
larger  or  smaller  than  the  ordinary,  more  concave  and  more 
crowded,  sometimes  with  a  small  accessory  incurved  lobule  on  the 
dorsal  margin  ;  antheridia  solitary.  Archegonia  several  or  nu- 
merous, terminal  on  a  much  shortened  or  sometimes  elongated 
branch  or  less  commonly  on  the  main  stem.  Bracts  free  from  the 
perianth,  crowded,  considerably  larger  than  the  leaves,  3 -ranked 
(including  bracteoles),  usually  in  three  sets,  2-4-lobed,  often  den- 
tate or  incised  ;  inmost  bracteole  always  present,  similar  to  the  cor- 
responding bracts  and  often  more  or  less  highly  connate  with  them 
on  one  or  both  sides,  the  outer  bracteoles  commonly  somewhat 
smaller,  rarely  wanting.  Perianth  more  or  less  highly  emersed, 
2-7  times  longer  than  broad,  distinctly  trigonous,  at  least  when 
young,  with  one  angle  always  ventral,  or  sometimes  4-6-angled, 
its  wall  of  one  layer  of  cells  or,  in  some  species,  2-  or  3-  stratose 
toward  the  base,  commonly  constricted  at  the  denticulate,  ciliate, 
laciniate,  or  subentire  mouth.     Calyptra  small,  thin,  or  rarely 


Cephalozia 


123 


fleshy,  surrounded  at  the  base  by  the  sterile  archegonia.  Capsule 
long-exserted  on  a  very  delicate  pellucid  seta,  cylindrical-oblong, 
or  oblong-globose,  quadrivalvular  to  the  base,  the  walls  bistratose, 
cells  of  the  inner  layer  with  semiannular  thickenings.  Elaters 
bispiral,  deciduous,  subobtuse,  mostly  150-200  n  long  and  8-10// 
wide.    Spores  smooth  or  minutely  roughened,  about  8-10  /i. 

Key  to  tlie  Species. 

Leaf- cells  30-60  //. 

Leaves  bifid  to  the  middle  or  a  little  below,  longer  than  broad  ;  perianth-wall  uni- 
stratose  unless  at  extreme  base.  i.  C.  bicuspidata. 

Leaves  bifid  about  V^,  their  length,  commonly  as  broad  as  long  or  broader  ;  peri- 
anth-wall of  2  or  3  layers  of  cells  for  Y^-Vi  its  length. 
Dioicous  ;  leaves  .  18-4  mm.  in  diameter,  decurrent,  nearly  horizontal  or 

subassurgent.  3.  C.  media. 

Monoicous  (polyoicous  ?)  ;  leaves  .3-.55  mm.  in  diameter,  scarcely  or  not  at 
all  decurrent,  assurgent  or  vertical-connivent.  2.  C.  pleniceps. 

Leaf-cells  10-20  11. 

Ramification  ventral  ;  leaves  distant  or  subimbricate,  their  lobes  usually  very  entire 
and  often  divergent.  4.  C.  divaricata. 

Ramification  commonly  lateral ;  leaves  mostly  approximate,  often  crowded  and  equi- 
tant,  complicate-bilobed,  the  margins  sharply  and  unequally  serrate-dentate. 

5.  C.  Turneri. 

I.  Cephalozla  bicuspidata  (L.)  Dumort.  Rec.  d'Obs.  Jung.  i8. 

1835- 

Jiingennamiia  bicuspidata  L.  Sp.  PI.  1132.  1753.  Hook. 
Brit.  Jung.       //.     18 16. 

Jiingennaiinia  Lauinicrsiana  Hiiben.  Flora,  15  :  306.  1832. 

Jungcrmannia  extensa  Tayl.  Lond.  Jour.  Bot.  5  :  279.  1846. 
CcpJialozia  extensa  Spruce,  On  Cephalozia,  44.  1882. 

Eticephalozia  bicuspidata  Schiffn.  ;  Eng.  &  Prantl,  Nat.  Pflan- 
zenfam.  i'^ :  97.  1895. 

Greenish  or  pallid,  more  rarely  rufescent  or  tinged  with  rose- 
color,  prostrate  or  assurgent-caespitose,  often  flagelliferous  :  stems 
rather  sparingly  branched,  4-20  mm.  long,  nearly  terete,  6  or  7 
cells  in  diameter,  the  cortical  cells  large  and  pellucid,  the  interior 
much  narrower  :  leaves  orbicular-ovate,  sometimes  oblong-ovate, 
mostly  .3-65  mm.  long,  obliquely  attached  or  subtransverse,  the 
inferior  distant  or  approximate,  the  superior  commonly  large  and 
subimbricate  or  crowded,  all  bifid  \-\  their  length,  concave,  often 
subcomplicate,  the  sinus  obtuse,  lobes  ovate-lanceolate,  triangular- 
lanceolate,  or  subtriangular,  connivent  or  patent,  subacuminate  or 
the  ventral  merely  acute,  sometimes — especially  the  dorsal — at- 
tenuate to  an  apiculus  2-4  cells  in  length  ;  leaf-cells  rather  large, 


124 


JUXGERMANNIACEAE 


thin-walled,  pellucid,  somewhat  thick,  quadrate-hexagonal,  be- 
coming elongated  toward  base,  30-60  // ;  underleaves  usually 
wanting  except  in  association  with  the  9  bracts  :  polyoicous  :  an- 
droecium  median,  terminal,  or  occupying  an  entire  branch,  the 
perigonial  leaves  much  like  the  others  but  more  concave,  usually 
closer,  and  sometimes  with  a  small  accessory  tooth  on  the  dorsal 
margin  near  the  base  :  archegonia  terminal  on  a  very  short  or 
much  elongated  branch  :  bracts  about  three  pairs  ;  the  inmost  2—3 
times  longer  than  the  leaves,  sometimes  slightly  connate  at  the 
base,  bifid  to  the  middle,  the  lobes  mostly  lanceolate-acuminate, 
entire,  repand,  or  w^ith  one  or  two  teeth  near  the  base  ;  bracteole 
similar  :  perianth  linear-subfusiform,  thrice  longer  than  the  bracts, 
1.8-2.5  mm.  X  .5  mrfi.,  at  first  tricarinate-prismatic,  later  often 
becoming  nearly  terete  below,  remaining  trigonous  toward  the 
constricted  denticulate  or  setulose  mouth,  its  wall  unistratose 
throughout  unless  at  the  extreme  base  :  calyptra  thin  :  capsule 
cylindrical-oblong,  .6-. 7 5  mm.  x  .3— .4  mm.;  spores  purple. 

On  decaying  wood,  moist  soil,  and  in  swamps  among  mosses. 

Mendocino  (Dr.  Bolander ;  Howe,  719)  and  North  Fork  of  the 
Little  River  (610,  615,  652,  659,  661,  689),  Mendocino  Co.; 
near  Eureka  (916,  963,  103  i). 

An  examination  of  Jiiugcnnannia  cxtensa  Tayl.  from  "  Observa- 
tory Inlet,  N.  America,  Dr.  Scouler,  1834'/  in  the  Taylor  collec- 
tion, now  incorporated  in  the  Gray  Herbarium,  Cambridge,  Mass., 
shows,  we  think,  no  satisfactory  and  reliable  grounds  for  its 
separation  from  C.  biciispidata.  Spruce  (On  Cephalozia,  45)  empha- 
sizes only  the  cladogenous  perichaetia  "  as  distinguishing  it 
from  C.  Laimnersiana,  but  we  find  the  9  branches  in  the  original 
material  often  much  elongated  as  in  C.  Lainincrsiaiia  ;  flagella  are 
not  wholly  wanting  ;  the  leaves  are  commonly  bifid  to  below  the 
middle,  but  they  can  be  matched  in  this  respect  by  European 
specimens  as  also  in  respect  to  the  apiculus  of  the  leaf-lobes. 
Most  of  the  Californian  specimens  are  dioicous  and  agree  closely 
with  the  original  of  Ta)'lor's  Jiingerinannia  cxtensa. 

The  specimens  which  v/e  once  reported  (Erythea,  4  :  50. 
1896)  from  California  under  the  name  Cephalozia  biciispidata  be- 
long with  C.  media  but  have  somewhat  thinner  perianths  than 
is  normal  with  that  species. 


Cephalozia 


125 


2.  Cephalozia  plexiceps  (Aust.)  Lindb.  Medd.  Soc.  Faun,  et  Fl. 
Fenn.  9  :  158.  1883. 

Jiingcnnannia  plciiiccps  Aust.  Proc.  Acad.  Nat.  Sci.  Philad. 
1869  :  222.  1869. 

Ccplialozia  crassiflora  Spruce,  On  Cephalozia,  40.  1882. 

Pale  green,  sometimes  lightly  tinged  with  brown,  densely  caes- 
pitose,  now  and  then  suberect  and  forming  compact  cushions  : 
stem  subramose,  rather  stout,  about  8  cells  in  diameter,  flattened 
dorsally,  the  cortical  cells  large  and  pellucid,  branches  occasion- 
ally stoloniform  :  leaves  obliquely  orbicular,  mostly  .3-.  5  5  mm.  in 
diameter,  strongly  concave,  assurgent,  often  vertical-connivent, 
subimbricate,  more  rarely  distant,  scarcely  or  not  at  all  decurrent, 
bifid  about  one  third  their  length,  the  sinus  obtuse,  less  commonly 
acute,  the  lobes  acute  or  the  ventral  occasionally  subobtuse,  often 
more  or  less  connivent  ;  leaf-cells  quadrate-hexagonal,  becoming 
oblong-hexagonal  toward  the  base,  rather  large  and  thick,  pellucid, 
thin-walled,  30— 60/i:  rudimentary  underleaves  rarely  present 
toward  the  apex  of  the  younger  branches  :  monoicous  (polyoic- 
ous  ?):  androecium  median  or  terminal,  dorsal  lobe  of  perigonial 
leaves  commonly  with  an  incurved  tooth  or  lobule  near  its  base  : 
9  branch  very  short  or  somewhat  elongated  ;  the  inmost  bracts 
bi-  to  quadri-fid,  distinct  or  slightly  connate  with  each  other  and 
with  the  bracteole,  often  bistratose  at  the  base,  the  segments  acute; 
bracteole  usually  bi-tri-fid ;  perianth  cylindrical-oblong,  2.3- 
2.8  mm.  X  .6-9  mm.,  contracted  at  the  plicate,  denticulate  mouth, 
carnose,  of  three  layers  of  cells  at  the  base  and  bistratose  to  the 
middle  or  even  for  two  thirds  its  length  ;  calyptra  mostly  unistra- 
tose. 

On  the  ground  beside  a  stream,  near  Sisson,  Siskiyou  Co., 
growing  among  Musci  {Aidacoinniujn,  etc.)  and  forming  a  com- 
pact turf. 

The  type  specimen,  from  the  White  Mountains  of  New  Hamp- 
shire (legit  Oakes)  is  in  the  possession  of  Mr.  W.  H.  Pearson. 

The  Californian  plant  has  a  rather  narrower  and  more  fleshy  per- 
ianth than  the  original,  the  wall  of  this  organ  being  in  the  former 
bistratose  for  two  thirds  its  length,  in  the  latter  only  about  to  the 
middle.  Cephalozia  crassiflora  Spruce — from  the  Pyrenees — an 
authentic  specimen  of  which,  also,  we  have  seen  in  herb.  Pearson, 
seems  to  differ  from  the  original  C.  plcniceps  in  scarcely  anything 
but  in  having  the  perianth  widest  above  the  middle,  while  in  both 
the  New  Hampshire  and  California  plants  the  perianth  is  com- 


126 


JUXGERMANNIACEAE 


monly  widest  at  the  middle  or  a  little  below.  We  are  of  the 
opinion,  first  ventured  as  a  suspicion  by  Dr.  Spruce  (On  Cepha- 
lozia,  41)  and  afterwards  confirmed  by  Mr.  Pearson  (List  of 
Canadian  Hepaticae,  9)  that  Cephalozia  crassiflora  is  a  synonym 
oi  C.  pleniceps. 

We  have  here  some  grounds  for  believing  that  our  Californian 
specimen  is  dioicous  as  well  as  monoicous  but  the  branches  of  a 
Cephalozia  can  be  so  easily  disarticulated  in  manipulation  that 
we  dare  not  assert  this  with  positiveness. 

Cephalozia  pleniceps  can  be  distinguished  from  its  near  ally  C. 
media  by  the  twice  larger,  scarcely  or  not  at  all  decurrent,  more 
nearly  orbicular  and  less  rhomboidal  leaves,  which  incline  to  be 
vertical-connivent  instead  of  horizontal  or  subassurgent,  by  the 
broader,  merely  acute  lobes,  the  thinner  calyptra,  the  rather 
more  fleshy  perianth,  etc.  This  species  seems  always  to  be  found 
in  company  with  mosses  in  swamps,  beside  streams,  and  on  moist 
rocks,  while  C  media,  in  California  at  least,  affects  logs  and  decay- 
ing wood  almost  exclusively. 

Plate  104.    Cephalozia  pleniceps. 

1.  Portion  of  plant  showing  perianth  and  androecium,  X  ^2. 

2.  Portion  of  a  sterile  stem,  with  leaves,  X  23. 

3.  9  branch  showing  perianth  and  bracts,  dorsal  view,  X  '2. 

4.  The  same,  ventral  view,  X  ^2. 

5.  Another  9  branch,  with  perianth  and  bracts,  dorsal  view,  X  ^2. 

6.  9  branch,  lateral  view,  the  perianth  undeveloped,  X  23. 

7.  A  thin  longitudinal  microtome  section  of  perianth,  X  23. 
8-1 1.  Cauline  leaves,  showing  the  outer  (convex)  face,  X  23. 
12-14.  Bracts,  X  23. 

15.  Bracteole,  X  23. 

16.  Second  (next  to  the  inmost)  bracteole,  X  23. 

17.  Portion  of  perianth  mouth,  X  4^- 

18.  Cross-section  of  stem,  X  53- 

19-21.  Bases  of  bracts  with  adjacent  parts  of  stem,  in  sectional  view,  X  4^- 
22.  A  cauline  leaf,  X  4i- 

Figs.  2-4  and  12-17  from  the  original  specimen  collected  in  the  White  Mts.  of 
New  Hampshire  by  Mr  Oakes,  ex  herb.  Pearson  ;  the  remaining  figures  from  our  speci- 
men collected  at  Sis.son. 

3.  Cephalozia  media  Lindb.  Medd.  Soc.  Faun,  et  Fl.  Fenn.  6 : 

242.  1881. 

C.  midtiflora  Spruce,  On  Cephalozia,  37.  1882. 
Eucephalozia  media  Schiffn.;  Eng.  &  Prantl,  Nat.  Pflanzenfam. 
97.  1895. 


Cephalozia 


127 


Slender,  bright-  or  pallid-green,  prostrate,  forming  close  de- 
pressed mats  or  rarely  laxly  creeping,  rather  sparingly  and  irregu- 
larly ramose,  the  apices  often  ascending  :  stem  somewhat  flattened 
dorsally,  the  cortical  cells  large  and  pellucid  :  leaves  small,  orbicu- 
lar-rhomboidal,  .  i  S-.4  mm.  in  diameter,  subimbricate  and  somewhat 
assurgent  or  often  distant  and  nearly  horizontal,  decurrent  dorsally, 
bifid  about  one  third  their  length  with  an  obtuse  or  rarely  lunate 
sinus  and  conniv^ent,  acute  or  subacuminate  lobes  ;  leaf-cells  pellucid, 
30—45  a  :  dioicous  :  androecia  occupying  the  end,  rarely  the  mid- 
dle of  mostly  rather  short  branches  :  inmost  bracts  of  the  very 
short  9  branch  about  three  times  larger  than  the  outer,  orbicular- 
oblong,  bi-quadri-fid,  the  segments  acute  or  acuminate,  entire  ; 
bracteole  nearly  similar,  sometimes  connate  with  the  bracts  :  per- 
ianth oblong-  to  linear-fusiform,  1.5-2.4  mm.  x  .4-8  mm.,  con- 
stricted and  denticulate  at  the  mouth,  of  2  or  3  layers  of  cells 
toward  the  base,  sometimes  2-stratose  as  high  as  the  middle,  the 
remainder  unistratose  ;  calyptra  2-  or  3-stratose  throughout :  cap- 
sule cylindrical-oblong,  .6-8  mm.  x  .28-.  5  mm.;  spores  cinnamon 
colored. 

On  decaying  logs  in  moist  shaded  places,  rarely  on  the  ground. 
Common  in  the  Coast  Range  Mountains,  north  of  San  Francisco. 

Redwood  Canon  near  Mill  Valley,  and  near  Olema  (28),  Marin 
Co.;  Duncan's  Mills  and  Turner's  Canon,  Sonoma  Co.;  Navarro 
(Miss  Byxbee),  North  Fork  of  the  Little  River  (656),  Mendocino 
(572,  573,  5-98,  665,  674),  Half-Way  House  (725),  Mendocino 
Co.;  Eureka  (904,  949),  Humboldt  Co.;  near  Hay  Fork,  Trinity 
Co.  (11 24);  Sisson,  Siskiyou  Co.  Collected  in  Marin  Co.  also 
by  Professor  Underwood  and  said  to  have  been  found  in  California 
also  by  Dr.  Bolander. 

The  perianth  in  our  specimens  is  usually  of  a  thinner  texture 
than  is  ascribed  to  CepJialozia  inultiflora  by  Dr.  Spruce.  It  is 
rarely  that  the  perianth- wall  is  2-stratose  so  far  as  the  middle.  In 
most  cases  the  perianth-wall  is  of  more  than  one  layer  of  cells  only 
in  the  lower  third  or  fourth,  sometimes  even  only  in  the  basal 
sixth,  yet  we  have  been  able  to  discover  no  other  character  to 
distinguish  the  Californian  plant  from  the  European  and  an  exam- 
ination of  European  specimens  seems  to  indicate  that  this  character 
is  susceptible  to  some  variation  even  there. 

4.   Cephalozia    divaricata  (Sm.)  Dumort.   Hep.   Europ.  89. 

1874. 

Jungermannia  divaricata  Sm.  Eng.  Bot. //.  7/p.  1800. 


128 


J  UNGERM  ANN  I  AC  E  AE 


Jungennannia  Starkii  Funck  ;  Nees,  Naturgesch.  Eur,  Leberm. 
2  :  215,  225.  1836. 

CepJialozidla  divaricata  Schiffn.  ;  Eng.  &  Prantl,  Nat.  Pflanzen- 
fam.      :  99.  1895. 

Very  small,  green,  olivaceous,  or  nearly  black,  sometimes 
tinged  with  rose-color,  closely  caespitose  or  laxly  creeping  among 
mosses,  prostrate  or  ascending,  eflagelliferous  ;  stem  stout  for  the 
size  of  the  plant,  2-10  mm.  long,  sparingly  branched,  about  7  or 
8  cells  in  thickness,  the  cortical  in  14-22  series,  not  different  from 
the  interior  or  a  little  smaller,  root-hairs  often  rather  abundant : 
leaves  small,  scarcely  longer  than  the  diameter  of  the  stem  or 
twice  as  long,  subquadrate  or  rotund-quadrate,  .08-18  mm.  in 
length,  .08-.  2  mm.  in  width,  distant  or  subimbricate,  nearly  trans- 
verse, bifid  to  the  middle  or  a  little  below,  the  lobes  triangular- 
ovate,  acute,  subacuminate,  or  nearly  obtuse,  divergent  orsubcom- 
plicate,  entire,  slightly  repand,  or  now  and  then  with  a  single  small 
tooth  on  one  side  or  the  other  ;  leaf-cells  subquadrate,  minute, 
10-16  fi,  subopaque  or  pellucid,  the  walls  thin  or  somewhat  thick- 
ened especially  at  the  angles,  cuticle  smooth  or  minutely  granu- 
late ;  stem-apex  and  margins  of  the  apical  leaves  sometimes 
gemmiferous :  underleaves  wanting,  or  present  in  the  younger 
parts,  entire  and  subulate  or  lanceolate,  or  sometimes  broader  and 
emarginate  or  bifid  :  dioicous  :  androecium  terminal  or  median, 
cauline  or  rameal,  the  perigonial  leaves  in  several  pairs,  concave, 
appressed-imbricate,  commonly  a  little  larger  than  the  ordinary 
leaves,  less  deeply  and  more  acutely  lobed  :  archegonia  terminal  on 
the  main  stem  or  on  an  elongated  branch  :  bracts  in  three  pairs,  2 
or  3  times  larger  than  the  leaves,  bilobed  or  rarely  trilobed,  usually 
distinct  dorsally  but  commonly  highly  connate  with  the  bracteole, 
the  lobes  acute  or  acuminate,  denticulate,  subspinulose,  or  nearly 
entire,  often  hyaline  at  the  apex  ;  bracteole  similar,  outer  bracteoles 
two,  somewhat  smaller  than  the  corresponding  bracts  and  less 
deeply  cleft ;  perianth  fusiform-ovoid  to  linear,  .8-1.4  mm.  X.2-.4 
mm.,  prismatic,  3-6-angled,  mostly  hyaline  toward  the  apex  and 
sometimes  purplish  below,  somewhat  contracted  at  the  denticulate 
or  subentire  mouth,  the  wall  unistratose. 

On  logs,  charred  stumps,  rocks,  and  moist  soil,  commonly  asso- 
ciated with  mosses  and  other  hepatics. 

Frequent  in  the  Coast  Range  region.  Clarendon  Heights, 
San  Francisco  ;  Mill  Valley,  Mt.  Tamalpais,  and  Lake  Lagunitas, 
Marin  Co.;  Cazadero,  Sonoma  Co.;  Mendocino  (544,  619,  669), 
North  Fork  of  the  Little  River,  and  Ukiah  (741),  Mendocino  Co.; 
Eureka  (956)  and  Blue  Lake  (995),  Humboldt  Co.;  Mt.  Diablo, 


Cephalozia 


129 


Contra  Costa  Co.;  Fruit  Vale,  Alameda  Co.  (Miss  Edith  S.  Byx- 
bee).    Also  gathered  by  Dr.  Bolander. 

In  the  Californian  specimens,  underleaves  can  nearly  always  be 
detected  toward  the  apex  of  the  younger  stems  or  branches. 

4/7.  Cephalozia  divaricata  scabra  var.  nov. 

Leaves  mostly  opaque,  often  of  two  or  three  layers  of  cells 
at  base,  especially  near  the  median  line,  the  back  near  the  fold 
usually  papillate,  verrucose,  spinulose-dentate,  subciliate,  almost 
lamelligerous,  or  subradicelliferous,  the  margins  now  and  then 
subdenticulate  or  'at  the  base  even  spinulose-dentate,  the  lobes 
sometimes  apiculate,  cuticle  of  the  leaf-cells  distinctly  roughened  : 
stems  exhibiting  rarely  here  and  there  minute  paraphyllia-like  ap- 
pendages, usually  composed  of  2  or  3  cells  in  a  lineal  series  : 
perianth-wall  often  bistratose  in  the  basal  fourth  or  fifth. 

On  rocks.  Near  Hay  Fork  (i  1 10),  Trinity  Co.;  Doolan's 
Canon  (776),  near  Ukiah,  Mendocino  Co.;  Cazadero,  Sonoma 
Co., 

This  variety,  especially  as  represented  in  the  first  two  specimens 
cited,  is  so  strikingly  different  from  the  normal  form  that  we  were 
at  one  time  on  the  point  of  treating  it  as  a  species,  which  we  were 
finally  deterred  from  doing  by  discovering  that  two  or  three  of 
our  Californian  specimens,  c.  g.,  those  cited  under  C.  divaricata 
from  Mt.  Diablo  and  (in  part)  from  Lake  Lagunitas,  seem  to  hold 
a  clearly  median  ground,  the  leaf  peculiarities  of  the  two  forms 
being  in  one  observed  case  even  exhibited  in  different  parts  of  a 
single  plant.  We  have  sometimes  seen  a  very  distinctly  roughened 
or  subpapillate  cuticle  in  European  specimens  of  C.  divaricata^ 
e.  g.,  Carr.  &  Pears.  Hep.  Brit.  261,  but  spinulose  teeth  or  even 
short  cilia  2  or  3  cells  long  from  the  back  of  the  leaf  we  have 
been  unable  to  discover  elsewhere  and  we  find  no  record  hitherto 
of  any  such  peculiarity  in  this  species  or  its  allies. 

5.  Cephalozia  Turxeri  (Hook.)  Lindb.  Acta  Soc.  Sci.  Fenn.  10: 

502.  1875. 

Jungerinannia  Turncri  Wook.  Brit.  Jung.//.  2g.     18 16. 
Anthelia  T?ir ner i  Dumort  Rec.  d'Obs.  Jung.  18.  1835. 
Prionolobtis  Ttirneri  ^^cYii^w.]  Eng.  &  Prantl.  Nat.  Pflanzenfam. 
i^:  98.  1895. 


130 


JUNGERMANNIACEAE 


Bright  green,  becoming  pallid  or  rusty-brown,  very  small,  de- 
pressed-caespitose  or  creeping  among  mosses ;  stems  2-6  mm. 
long,  terete,  6c- 100  a  in  diameter,  usually  ascend-ng  at  the  apex, 
subramose,  the  branches  lateral,  very  rarely  strictly  ventral,  cortical 
cells  subpellucid,  root-hairs  rather  few  :  leaves  mostly  approximate, 
often  crowded  and  equitant,  complicate-bilobed  ^—7^  their  length, 
the  margin  everywhere  sharply  and  unequally  serrate -dentate,  the 
lobes  ovate  or  ovate-lanceolate,  acute  or  apiculate,  the  dorsal  sub- 
erect  and  somewhat  parallel  to  the  stem,  the  slightly  broader 
ventral  usually  patent ;  leaf-cells  quadrate- hexagonal,  14-20  a,  the 
walls  strongly  thickened  especially  near  the  angles :  underleaves 
wanting  :  polyoicous  :  androecium  usually  median,  the  perigonial 
leaves  similar  to  the  others  but  sometimes  a  little  larger  and  more 
closely  imbricate :  archegonia  terminal  on  a  commonly  much 
elongated  branch  :  bracts  two  or  three  pairs,  increasing  in  size  up- 
ward, the  inmost  twice  the  size  of  the  leaves,  often  highly  connate 
with  each  other  and  with  the  bracteole,  spinulose-dentate,  bifid, 
the  lobes  acute  or  subacuminate ;  bracteole  ovate,  subacuminate, 
spinulose-dentate,  otherwise  usually  entire,  sometimes  connate 
with  only  one  of  the  bracts ;  outer  bracts  commonly  distinct,  outer 
bracteoles  wanting  or  only  one  present ;  perianth  linear-oblong,  i 
mm.  X  .25-35  mm.,  pentagonal-prismatic  with  pronounced 
carinae,  of  a  single  layer  of  cells  or  sometimes  bistratose  at  the  ex- 
treme base  or  at  the  angles,  often  decolorate  toward  the  rounded 
apex,  the  mouth  nearly  closed,  denticulate  or  very  obscurely  cilio- 
late  :  capsule  oblong-oval,  .35-.4  mm.  x  .22  mm. 

On  stony,  lightly  shaded  banks  and  on  sandy  soil,  sometimes 
associated  with  Claopodiuin  Icitconciiron  and  Catliarinca  iindulata 
or  with  Nardia  crcmdata,  rarely  on  logs  or  ascending  the  bases  of 
stumps  of  Sequoia  scmpervireiis. 

Marin  Co.  (Miss  M.  E.  Parsons,  May,  1888,  in  herb.  Under- 
wood), Mill  Valley  (Howe,  May  3,  1894,  Dec.  31,  1895,  Feb.  22, 
1896),  Redwood  Cafion  "  on  the  slopes  of  Mt.  Tamalpais  ;  Dun- 
can's Mills  and  near  Cazadero,  Sonoma  Co.;  Mendocino  (584, 
618),  North  Fork  of  the  Little  River  (68;),  and  Half-Way  House 
(732),  Mendocino  Co.;  near  Eureka  (974,  1030),  Humboldt  Co.; 
near  Lake  San  Andreas,  San  Mateo  Co.;  Twin  Oaks,  San  Diego 
Co.  (F.  W.  Koch,  Dec.  28,  1895). 

Exsicc.  Hep.  Am.  174. 

The  above  species,  first  described  from  Bantry,  Ireland,  and 
since  recorded  from  a  few  other  localities  in  Ireland,  England, 
France,  the  Canary  Islands,  and  northern  Morocco,  seems  to  be 


Kaxtia 


131 


more  common  in  the  coast  counties  of  California  than  in  the  Old 
World.  The  only  European  specimen  that  we  have  been  able  to  use 
for  comparison  is  one  collected  by  G.  Davies  in  Ashdown  Forest, 
Sussex,  England,  in  1881,  which  we  owe  to  the  generosity  of 
Matthew  B.  Slater,  Esq.,  of  Malton  ;  this  unfortunately  lacks  ma- 
ture perianths,  yet  the  correspondence  of  the  Californian  plants 
with  it,  so  far  as  it  goes,  and  with  the  detailed  description  of 
the  species  given  by  Dr.  Spruce  is  so  close  that  we  have  no 
ground  for  doubting  their  specific  identity.  A  slight  difference 
possibly  lies  in  the  more  frequent  occurrence  in  the  Californian 
specimens  of  concrescence  on  both  sides  between  bracteole  and 
bracts,  forming  a  complete  cup  with  an  erect  acutely  lobed  limb, 
yet  Spruce  says  of  the  specimens  collected  in  Ireland  by  Lindberg, 
"  bractae  intimac  ami  bracteola  biloba  in  excipiihnn  alte  connatae 
on  the  other  hand,  the  bracteole  in  our  plants  is  sometimes  free  on 
one  margin  and  the  bracts  may  even  be  found,  though  very  rarely, 
to  be  quite  distinct  dorsally. 

The  collection  of  this  species  in  San  Diego  County  by  Mr. 
Koch  suggests  that  it  probably  occurs  also  in  the  intervening  coast 
region. 

27.  KANTIA  S.  F.  Gray  (as  AW///^)  Nat.  Arr.  Brit.  PI.  i  :  706. 

1821. 

Calypogeja  Raddi  /.  min.  Atti  Soc.  Ital.  Sci.  Mod.  18  : — (31). 
1 818.    Mem.  Mat.  e  Fis.  Soc.  Ital.  Sci.  Mod.  18:  42.  1820. 

Cincinimliis  V)\xmox\,.  Comm.  Bot.  113.  1822. 

Calypogeia  Corda  ;  Opiz,  Beitr.  653.  i829.t 

Plants  medium -sized,  depressed-caespitose,  light  green,  some- 
times darkening  on  drying.  Stems  prostrate,  or  ascending  at  the 
often  gemmiparous  apices,  subsimple  or  with  irregularly  disposed 
branches,  these,  with  rare  exceptions,  arising  somewhat  laterally 
from  the  axils  of  the  underleaves  ;  root-hairs  very  long,  usually 
numerous,  springing  in  clusters  from  the  base  of  the  underleaves. 
Leaves  alternate,  incubous,  plane  or  slightly  convex,  oblong,  ovate, 
or  subrhomboidal,  rounded  or  retuse  at  the  apex,  less  commonly 
acute,  bidentate,  or  bilobed,  the  margins  entire;  leaf-cells  rather 
large,  chlorophyllose  or  pellucid,  rarely  with  trigones.  Under- 


*  Spruce,  On  Cephalozia,  72  1882. 
t  See  footnote,  page  33. 


132 


JUNGER  M  ANN  I ACEAE 


leaves  present  throughout,  usually  ^-^  the  size  of  the  leaves,  sub- 
orbicular,  or  sometimes  reniform  or  ovate,  the  apex  bifid,  retuse, 
or  entire.  Dioicous,  autoicous,  or  paroicous.  Sexual  branches 
arising,  singly,  in  pairs,  or  in  groups  of  three,  from  the  axils  of 
the  underleaves,  the  ^  branches  always  originating  near  the  basal 
margin  of  the  underleaf,  the  9  commonly  median.  Androecium 
(in  autoicous  and  dioicous  sp.)  small,  capitate  or  hnear-amenti- 
form  ,  ^  bracts  very  small,  bifid.  9  bracts  2  or  3  pairs  with 
similar  bracteoles,  much  smaller  than  the  leaves,  suborbicular, 
ovate,  or  lanceolate,  subentire  or  2-4-lobed  ;  archegonia  4-12. 
Perianth  wanting  ;  the  9  branch  developing  into  a  fleshy  pfendu- 
lous  subterranean  radicelliferous  cylindrical  or  clavate  pouch 
(perigynium),  crowned  at  the  mouth  with  the  persistent  scale-like 
bracts  and  lined  internally  with  clavate-papilliform  cells,  Calyp- 
tra,  at  maturity,  adnate  for  three  fourths  or  more  of  its  length  with 
the  perigynium,  and  almost  equally  long,  its  free  portion  sur- 
rounded by  the  abortive  archegonia.  Capsule  cylindrical,  dehis- 
cing to  the  base  by  linear  spirally  wound  valves,  these  bistratose, 
the  w^alls  of  the  outer  layer  of  cells  nearly  or  quite  destitute  of 
local  thickenings,  those  of  the  inner  layer  with  numerous  semi- 
annular  bands  ;  seta  long  ;  involucellum  highly  developed,  finally 
adnate  to  the  calyptra  and  reaching  nearly  to  the  line  of  departure 
of  calyptra  and  perigynium-wall,  its  cells  elongated  and  sinuous. 
Elaters  bispiral. 

I.  Kantia  trichomanis  (L.)  S.  F.  Gray,  /.  c. 

Mniiim  tricJiomanis  L.  Sp.  PI.  11 14.  1753. 

Jiingcrviannia  Tidchonianis  Dicks.  PI.  Crypt.  3:  10.  1793. 
Hook.  Brit.  Jung.//,  yg.  1816. 

Jungcninxunia  Sprcngclii  Mart.  Fl.  Crypt.  Erlang.  133.  pi-  3- 
f.  6.  1817. 

Cincinmdiis  TricJiomanis  Dumort.  /.  c. 

Calypogcia  TricJiomanis  Corda,  /.  c.  Nees,  Naturgesch.  Eur. 
Leberm.  3:8.  1838. 

Densely  depressed-caespitose  or  straggling  among  mosses  : 
stems  8-25  mm.  long,  .1-36  mm.  in  diameter,  simple  or  spar- 
ingly branched,  prostrate  and  terminating  obtusely,  or  often  atten- 
uate, ascending,  and  microphylline  at  the  apex  and  crowned  with 
a  subglobose  cluster  of  gemmae  :  leaves  usually  closely  imbricate, 
ovate,  .9-.  18  mm.  x  .63-1.4  mm.,  slightly  decurrent  ventrally,  be- 
coming, in  the  more  slender  conditions,  smaller,  merely  approxi- 
mate or  even  somewhat  distant,  subrhombic,  dimidiate-ovate,  and 
strongly  decurrent,  apex  in  all  commonly  rounded,  occasionally 


Kantia 


133 


retuse,  sometimes  acute,  rarely  bidentate  ;  superior  and  median  leaf- 
cells  35-65  /i,  the  chloroplasts  mostly  close  to  the  vertical  walls, 
trigones  wanting  or  indistinct ;  gemmae  ovoid,  of  one  or  two  cells  : 
underleaves  orbicular  to  broadly  ov^ate,  1-3  times  as  broad  as  the 
stem,  distant,  approximate,  or  imbricate,  bifid  to  below  the  middle 
or  merely  retuse,  the  lobes  acute  or  obtuse,  entire  or  rarely  with  a 
tooth  on  the  outer  margin  :  autoicous  (polyoicous  ? ):  androecium 
short  and  capitate  or  becoming  linear-amentiform  and  attaining  a 
length  of  I  mm.  or  more,  arising  from  the  same  axil  with  the  9 
branch  or  at  a  distance  from  it ;  antheridia  solitary,  ovoid,  about 
.  I  mm.  in  greatest  diameter,  short-stalked  :  archegonia  4-7  :  per- 
igynium  cylindrical,  1.5-2.5  mm,  x  .6-.9  mm.  :  capsule  1.5-2.5 
mm.  X  .5-5  mm.;  seta  1-2.5  cm.  long;  spores  12-16  u.,  very  mi- 
nutely punctulate  ;  elaters  180-350//  x  11-15      rather  obtuse. 

On  moist  banks  and  decaying  logs.  Common  in  the  Coast 
Range  Mountains.  Mill  Valley,  Redwood  Canon,  and  Olema, 
Marin  Co.  ;  Duncan's  Mills  {c.  fi\)  and  Cazadero  {c.fr.),  Sonoma 
Co.;  North  Fork  of  the  Little  River  (640,  660,  696)  and  Men- 
docino (583),  Mendocino  Co.;  Eureka  (903,  1221),  Humboldt  Co. 
Collected  also  by  Dr.  Bolander  at  Mendocino  and  in  "  Devil's 
Cafion,  Forest  Hill,  Sierra  Nevada,  no.  4620." 

The  Californian  specimens  belong  chiefly  to  what  is  sometimes 
treated  as  a  variety  or  species  under  the  name  Sprcngelii,  depart- 
ing from  the  form  ordinarily  considered  typical  in  the  smaller 
more  ovate  underleaves,  which  are  deeply  bifid,  with  the  lobes 
acute.  The  underleaves,  however,  often  show  such  extremes  of 
form  and  size  in  a  single  tuft  and  such  gradual  transitions  between 
these  extremes  that  we  are  convinced  that  any  attempt  to  div^ide 
this  group  into  two  species  according  to  the  character  of  the 
underleaves  is  both  unnatural  and  inconvenient.  While  the  Cali- 
fornian specimens  represent,  in  our  opinion,  but  a  single  species, 
the  determination  of  what  name  this  species  shall  bear  is  attended 
with  some  little  difficulty  owing  principally  to  the  various  ways 
in  which  Kantia  tricJiomanis  has  been  interpreted  and  defined  by 
European  authors.  Lindberg  *  has  made  the  relation  of  the 
antheridia  to  the  archegonia  a  ground  for  distinguishing  Kantia 
calypogeia  (Raddi)  Lindb.  from  Ka7itia  tricJiomanis,  describing  the 
former  as  autoicous  and  the  latter  as  paroicous,  and  Kaalaas  f  has 

*  Hep.  Utveck.  32.     1877.    Kongl.  Sv.  Vet.-Akad.  Handl.  23^:  25.  1889. 
t  Nyt  Mag.  for  Naturvidensk.  33:  201.  1893. 


134  JUNGERMANNIACEAE 

correlated  with  these  characters  others  drawn  chiefly  from  the 
underleaves  and  leaf-apices.  Nees  von  Esenbeck,*  however,  gives 
the  impression  that  his  Calypogeia  tricJiouianis  is  dioicous  and  the 
species  is  so  described  by  Limpricht  f  and  by  Stephani.  J  Gottsche  § 
on  the  other  hand,  twice  states  that  the  $  and  9  branches  may 
arise  side  by  side  from  the  axil  of  a  single  underleaf.  Spruce 
remarks  \\  "Our  Kantia  TricJwinanis  {Calypogia  Tr.,  Syn.  Hep.) 
is  also  certainly  monoicous,  and  fruits  freely  in  the  Castle- Howard 
woods  in  the  springtime.  The  flowers  stand  normally  three  to- 
gether, in  the  axil  of  a  foliole,  thus  ( ^  9  S)\  but  one  or  other 
flower  in  each  triplet  is  apt  to  be  obsolete,  thus  obscuring  the 
monoicous  character.  We  have  probably  a  second  species,  in 
bogs  and  wet  places,  with  dioicous  inflorescence,  and  large,  very 
slightly  notched  (or  even  quite  entire)  folioles  ;  but  I  have  seen 
no  fertile  specimen."  In  a  reprint  of  this  paper  On  Anomoclada") 
in  possession  of  Professor  Underwood,  Dr.  Spruce  has  written 
K.  TricJiomanis''  on  the  margin  opposite  the  words  "probably 
a  second  species"  and  "A".  Sprejigelii''  (Mart,)"  opposite  the 
reference  to  the  monoicous  species.  In  none  of  the  Californian 
specimens  (and  in  none  from  the  eastern  United  States,  so  far  as 
investigations  have  been  made)  have  we  been  able  to  demonstrate 
paroicous  inflorescence.  By  persistent  searching  we  have  always 
been  able  to  find  antheridia,  when  they  are  to  be  found  at  all, 
occupying  purely  $  branches  on  plants  which  bear  also  9  branches 
There  is,  however,  an  apparent  tendency  to  become  dioicous,  which 
is  especially  pronounced  in  plants  with  large,  merely  emarginate 
underleaves.  In  European  specimens,  also,  we  are  satisfied  that 
the  correlation,  as  defined  by  Kaalaas,  between  the  position  of  the 
antheridia  and  the  characters  of  the  underleaves,  leaf-apices,  etc., 
does  not  always  exist.  For  example,  an  autoicous  or  perhaps 
polyoicous  specimen  collected  in  Germany  by  Herr  F.  Stephani 
has  leaves  and  underleaves  which  are  very  decidedly  of  the  type 


*  Naturgesch.  Eur.  Leberm.  3  :  19.  1838. 
fCohn,  Krypt.-Fl.  Schles.  i  :  310.  1877. 
X  Berichte  Bot.  Ver.  Landshut,  7:  143  (51).  1879. 

^  Acta    Acad.     Caes.    Leop. -Carol.  Nat.   Cur.   2i'2 :   427.    1845.  Abhandl. 
Naturw.  Ver.  Hamb.  7  :  41.  1 880. 
^  Jour.  }3ot.  14  :  164.  1876. 


Kantia 


135 


conceived  by  Kaalaas  to  be  peculiar  to  Kantia  trichouianis.  It  should, 
perhaps,  be  remarked  that  in  European,  as  in  American,  specimens 
of  Kantia  trichonianis,  we  have  as  yet  been  unable  to  discover  anther- 
idia  in  the  axils  of  the  9  bracts — doubtless  due  to  the  fact  that  most 
of  the  European  specimens  of  Kantia  trichonianis  accessible  to  us  are 
in  an  unfavorable  condition  for  the  demonstration  of  antheridia. 
To  what  extent  paroicous  inflorescence  prevails  in  the  European 
plants  and  in  how  far  and  how  constantly  it  may  be  associated 
with  other  characters  are  matters  which  we  hope  will  soon  be  ex- 
haustively investigated  by  European  hepaticologists.  Inasmuch 
as  Linnaeus'  specimen  of  Mninni  tricJiomanis,  if  he  ever  had  one, 
is  not  preserved,  it  would  appear  that  the  only  way  of  interpreting 
the  diagnosis,  Mniiun  foliis  disticJiis  intcgcrrimis''  is  by  calling  to 
our  assistance  the  only  synonym  that  he  quotes,  viz.:  '^Mninni 
trichomanis  facie,  foliolis  integris.  Dill.  muse.  236.  t.  31.  f.  5." 
This  Dillenian  plant  Lindberg  has  declared  ^  to  be  Kantia  caly- 
pogea  (Dood.;  Radd.)  Lindb.  saccifera. — Haec  species  est  autoica, 
K.  trichonianis  (L.)  Lindb.  tamen  paroica."  On  this  showing  it 
seems  a  little  strange  that  Lindberg  should  not  have  pinned  the 
name  trichonianis  (L.)  "  down  to  the  autoicous  form  and  given 
his  paroicous  species  a  new  name.  It  is,  however,  to  be  noted 
that  this  determination  is  out  of  harmony  in  one  or  two  particulars 
with  his  earlier  determinations  of  the  plants  preserved  in  the  Dil- 
lenian herbarium,  for  he  once  wrote  :t  "  Secundum  specimina,  in 
herbario  ejus  nunc  asserv^ata,  Mninni  Dillen.  Hist.  muse.  tab.  31, 
figg.  et  5  et  6  ad  K.  trichonianis  a  2  fissani  [afterwards  K.  calypogeia 
(Raddi)  Lindb.]  pertinent !  Sed  (fig.  5)  caespes  componitur  et  ab 
hac  forma  (dioica),  $  etsacculos  juveniles  gerente,  et  ab  intermixta 
K.  argnta,  ster.  et  gonidiifera  !"  One  will  observe  that  the  sacci- 
ferous  plant  is  here  declared  to  be  dioicous  !  Whether  the  determ- 
ination of  three  years  later  was  a  rectification  based  upon  a  reexami- 
nation of  the  Dillenian  specimen  we  are  not  informed.  But 
whether  autoicous  or  dioicous,  it  certainly  appears  quite  unjustifiable 
to  limit  the  name    tricJionianis''  to  an  exclusively  paroicous  form. 

*Hep.  Utveck.  32.  1877. 

t  Not.  Sallsk.  pro  Fauna  et  Flora  Fenn.  Forhandl.  13:  364.  1874. 


136 


JUNGERMANNIACEAE 


28.  LEPIDOZIA*  Dumort.  Rec.  d'Obs.  Jung.  19.  1835.  G. 
L.  &  N.  Syn.  Hep.  200.  1845. 

Pleuroschisma  §  Lepidozia  Dumort.  Syll.  Jung.  69.  1831. 

Mastigophora  Nees,  Naturgesch.  Eur.  Leberm.  i  :  95,  10 1. 
1833- 

Hcrpcthnn  §  Lepidozia  Nees,  op.  cit.  3:31.  1838. 

Plants  small  or  moderately  large,  pallescent  or  yellowish-green, 
in  dense,  often  depressed  mats.  Stems  prostrate  or  ascending, 
rarely  erect  or  pendulous,  pinnately  or  bipinnately  branched,  often 
beautifully  plumose,  the  vegetative  branches  lateral  in  origin, 
rarely  ventral ;  root-hairs  rather  scanty,  mostly  from  the  ventral 
surface  of  the  underleaves.  Leaves  incubous.  small,  ventrally  con- 
cave, decurved,  unsymmetrical,  usually  as  broad  as  long,  palmately 
4-  (rarely  2-  or  3-,  or  5-  or  6-)  cleft  or  -parted,  the  segments 
lanceolate  or  subulate  and  (in  our  species)  entire  ;  leaf-cells  rather 
small  or  medium-sized.  Underleaves  present  throughout,  similar 
to  the  leaves  but  symmetrical  and  somewhat  smaller.  Dioicous 
or  autoicous.  Androecium  occupying  a  short  ventral  ramulus  or 
rarely  terminal  on  a  lateral  branch  ;  $  bracts  suborbicular,  saccate- 
concave,  larger-celled,  mostly  bilobed,  bracteoles  much  smaller  ; 
antheridia  single.  Archegonia  few  or  numerous,  terminal  on  a 
short  ventral,  often  strongly  radicelliferous,  branch.  9  bracts  5-5 
pairs,  the  inmost  mostly  three  times  larger  than  the  cauline  leaves, 
all  larger-celled  and  more  translucent,  2-4-lobed  or  -dentate  at 
apex,  the  margins  entire,  denticulate,  or  spinose  :  bracteoles  simi- 
lar. Perianth  free,  cylindrical-ovoid  to  narrowly  fusiform,  more 
or  less  distinctly  trigonous  above,  with  the  third  angle  ventral, 
gradually  contracted  at  the  apex  toward  the  rather  small  denticu- 
late or  ciliate-laciniate  mouth.  Calyptra  as  long  as  the 
perianth.  Capsule  cylindrical -ovoid,  dehiscing  to  the  base  by 
straight  valves,  these  2-4-stratose,  the  walls  of  the  external  cells 
with  columnar  or  nodular  thickenings,  those  of  the  inmost  layer 
with  semiannular  bands  ;  seta  moderately  long.     Elaters  bispiral. 

*  In  order  to  retain  in  use  Lepidozia  as  a  generic  name,  we  recognize  its  subgeneric 
priority.  The  Maslis^ophora  Nees  of  1833  ^^'^^  rather  briefly  defined  and  no  species 
were  cited  under  it,  yet  in  1838  Nees  mentions  Afastigop/ioj-a  rcptans  as  a  synonym  of 
Herpetitan  reptans  and  refers  to  page  loi  of  the  first  volume  of  the  Eur.  Leberm.  as  the 
place  of  publication,  where,  however,  no  specific  names  are  to  be  found.  There  can, 
therefore,  be  no  possible  doubt  as  to  what  Nees  intended  to  have  understood  under  his 
Mastif^ophora  of  1833.  But  in  1 838  Nees  reverted  to  Dumortier's  sectional  name 
Lepidozia  (using  it  for  a  section  of  Herpetiuui,  and  without  any  allusion  to  Dumortier's 
work  of  1835,  in  which  Lepidozia  was  raised  to  generic  rank)  and  forthwith  transferred 
the  name  Mastigophora,  with  full  diagnosis,  io  a  very  different  plant  [Jungcnnannia 
VVoodsii  Hook. ). 


Lepidozia 


137 


I.  Lepidozia  reptaxs  (L.)  Dumort.  Rec.  d'Obs.  Jung.  19.  1835. 
Jtingcrmannia  rcptans      Sp.  PI.  1133.  1753. 
PlcnroscJiisma  rcptans  Dumort.  Syll.  Jung.  69.     1 83  I. 
HcrpctiiLVL  reptans  Nees,  Naturgesch.  Eur.  Leberm.  3:  31. 

1838. 

In  depressed  mats  or  creeping  among  mosses  :  stems  prostrate, 
6-25  mm.  long,  .18-36  mm.  in  diameter,  subpinnately  to  bipin- 
nately  ramose,  the  branches  ending  obtusely  or  often  becoming 
flagelliform  :  leaves  obliquely  inserted,  patent-horizontal  to  erecto- 
patent,  mostly  approximate  or  subimbricate,  rarely  distant,  sub- 
quadrate,  inclining  occasionall)^  to  ovate  or  obovate,  .36—45  mm. 
long,  .25-54  mm.  broad,  decurved,  3-  or  4-  (rarely  2-)  cleft  for 

Yl-Yz  their  length,  the  lobes  unequal,  mostly  lanceolate-acute  or 
subulate,  incurved,  4-8  cells  wide  at  base  ;  leaf-cells  24-48  a,  their 
firm  walls  obscurely  or  not  at  all  incrassate  at  the  angles,  cuticle 
smooth  :  underleaves  distant  or  contiguous,  subquadrate,  broader 
than  long,  -l-l-  the  size  of  the  leaves,  mostly  4-lobed  to  about  the 
middle,  the  lobes  usually  obtuse  :  autoicous  :  androecium  typically 
on  a  short  ventral  branch,  rarely  borne  on  a  branch  of  lateral 
origin  :  axis  of  9   branch  about  .  5  mm.  long,  archegonia  2-6  : 

9  bracts  2-3-dentate  at  apex,  margins  entire  or  slightly  denticu- 
late :  perianth  subcylindrical  or  narrowly  ovoid-conical,  2-2.7 
long,  .63-1  mm.  in  greatest  width  (below  the  middle),  rounded- 
trigonous  above,  the  wall  of  2  or  3  layers  of  cells  in  the  basal  half 
or  third,  otherwise  unistratose,  the  mouth  denticulate  :  capsule 
about  1.2  mm.  X.7  mm.;  seta  1-2  cm.  long;  spores  14-16  dis- 
tinctly granulate-papillate;  elaters  200-325  /^.x  10-16 rather 
obtuse. 

On  old  logs  in  moist  woods,  on  banks  of  streams,  among 
mosses,  etc.    Not  especially  common  in  California.    "  Redwood 
Cafion,"  near  Mill  Valley,  Marin  Co.;  near  Mendocino  (720)  and 
j"    North  Fork  of  the  Little  River  (658),  Mendocino  Co.;  Eureka 
I     (906,  929),  Humboldt  Co.;  near  Hay  Fork  (1120),  Trinity  Co.; 
I    Sisson  (36),  Siskiyou  Co.    Collected  at  Mendocino  also  by  Dr. 
Bolander  and  in  Marin  County  by  Professor  Underwood  (June, 
1888). 

Dr.  Spruce, in  describing  the  dioicous  Lepidozia  Pcarsoni,  in 
which  the  androecia  normally  terminate  lateral  branches,  has  stated 
that  in  Lepidozia  nptaiis  the  male  spike  always  occupies  a  postical 
branch.    The  Californian  specimens  of  Lepidozia  reptans,  however 


*Jour.  Bot.  19  :  35.  l88l. 


138 


J  UNG  ER  M  ANN  I ACEA  E 


— though  apparently  always  autoicous — occasionally  show  an 
antheridial  spike  at  the  end  of  a  lateral  branch,  and  we  have  ob- 
served the  same  conditions  in  certainly  autoicous  plants  of  G.  &  R. 
Hep.  Eur.  282. 

Lcpidozia Jilauientosa'^  (Lehm.  &  Lindenb.)  Lindenb. — a  very 
distinct  species,  now  certainly  known  only  from  Alaska — will  prob- 
ably yet  be  found  to  extend  farther  down  the  coast,  though  we  are 
inclined  to  doubt  its  occurrence  Avithin  the  limits  of  California. 
This  species  is  larger  in  all  its  parts  f  than  Lepidozia  rcptans  and 
more  rigid  and  brittle  in  drying.  The  main  stems  are  2-3  cm. 
long  and  .36-. 6 5  mm.  in  thickness.  The  leaves  are  almost  trans- 
versely inserted  and  erecto-patent  to  suberect ;  the  underleaves  are 
proportionally  larger  than  in  L.  reptans,  being  -|-|  the  size  of  the 
lateral  leaves  ;  the  fronds  have  therefore  the  appearance  of  being 
almost  uniformly  developed  on  all  sides  instead  of  being  distinctly 
dorso-ventrally  flattened.  Fig.  4  in  Lindenb.  &  Gottsche's  plates 
scarcely  does  justice  to  the  similarity  in  size  of  the  leaves  and  under- 
leaves ;  and  Taylor's  stipulis  minutis  "  of  ''Lepidozia  attcmiata  "  is 
difficult  to  understand  in  view  of  the  undoubtedly  authentic  original 
specimen  existing  in  his  herbarium,  which  is  an  exact  counterpart  of 
the  original  oi  Jiingcrmannia  filanientosa,  now  preserved  in  the  Nat- 
ural History  Museum  of  Vienna.  In  the  original  oi  L.  filaincntosa 
the  perianths  are  4-6  mm.  long  and  are  widest  at  the  middle  or  a 
little  above  ;  the  only  capsule  seen  is  2  mm.  long.  Menzies  was 
probably  the  first  collector,  though  his  name  is  not  cited  by  Linden- 
berg  and  does  not  appear  on  the  Vienna  specimen.  The  species 
has  since  been  collected,  so  far  as  we  know,  but  twice  :  by  J.  M. 
Macoun,  Prince  of  Wales  Sound,  Alaska,  September,  29,  1891 
(no.  389,  associated  with  Scapaiiia  Bolandcri,  in  herb.  Underwood), 
and  by  Miss  Jessie  Trowbridge,  Sumdum,  Alaska,  1895  (in  herb. 
Howe).  The  species  is  apparently  dioicous.  We  have  seen  peri- 
anths only  upon  the  original  specimen.  The  androecia  are  upon 
ventral  branches.   The  main  stems  and  lateral  branches  (especially 

^Lkpidozia  i  iLAMENTOSA  Lindenb.  (excluding  var.  G.  L.  &  N.  Syn.  Hep. 

206.     1845.    Lindenb.  cK:  Gottsche,  Sp.  Hep.  fasc.  6  &  7  :  36.  //.  6.  1846. 
Juni^ey7)tanitia  filamentosa  Lehm.  &  Lindenb.;  Lehm.  Pugill.  6:  29.  1834. 
Lepidozia  atteniiata  Tayl.  Lond.  Jour.  Bot,  5  :  369.  1846. 

t  With  the  exception  of  the  leaf-cells,  which  are  a  trifle  smaller  and  more  opaque 


Blepharostoma 


139 


in  9  plants  ?)  sometimes  terminate  in  a  somewhat  enlarged,  rather 
dense,  rosette  of  leaves. 

Lcpidozia  luiDiillima  Tayl.  (/.  c.)  Northwest  coast  of  America  ? 
Menzies  "  is  a  CcpJialozia. 

29.  BLEPHAROSTOMA  Dumort.  p.  p.  Rec.  d'Obs.  Jung.  18. 
1835.    Spruce,  On  Cephalozia,  86.  1882. 
Jimgcrniannia  §  Blepharostoma  Dumort.  /.  /.  Syll.  Jung.  65. 

1831. 

CJiactopsis  Mitt.  Jour.  Linn.  Soc.  8:  53.  1865. 

Blepharostoma  ^  Chaetopsis  Schiffn.;  Eng.  &  Prantl,  Nat. 
Pflanzenfam.      :  104,  105.  1895. 

Plants  small,  sometimes  very  slender  and  delicate.  Stems 
usually  prostrate-entangled,  the  branching  normally  lateral  or  sub- 
dichotomous.  Leaves  transversely  inserted,  divided  very  nearly 
or  quite  to  the  base  into  2-5  rigid  and  setaceous  or  soft  and  capil- 
lary, simple  or  less  commonly  furcate  laciniae,  each  consisting  of 
a  single  row  of  cells  throughout.  Underleaves  very  similar,  their 
laciniae  sometimes  fewer  by  one  and  occasionally  a  little  shorter. 
Archegonia  few,  terminal  on  the  main  stem  or  on  the  principal 
branches.  bracts  usually  more  crowded  than  the  leaves,  their 
laciniae  sometimes  more  numerous  and  often  forked  ;  antheridia 
ovoid,  solitary  or  very  rarely  binate.  5  bracts  gradually  larger 
than  the  leaves,  free  from  the  perianth,  distinct  or  the  inmost 
slightly  connate  at  the  base,  the  often  much  branched  laciniae 
springing  from  a  basilar  membrane  several  cells  in  height ;  brac- 
teoles  similar.  Perianth  subpyriform  to  elongate-cylindrical,  its 
wall  composed  of  a  single  layer  of  cells,  the  mouth  ciliate.  Cap- 
sule dehiscing  to  the  base  by  four  straight  valves.  Seta  rather 
short  or  moderately  long.    Elaters  bispiral,  obtuse. 

Key  to  the  Species. 

Leaves  mostly  with  3  or  4  laciniae,  their  cells  40-70  a  X  16-32  //,  1^-21^  times  as 
long  as  broad,  thick-walled,  the  septa  usually  strongly  thickened  toward  the  pe- 
riphery and  slightly  protuberant.  I.  B.  trichophyllum. 

Leaves  mostly  with  2  or  3  laciniae,  their  cells  50-112  a  X  25-50  ,u,  lj^-4  times  as  long 
as  broad,  thin- walled,  slightly  contracted  at  the  septa  (which  are  never  at  all  pro- 
tuberant). 2.  B.  aracJmoideum. 

I.  Blepharostoma  trichophyllum  (L.)  Dumort.    Rec.  d'Obs. 

Jung.  18.  1835. 
Jiingcnnannia  trichophylla  L.  Sp.  PL  1 1 3 5.  1753- 
Chaetopsis  trichophylla  Mitt.  Jour.  Linn.  Soc.  8:  53.  1865. 


140 


JUXGERMAXXIACEAE 


Green  or  yellowish-green,  intricately  caespitose  or  straggling 
among  other  bryophytes  :  stems  rather  rigid,  4-20  mm.  long,  .06- 
.27  mm,  in  diameter,  with  often  copious  lateral  or  dichotomous 
branching,  cortical  cells  subopaque,  7— 14-seriate,  28—96  /^XI5— 
32  n,  axile  cells  4-1 2-seriate  ;  root-hairs  infrequent:  leaves  close 
or  somewhat  widely  spaced,  erecto-patent  or  suberect,  divided 
nearly  to  the  base  into  3  or  4  rigid  setaceous  laciniae  (occasionally 
five  or  in  very  slender  conditions  rarely  reduced  to  two),  these  .4- 
.65  mm.  (8-13  cells)  long,  usually  connate  for  the  height 

of  the  basal  cells,  or  sometimes  with  a  basilar  membrane  one  cell 
in  height  ;  leaf-cells  40-70 /7.x  16-32  ja,  1^-2^  times  as  long  as 
broad,  thick-walled,  the  septa  usually  strongly  thickened  toward 
the  periphery  and  slightly  protuberant,  giving  the  laciniae  a  some- 
what nodulose  outline  ;  cuticle  granulate  or  striatulate  :  underleaves 
similar  to  the  lateral  .leaves,  their  laciniae  commonly  one  or  two 
cells  shorter  :  paroicous,  or  with  antheridia  on  the  same  axis  but  a 
little  removed  from  the  9  bracts,  or  sometimes  dioicous  :  $  bracts 
with  a  narrow  basal  membrane,  the  laciniae  usually  once  furcate  ; 
antheridia  about  .  1 5  mm.  in  diameter,  the  short  stalk  composed  of 
a  single  row  of  cells  :  archegonia  4—7  :  9  bracts  4— 6-laciniate 
with  a  basilar  membrane  2-6  cells  high,  the  laciniae  furcate  or  often 
with  an  antler-like  ramification  :  perianth  highly  exserted,  cylin- 
drical or  pyriform-cylindrical,  1.5-2.2  mm.  x. 45-8  mm.,  inflated, 
with  usually  one  or  two  deep  furrows  toward  the  apex,  the  rather 
wide  mouth  somewhat  abruptly  drawn  together  and  fringed  with 
long,  mostly  connivent  cilia  :  calyptra  thin,  free,  about  half  as  long 
as  the  perianth  :  capsule  ellipsoid-oval,  .6-.g  mm.  long,  the  valves 
bistratose,  the  cell-walls  of  the  outer  layer  with  purplish-brown 
columnar  thickenings,  those  of  the  inner  layer  with  similar  thicken- 
ings and  with  imperfect  semiannular  fibers  ;  seta  5-12  mm.  long  ; 
foot  deeply  penetrating  the  stem-apex,  descending  to  the  plane  of 
insertion  of  the  third  set  of  bracts  ;  spores  14-18  //,  minutely  granu- 
late-papillate ;  elaters  1 30-300  ii  x  9-1 1  H- 

On  logs  and  moist  soil. 

Fish  Creek,  Mariposa  Co.  (Miss  Edith  S.  Byxbee,  July,  1895) ; 
near  Mendocino  (668,  722);  Blue  Lake,  Humboldt  Co.  (1009). 

2.  Blepharostoma  arachnoideum  sp.  nov. 
Pallid  or  ycllowish-grccn,  depresscd-caespitose,  very  delicate, 
confervoid,  mostly  pulverulent-nitcnt  owing  to  the  presence  of 
gemmae  :  stems  flaccid,  2-5  mm.  long,  .05-. 22  mm.  in  diameter, 
simple  or  sparingly  branched,  the  branching  dichotomous,  lateral, 
or  sometimes  strictly  ventral,  cortical  cells  4-1  o-(mostly  6-  or  7-) 
seriate,  pellucid,  48-160//  x  25-60  //.,  the  dorsal  a  little  larger 


Blepharostoma 


141 


than  the  ventral,  axile  cells  i-6-seriate;  root-hairs  very  long,  color- 
less or  yellowish,  usually  wanting,  springing,  when  present,  singly 
or  in  groups  of  2  or  3  from  near  the  places  of  insertion  of  the 
underleaves  :  leaves  rather  widely  spaced,  patent  or  erecto-patent, 
divided  nearly  or  quite  to  the  base  into  2  or  3  capillary  laciniae 
(very  rarely  four,  or  in  attenuate  conditions  sometimes  reduced  to 
one),  the  laciniae  .36-75  mm.  (6-1 1  cells)  long,  often  once  forked 
at  about  the  middle  or  above,  discrete  to  the  very  base  or  more 
commonly  connate  for  the  height  of  the  basal  cells,  slightly 

contracted  at  the  septa;  leaf-cells  mostly  50-11 2 /y.  x  25-50 
I  times  as  long  as  broad,  hyaline,  thin-walled,  the  septa  some- 
times slightly  thickened  toward  the  periphery  but  never  at  all 
protuberant  ;  cuticle  (of  the  cortical  cells  as  well)  minutely 
striatulate  :  underleaves  very  similar  in  all  respects  to  the  lateral 
leaves  :  gemmae  frequent,  especially  toward  the  stem-apex,  uni- 
cellular, oblate -ellipsoidal,  25  a  in  greatest  diameter,  formed  by  the 
multiplication  in  a  single  series  and  reduction  in  length  of  the  ter- 
minal cells  of  the  leaf-segments,  the  resulting  submoniliform 
chains  composed  often  of  12-18  cells  :  remaining  parts  unknown. 

On  old  logs  in  moist  woods,  ''Russian  Gulch,"  near  Mendocino 
(703 — the  type)  and  North  Fork  of  the  Little  River  (1222), 
Mendocino  County. 

The  above  species  is  strikingly  distinct  from  Blepharostoma 
tncJiophylliLni.  Blepharostoma  sejunetum  Angst.,  from  Brazil, 
is  doubtless  a  nearer  ally,  judging  from  description  alone,  but 
the  Californian  plant  evidently  differs  from  this  in  the  shorter  and 
rather  stouter  stems,  in  having  often  3  leaf-laciniae,  the  basal  cells 
of  which  are  commonly  more  or  less  connate,  in  the  broader  leaf- 
cells,  and  probably  also  in  the  ramification.  In  habit  and  delicacy 
of  structure,  Blepharostoma  aracJinoideum  is  suggestive  of  Dr. 
Spruce's  genera  Telaranea  and  AracJiniopsis.  The  occasional  oc- 
curence of  a  postical  branch  is  an  aberrant  character  in  Blepharo- 
stoma, with  which  genus,  however,  the  equally  developed  three 
ranks  of  leaves  plainly  unite  our  plant.  The  species,  in  a  sense, 
connects  Blepharostoma  with  Telaranea,  which,  in  turn,  as  Spruce 
remarks,  is  with  difficulty  kept  separate  from  Lepidozia. 

-Ofvers.  Kongl.  Vetensk.-Akad.  Forhandl.  1876":  78.  1876.  The  width  of 
the  laciniae  is  here  given  as  "0,003  niiHim."  which  is  evidently  a  misprint  for  0,03 
millim. 


142 


J  UNG  E  R  M  ANN  I ACEAE 


30.  ANTHELIA  Dumort. />.  p.  Rec.  d'Obs.  Jung.  18.  1835. 

Schiffn.;  Eng.  &  Prantl,  Nat.  Pflanzenfam.  i'^:  106.  1895. 

J2ingc7'nia]uiia  §  AiitJiclia  Dumort. /. /.  Syll.  Jung.  63.  183 1. 

AntJielia  §  A.   Spruce,  On  Cephalozia,'  80.  1882. 

Plants  small,  densely  caespitose,  olivaceous-green  or  glauces- 
cent.'*'  Stems  stout  for  the  size  of  the  plant,  opaque,  irregularly 
pinnate,  the  branches  all  lateral,  cortical  cells  multiseriate,  small 
to  medium -sized  ;  root-hairs  few  or  abundant.  Leaves  small, 
transversely  inserted,  erect  or  erecto-patent,  somewhat  complicate- 
carinate,  commonly  equitant-imbricate,  bifid  to  the  middle  or  be- 
low, the  lobes  subacuminate,  several  cells  in  width  except  at  the 
extreme  apex,  the  margins  entire  or  erose-denticulate.  Under- 
leaves  similar  to  the  lateral  and  scarcely  smaller,  the  stems,  there- 
fore, appearing  uniformly  tristichous-foliate.  Dioicous  or  paroi- 
cous.  Antheridia  large,  solitary  in  the  axils  of  but  slightly 
modified  leaves.  Archegonia  few,  terminal  on  the  main  stem  or  the 
leading  branches.  9  bracts  gradually  larger  than  the  leaves, 
densely  crowded,  more  often  denticulate,  more  or  less  fused  with 
the  base  of  the  perianth  and  the  hollowed-out  stem-apex,  distinct  or 
rarely  somewhat  connate  at  base,  the  apices  commonly  hyaline. 
Perianth  ovoid  to  short-cylindrical,  deeply  unisulcate  dorsally  and 
2-  or  3-carinate  ventrally,  8-io-plicate  at  the  slightly  narrowed 
lobate-dentate  mOuth,  the  wall  of  2  or  3  layers  of  cells  near  the 
base,  otherwise  unistratose.  Calyptra  2  or  3  cells  thick,  carry- 
ing on  its  surface  the  sterile  archegonia  and  sometimes  also 
small  scale-like  paraphyses.  Capsule  subglobose,  dehiscing  to 
the  base  by  straight  valves,  which  are  composed  of  two  layers  of 
cells,  the  walls  of  the  outer  layer  with  columnar  or  nodular  thick- 
enings, those  of  the  inner  layer  similarly  marked  toward  the  base 
of  the  capsule  but  bearing  toward  the  apex  more  or  less  perfect 
semiannular  fibers  ;  seta  short.     Elaters  2-  or  3 -spiral. 

I.  Anthelia  Juratzkana  (Limpr.)  Trevis.  Mem.  r.  1st.  Lomb. 
III.  4:  416.     1877.    Spruce,  On  Cephalozia,  82.  1882. 
Jungcrinaiinia  Juratzkana  Limpr.;  Cohn,  Krypt.-Fl.  Schles.  I  : 
289.     1877.     Hedwigia,  15:  18.  1876. 

*  Herbarium  specimens  of  Anthelia  are  often  thickly  covered  with  a  delicate  white 
cobwebby  growth,  which  under  a  magnification  of  500  diameters  is  found  to  consist  of 
very  slender  short  and  rigid  or  long  and  flexuous  threads  about  i  a  in  diameter  and  of 
homogeneous  structure.  The  organism  is  evidently  one  of  the  colorless  schizophytes, 
but  further  than  this  we  would  not  attempt  to  express  an  opinion  as  to  its  systematic 
position.  Schizophyceae  and  mycelia  of  fungi  are  usually  also  present  in  greater  or  less 
abundance. 


Anthelia 


148 


i^^  Jiingeniiannia  nivalis  S\v.;  Wahlenb.  Fl.  Carp."^  363.  18 14. 
Wahlenb.  Fl.  Suec.  779.  1826. 

Anthelia  nivalis  Lindb.  Muse.  Scand.  5.  1879.  Kaalaas, 
Nyt  Mag.  for  Naturvidensk.  33  :  232.  1893. 

Plants  dark  green  or  glaucescent  at  the  apices,  brownish  below, 
forming  wide  dense  mats  2-4  mm.  in  thickness  :  primary  stems 
prostrate  or  decumbent,  the  principal  branches  erect,  .08-.  26  mm. 
in  diameter,  sparingly  radicelliferous ;  leaves  erect-appressed, 
usually  crowded,  more  distant  on  slender  sterile  stems,  ov^ate, 
.3-. 3 5  mm.  long,  bifid  their  length,  the  lobes  ovate-lanceo- 

late, subacuminate,  the  margins  entire  or  v^ery  slightly  erose-den- 
ticulate ;  leaf-cells  rectangular-oblong  to  subquadrate,  16-40  fi 
in  greatest  diameter,  rather  thin-walled  :  paroicous  :  bracts  sev- 
eral pairs,  a  little  saccate  at  base,  apices  denticulate,  bracteoles 
similar  :  perianth  scarcely  exserted,  ov^oid,  1-1.6  mm.  x  .6-. 7  mm., 
lobulate  at  the  somewhat  contracted  mouth  :  capsule  .^—.y  mm.  in 
diameter,  on  a  seta  1-3  mm.  long;  spores  16-24  maximum 
diameter,  granulate-papillate;  elaters  120-175  //.  long,  11  —  14 
greatest  diameter,  somewhat  attenuate  at  the  extremities,  mostly 
trispiral. 

On  rocks  in  alpine  regions.  ''California,  Bolander"  in  herb. 
U.  S.  National  Museum  (under  the  name  Anthelia  jnlacea).  It 
seems  probable  that  Dr.  Bolander's  specimen  was  collected  on  Mt. 
Dana  in  September,  1866. 

This  citation  is  as  given  by  authors.  We  have  been  unable  to  consult  this  earlier 
work  of  Wahlenberg's  and  have  also  failed  to  see  the  W.  jNI.  Ind.  muse."  in  which 
Jungcnnannia  Jiivalis  as  a  nonien  midutii  is  said  to  have  made  its  first  appearance. 
Though  we  have  not  been  able  to  see  all  the  literature  bearing  upon  the  point,  we  are 
of  the  opinion  that  Jiingerniannia  nivalis  Sw.  should  be  treated  as  an  exact  synonym  of 
Jiingerniannia  jtilacea  Lightf.,  inasmuch  as  we  are  in  possession  of  no  evidence  (as  has 
already  been  said,  in  effect,  by  Dr.  Spruce)  that  Lightfoot,  Swartz,  or  Wahlenberg  dis- 
tinguished between  what  we  now  recognize  as  Anthelia  jnlacea  and  Anthelia  Juratz- 
kana.  Junge7-T7iannia  nivalis  could  then  be  cited  as  a  synonym  for  Jungej-jnamiia 
jnlacea^  in  the  restricted  sense  of  Limpricht,  with  as  much  justice  as  for  Jnngermannia 
Jni'atzkana.  At  all  events,  in  Wahlenberg's  Flora  ^\\e.z\z2i.,  Jungermanjiia  nivalis  is 
used  in  contradistinction  to  Jungermannia  julacea  which  is  applied  to  the  Jiinger- 
niannia concinnata  of  Lightfoot  and  of  Hooker,  and  the  diagnosis  of  Jungerman7iia 
nivalis  as  there  given,  would,  on  the  whole,  [^^  surculis  setaceis  ramosis '■^  iurculi 
*  *  ^  Jlexuosi''^)  be  more  suggestive  of  the  dioicous  species.  Lindberg,  it  is  true- 
states  (Kongl.  Sv.  Vet.  Akad.  Handl.  23^:  29.  1889)  that  his  Anthelia  7iivalis  (the 
paroicous  species)  is  very  common  in  Scandinavia  while  A.  Julacea  is  very  rare,  but 
Kaalaas  (Xyt  Mag.  for  Naturvidensk.  33  :  230,  231,  233,  234.  1893)  finds  the  two 
species  about  equally  abundant  in  Norway.  Until  proof  is  forthcoming  that  Jungei- 
mannia  nivalis  was  originally  applied  to  the  paroicous  species  we  prefer,  with  Spruce, 
to  use  a  specific  name  concerning  whose  original  meaning  there  is  no  doubt. 


144 


JUNGERMANNIACEAE 


Antliclia  julacca{\^.\  Limpr.)  Dumort. — a  nearly  related  species, 
which  may  be  expected  to  occur  in  California — is  distinguished  by 
its  larger  size,  dioicous  inflorescence,  thicker-walled  leaf-cells,  and 
bispiral  elaters. 

31.  PTILIDIUM  Nees,  Naturgesch.  Eur.  Leberm.  i  :  95.  1833. 
BlcpJiarozia  Dumort.  Rec.  d'Obs.  Jung.  16.  1835. 

Usually  in  dense  and  commonly  brownish  mats.  Stem  pros- 
trate or  ascending,  i— 2-pinnate  or  irregularly  and  sparingly 
branched,  eflagelliferous,  the  branches  lateral :  root-hairs  few  and 
short.  Leaves  obliquely  inserted,  twice  bifid  to  below  the  middle 
or  palmately  3-5 -cleft,  the  dorsal  segments  larger  and  incubous, 
all  filiform-acuminate,  the  segment-margins  entire,  or  more  com- 
monly fringed  with  long,  simple  or  branched  cilia.  Underleaves 
much  smaller  but  otherwise  somewhat  similar,  more  or  less  dis- 
tinctly 2-  or  3 -parted.  Antheridia  short-stalked,  on  the  main  stem 
or  lateral  branches,  in  the  axils  of  more  concave  and  more  closely 
imbricate  but  otherwise  scarcely  modified  leaves.  Archegonia 
terminal  on  what  is  primarily  the  main  stem  or  a  principal  branch, 
the  perianth,  however,  through  innovation  below,  finally  apical  on 
a  short  apparently  lateral  or  dichotomously  divergent  branch. 
Bracts  commonly  one  or  two  pairs,  similar  to  the  leaves  or  often 
more  profusely  eiliate-laciniate.  Perianth  free,  several  times  longer 
than  the  bracts,  cylindrical -obovate,  ciliate  and  subplicate  at  the 
somewhat  contracted  mouth.  Calyptra  free.  Capsule  ovoid  on  a 
moderately  long  seta,  dehiscing  to  the  base  by  four  rather  rigid 
valves,  the  cells  of  the  latter  bearing  semiannular  and  also  brownish 
nodular  thickenings.  Spores  punctulate,  several  times  broader 
than  the  2-  (3-)  spiral  elaters. 

I.  Ptilidium  Californicum  (Aust.)  Undervv.  &  Cook,  Hep.  Am. 
69.    May,  1890.    Pearson,  List  Can.  Hep.  7.    16  Je.  1890. 
Lcpidozia  ?  Calif ornica  Aust.  Bull.  Torn  Bot.  Club,  6:  19. 

1875. 

Mastigophora  CaUforiiica  Aust.  Bull.  Torr.  Bot.  Club.  6  :  302. 
1879. 

Brownish  or  greenish-yellow,  usually  in  dense  appressed  mats  : 
stems  subfiliform  and  flaccid  or  becoming  somewhat  stout,  mostly 
2-3  cm.  long,  irregularly  and  sparingly  branched,  rarely  subpin- 
nate  or  subbipinnate  :  leaves  loosely  imbricate,  palmately  3-  or  4- 
cleft  to  below  the  middle,  the  segments  lanceolate  or  more  com- 
monly linear-lanceolate,  filiform-attenuate,  unequal,  the  ventral 


Ptilidium 


345 


smaller  and  subinflexed,  the  segment-margins  very  entire,  repand, 
or  here  and  there  again  incised  or  sometimes  bearing  a  few  (2  or 
3  on  a  side)  long  and  rather  rigid  cilia  :  underleav^es  usually  a  little 
broader  than  the  stem,  deeply  2-  or  3 -parted,  the  segments  in- 
ciso-ciliate  ;  leaf-cells  40-45  hexagonal-  or  pentagonal-rotund, 
the  walls  strongly  thickened  at  the  angles,  cuticle  smooth : 
dioicous  :  ^  plants  commonly  more  slender,  androecium  cauline 
or  rameal,  antheridia  ovoid  or  ellipsoidal,  single  or  in  pairs,  the 
stalks  finally  much  shorter  than  their  long  diameter  :  9  branch 
short,  bearing  about  2—4  pairs  of  leaves  inclusive  of  bracts  :  bracts 
one  or  two  pairs,  a  little  larger  than  the  leaves  and  more  inciso- 
ciliate  ;  bracteole  similar,  rarely  connate  with  bracts  on  one  or  both 
sides  ;  perianth  cylindrical-obovoid  to  subfusiform-oblong  from  an 
obconic  base,  slightly  narrowed  at  the  ciliate  subplicate  mouth  : 
spores  brown,  21-30  u. ;  elaters  120-225  ux  7-10  a,  2-  (rarely  3-) 
spiral,  contorted,  subobtuse. 

Found  in  California  only  by  its  first  collector,  Dr.  Bolander. 
The  plants  from  which  the  original  description  was  drawn  were 
picked  out  from  specimens  of  Hypmun  circinale  in  Sulliv.  &  Lesq. 
Muse.  Bor.-Am.  Exsicc,  No.  474,  ed.  2  ;  these  are  now  preserved 
in  herb.  Pearson.  Duplicates  of  this  original  material  may  often 
be  found  in  connection  with  no.  474  of  the  Exsicc.  cited.  Habitat 
and  locality  were  given  as  ad  corticem  arborum  in  sylvis  Cali- 
forniae  montanae."  The  specimens  doubtless  came  from  Hum- 
boldt or  Mendocino  counties,  and  the  plant  is  to  be  looked  for  on 
the  bark  of  living  trees  and  fallen  trunks  of  Sequoia  senipervirens 
and  of  other  conifers  in  the  northern  coast  region.  The  species 
extends  northward  to  British  Columbia  (Macoun)  and  to  Idaho 
(Leiberg).  Some  of  the  northern  specimens  make  a  nearer  ap- 
proach to  Ptilidiitin  ciliare,  which  also  occurs  in  the  same  region,  but 
are  always  very  distinct  in  the  less  branched  stem,  the  more  rigid 
leaves,  with  much  more  pronouncedly  filiform-attenuate  segments 
and  longer,  more  rigid,  and  always  much  less  numerous  cilia. 

Plate  105.    Ptilidium  Califormcum. 

1.  Plants,  natural  size. 

2.  Portion  of  stem,  with  branch,  X  12. 

3.  Cauline  leaf,  viewed  from  above,  X  23. 

4.  Cauline  leaf,  viewed  from  below,  X  23. 

5.  A  3-cleft  cauline  leaf,  viewed  from  below,  X  23. 

6.  A  typical  apex  of  leaf-segment,  X  4i- 

7.  Cauline  leaf,  ventral  view,  X  23. 


146 


JUXGERMANXIACEAE 


8.  Rameal  leaf,  ventral  view,  X^S- 
9  and  lo.  Underleaves,  X^S- 

11.  Androecium,  X  12. 

12.  ^  bract,  X  23. 

13.  Antheridia,  X  23. 

14.  Bracteole,  X  23. 

15.  Second  (next  to  the  inmost)  bracteole  from  another  plant,  X  23. 

16.  Inmost  bract,  X  23. 

17  and  18.  Ventral  and  dorsal  aspects  of  perianth  and  bracts,  \  12. 

19.  Portion  of  perianth- mouth,  X  4^- 

20.  Leaf-cells,  X  225. 

Figs.  1-6,  9,  10,  and  20,  drawn  from  the  type  specimen  collected  in  California  by 
Bolander,  ex,  herb.  Pearson  ;  the  remaining  tigures  from  Hep.  Am.  69  (Idaho,  J.  B. 
Leiberg). 

32.  SCAPANIA  Dumort.  Rec.  d'Obs.  Jung.  14.  1835. 

Martinclli2is  S.  F.  Gray,  p.  p.  max.  Nat.  Arr.  Brit.  PI.  I  :  690. 
1821.  Lindb.  (as  Martinellia)  Act.  Soc.  Sci.  Fenn.  10:  518. 
1875.    O.  Kuntze  (as  Martinellia),  Rev.  Gen.  PI.  83;.  1891. 

Radid a  Dumovt.  p.  p.  Comm.  Bot.  112.  1822. 

Radula  §  Scapania  Dumort.  Syll.  Jung.  38.     183  i. 

Plants  usually  large,  caespitose,  green,  brown,  rose-red,  or 
dark-purple ;  primary  stems  defoliate,  creeping,  the  secondary 
mostly  ascending,  sometimes  erect,  rarely  prostrate,  simple  or 
dichotomously  branched,  apex  commonly  decurved.  Leaves  al- 
ternate, distichous,*  complicate-bilobed,  the  fold  rounded  out- 
wardly or  more  often  with  an  acute  or  somewhat  winged  keel,, 
margins  of  the  lobes  ciliate,  dentate,  or  entire  ;  ventral  lobes  mostly 
larger,  convex  dorsally,  succubous ;  dorsal  lobes  incumbent ; 
underleaves  wanting.  Stem-apex  and  upper  leaves  sometimes 
gemmiferous.  Dioicous  or  rarely  monoicous ;  androecium  ter- 
minal or  interrupted ;  $  bracts  ventricose,  usually  smaller  and 
with  subequal  lobes  ;  antheridia  (mostly  1-6)  ovoid  or  ellipsoidal, 
on  pedicels  of  about  their  own  length,  often  accompanied  by  hair- 
like or  leaf-like  paraphyses.  Archegonia  few,  at  the  apices  of 
the  main  branches.  9  bracts  hardly  different  from  the  ordinary 
leaves,  though  a  little  larger  and  more  equally  lobed.  Perianth 
oblong  or  obovate,  strongly  compressed  dorso-ventrally  (parallel 
with  the  plane  of  the  stem),  smooth,  or  rarely  subplicate,  the 
mouth  broad,  truncate,  entire,  dentate,  or  ciliate,  decurved  when 
young.  Capsule  oblong-ovoid,  long-exserted,  valves  thick-walled, 
the  inner  layer  of  cells  with  semiannular  or  nodose  thickenings. 
Elaters  bispiral,  deciduous. 


*  See  Scapania  heteropJiylla. 


SCAPANIA 


147 


Key  to  tlie  Species. 

Uppermost  leaves  (in  9  plants)  with  long  curved  teeth  on  the  carina;  perianth -mouth 

denticulate  or  subentire.  3.  S.  Oak  sii. 

Leaves  without  teeth  (or  very  rarely  a  single  tooth)  on  the  carina  or  fold. 

Basal  margin  of  dorsal  leaf- lobes  with  long,  decurved,  often  compound  cilia  ;  leaves 

coarsely  dentate  ;  ventral  lobes  oblong-ovate.  I .  S.  Bolandei-i. 

Basal  margin  of  dorsal  leaf-lobes  entire  or  with  small  teeth. 

Plant  ("  frond")  distinctly  complanate  ;  leaves  always  distichous. 

Stems  4-15  mm.  long;  ventral  leaf-lobe  very  slightly  or  not  at  all  de- 
cunent. 

Leaves  bilobed  their  length  ;  ventral  lobes  oblong-ovate,  acute 

or  subacuminate,  serrate-dentate,  deflexed  and  often  somewha. 
secund  ;  dorsal  lobes  nearly  parallel  to  the  stem,  appressed-imbri- 
cate  above;  perianth-mouth  entire  or  repand.     5.  S.  tmibrosa. 

Leaves  bilobed  j4.  their  length  ;  ventral  lobes  orbicular-ovate  to  obo- 
vate,  often  obtuse,  entire  or  sparingly  dentate  ;  dorsal  lobes  erecto- 
patent  or  patent-horizontal,  ascending,  never  appressed-imbricate  ; 
perianth  mouth  in  most  cases  shortly  ciliate-dentate. 

6.  S.  curt  a. 

Stems  l-io  cm.  long,  ventral  leaf-lobe  decurrent,  obtuse,  rarely  subacute. 
Plants  aquatic  ;  leaves  flaccid,  lobes  often  subequal  at  apex,  some- 
times entire,  the  ventral  round- trapezoidal  ;  perianth-mouth  repand 
or  subdentate.  4.  S.  tindulata. 

Plants  non-aquatic  ;  leaves  somewhat  stiff,  ciliate-dentate ;  ventrat 
lobes  obovate,  1-2.]/,  times  the  size  of  the  dorsal ;  perianth-mouth 
ciliate-dentate.  2.  .5".  neuiorosa. 

Plant  ( "  frond  "  )  obscurely  complanate  ;  leaves  sometimes  irregularly  3-ranked 
by  interpolation  of  unlobed  leaves,  erecto-patent,  often  with  squaiTose  tips. 

7.  S.  heterophylla. 

I.  ScAPAXL\  BoLAXDERi  Aust.  Proc.  Acad.  Nat.  Sci.  Philad.  1869  : 

218.  1869. 
Scapaiiia  c  mi  data  Tayl.  in  herb. 

Scapania  Calif ornica  Gottsche ;  Cal.  Med.  Gaz.  1870:  184 
(40)  (name  only).  1870. 

Scapania  albescens  Steph.  Bot.  Jahrb.  8  :  96.  1886. 

Olive-green  to  yellowish-brown,  caespitose  :  stems  rigid,  as- 
cending, nearly  prostrate  or  subpendulous,  1-4  cm.  long,  dichot- 
omous,  9  plant  innovating  from  near  base  of  the  perianth  : 
leaves  scarcely  increasing  in  size  upward,  approximate  or  close, 
stiff,  hardly  changed  in  form  or  position  on  drying,  acutely  com- 
plicate, narrowly  alate-carinate,  the  carina  entire  or  rarely  fur- 
nished with  a  single  tooth  ;  ventral  lobes  obliquely  oblong-ovate, 
patent,  strongly  convex,  obtuse,  slightly  decurrent,  coarsely  den- 
tate, 2^  times  greater  than  the  dorsal,  apex  and  ventral  margin 
deflexed  ;  dorsal  lobes  broadly  and  obliquely  ovate,  more  or  less 


148 


JUNGERMANNIACEAE 


imbricate,  erect  or  subappressed,  slightly  conv^ex,  somewhat  acute, 
with  fewer  and  larger  teeth  and  bearing  at  the  base  several  long 
decurved  often  compound  cilia  ;  leaf-cells  subquadrate-oval,  more 
elongated  toward  base,  i  5-40  with  conspicuous  trigones,  cu- 
ticle minutely  roughened  :  gemmae  at  apex  of  stem,  of  one  or  two 
cells,  oval  or  ellipsoidal,  15-25  ax  15  a:  dioicous,  $  and  $ 
plants  in  the  same  tuft :  androecium  terminal  or  often  interrupted  ; 
antheridia  2-6,  ellipsoidal  or  obovoid,  on  stalks  of  about  their 
own  length  or  longer,  accompanied  by  very  numerous,  branching, 
capillary,  or  leaf-like  paraphyses  ;  $  bracts  scarcely  modified 
or  smaller,  saccate,  densely  imbricate,  with  subequal  inciso- 
dentate  lobes:  perianth  oblong,  more  rarely  obovate,  2-3.5 

X  1. 5-1.8  mm.,  ciliate-lacinulate  or  doubly  ciliate-dentate  at 
mouth  :  capsule  oblong-ovoid,  brown,  seta  3—10  mm.  long  ;  spores 

10-12  u.,  punctulate  ;  elaters  bispiral,  obtuse,  contorted,  100-175 
fi  X  8- 1  o  fj.. 

Exsicc.  Hep.  Bor.-Am.  19. 

Common  in  the  Coast  Range  Mountains,  north  of  San  Francisco'^ 
on  logs  and  stumps  of  the  Coniferae,  especially  of  Sequoia  scnipcr- 
virens — more  rarely  on  the  bark  of  living  trunks. 

Redwood  Canon,  Marin  Co.  (Miss  Edith  S.  Byxbee);  Dun- 
can's Mills  (i  193)  and  Turner's  Canon  (i  194),  Sonoma  Co.;  Men- 
docino (596),  Little  River  (648,  684),  and  Half- Way  House  (727), 
Mendocino  Co.;  Eureka  (911,  928,  938,  984),  Humboldt  Co. 
Frequently  with  sporogonia.  Apparently  closely  confined  to  the 
Pacific  Coast,  ranging  as  far  northward  as  Alaska.  Specimens 
from  Nova  Scotia,  Ontario,  and  Lake  Superior,  referred  to  this 
species  by  Mr.  Pearson  *  are  probably  something  else.  Scapaiiia 
Bolaiideri  is  strongly  marked  by  the  rigid  oblong-obovate  ventral 
lobes  and  by  the  coarsely  dentate  dorsal  lobes,  furnished  at  the 
base  with  long  curved  often  branched  cilia.  These  cilia  often  lie 
close  to  the  stem  and  are  covered  by  the  dorsal  lobe  next  below, 
so  that  a  careful  separation  of  the  leaves  from  the  stem  is  usually 
necessary  for  a  satisfactory  demonstration  of  this  character. 

The  plant  when  growing  in  tufts  of  Dicraimui  is  sometimes 
erect.  We  have  rarely  met  with  gemmiferous  conditions.  The 
original  specimens  w^cre  collected  in  California  by  Dr.  Bolander, 
and  were  distributed  by  Mr.  Austin  as  no.  19  of  the  Hepaticac 
Boreali-Americanae. 


*  List  of  Can.  Hep.  13.  1890. 


SCAPANIA 


149 


Unpublished  figures  of  this  species,  drawn  by  Dr.  Gottsche, 
are  preserved  in  the  Gray  Herbarium  at  Cambridge,  Mass.  The 
sheet  on  which  the  figures  are  drawn  bears  the  inscription  "  Sca- 
pania  Cahfornica  G.  MS.,"  in  addition  to  the  name  given  the 
species  by  Mr.  Austin. 

Scapania  Bolaiideri  is  reported  from  Japan  by  Herr  Stephani, 
(Bull.  Herb.  Boiss.  5  :  82.  1897). 

Plates  106  and  107.    Scapania  Bolanderi. 

1.  Plants,  natural  size. 

2.  Apical  portion  of  a  sterile  plant,  y^ii. 

3.  Portion  of  a  ^  plant,  showing  position  of  androecia,  X  ^2. 

4.  Ciliate  margin  of  the  base  of  the  dorsal  lobe,  X  ^o^- 

5.  Apex  of  ventral  lobe,  X  106. 

6.  Leaf,  dorsal  view,  X  23. 

7.  Leaf,  ventral  view,  X  23- 

8.  Young  perianth  and  bracts,  ventral  view.  X  12. 

9.  Perianth,  bracts,  and  sporogonium,  X  ^2. 
10   and  II.   ^  bracts  and  antheridia,  X  ^2. 

12.  Antheridia,  X  53- 

13.  Antheridial  paraphyses,  X  106 

14.  Outline  of  a  portion  of  perianth-mouth,  X  53- 
15    Leaf  cells  near  apex  of  ventral  lobe,  X  305- 

Figs  I,  3,  and  10-12,  from  no.  938  (Eureka);  2,  6  and  7,  from  no.  984  (Eureka); 
4,  5,  8,  9,  14  and  15,  from  no.  596  (Mendocino);  13  from  no.  928  (Eureka). 

2.   Scapania  xemorosa  (L.)   Dumort.   Rec.  d'Obs.  Jung.  14. 

1835. 

Jiingermannia  nciiiorosa  L.  Sp.  PI.  2:  1132.  1753- 
Brownish  or  olive-green,  usually  in  broad,  compact  cushions  : 
stems  rather  rigid,  1-6  cm.  long,  sparingly  branched,  ascending, 
with  the  apex  more  or  less  decurved  :  leaves  somewhat  stiff,  ap- 
proximate or  slightly  imbricate,  increasing  a  little  in  size  upward, 
irregularly  ciliate-dentate,  more  or  less  evidently  winged  along  the 
fold,  the  ventral  margin  decurrent ;  ventral  lobes  2—2.5  times 
larger  than  the  dorsal,  obovate,  convex,  obtuse,  apex  and  ventral 
margin  commonly  deflexed  ;  dorsal  lobes  reniform  or  broadly  and 
obliquely  ovate,  rounded  at  apex  or  with  a  short  point,  usually  a 
little  vaulted,  the  margin  somewhat  appressed ;  leaf-cells  nearly 
uniform,  oval-quadrate,  thick-walled,  cuticle  roughened  :  gemmae 
sometimes  abundant  at  stem-apex  and  on  margins  of  upper  leaves, 
yellowish-brown,  oval,  unicellular  or  uniseptate,  15-24//  x  1 5 />« 
occasionally  in  moniliform  threads  :  dioicous  :  androecium  terminal, 
^  bracts  scarcely  different,  antheridia  2-6,  accompanied  by  nu- 
merous long,    multiseptate,  simple  or  branched,  hair-like  para- 


150 


JUNGERMANNIACEAE 


physes,  brown  trichomes  of  a  similar  structure  sometimes  occur- 
ring on  other  parts  of  the  stem  :  perianth  long-obovate,  2-3.5  t^^^- 
X  1.5-2  mm.,  now  and  then  subpHcate,  ciliate-dentate  at  mouth  : 
capsule  ovoid-oblong,  brown,  seta  5-1  5  mm.  in  length  ;  spores  12- 
16  fj.,  punctulate  ;  elaters  contorted,  100- 180//  x  y-io/Jt. 

On  rocks  and  moist  banks.  Mt.  Tamalpais  (21,  ^  and  gem- 
miferous) ;  Lake  San  Andreas  (1192,  c./r.)  San  Mateo  Co.;  near 
Cazadero  (1191,  c./r.),  Sonoma  Co.;  Crescent  City  (Mr.  Thomas 
Howell),  Del  Norte  Co.  A  few  slender  gemmiferous  plants  from 
near  Lake  Lagunitas,  Marin  Co.,  appear  to  belong  with  this  species 
also,  though  the  dorsal  lobes  are  commonly  squarrose  and  the 
teeth  are  shorter  and  less  numerous  than  is  usual. 

The  cuticle  in  the  Californian  specimens  is  for  the  most  part 
distinctly  granulate  or  verruculose,  often  as  much  so  as  in  a  speci- 
men of  Scapania  aspera  Miiller  &  Bernet  *  collected  and  deter- 
mined by  Loitlesberger  and  in  another  collected  and  determined 
by  Pearson,  but  we  are  unable  as  yet  to  distinguish  Scapania  aspera 
from  wS.  ncmorosa.  A  specimen  collected  by  Loitlesberger  in 
"  Saminathal  (Vorarlberg)  800  m."  and  named  by  him  Scapania 
nemorosa  has  its  leaf-cells  more  decidedly  scabrous  than  has  either 
of  these  specimens  which  have  been  referred  to  Scapania  aspera. 

3.  Scapania  Oakesii  Aust.  Bull.  Torn  Bot.  Club,  3  :  10.  1872. 

Light-green,  brownish,  or  fuscous-purple,  caespitose  :  stems 
somewhat  rigid,  sparingly  branched,  ascending,  1-2.5  cm.  long: 
leaves  tense  or  a  little  flaccid,  approximate  or  close,  crowded  and 
larger  at  apex  in  9  plants,  alate-carinate,  the  carina  in  the  upper 
leaves,  especially  the  involucral,  broad  and  furnished  with  a  single 
or  sometimes  a  double  series  of  long,  usually  curved,  teeth,  these 
wanting  or  infrequent  in  $  plant,  ventral  margins  decurrent ;  ven- 
tral lobes  obliquely  obovate  or  rhombic-trapezoidal,  2-2.5  times 
greater  than  the  dorsal  below,  subequal  at  the  apex,  convex,  obtuse 
or  sometimes  slightly  pointed,  serrate-dentate  ;  dorsal  lobes  round- 
ovate,  scarcely  imbricate  except  at  stem-apex,  convex,  obtuse  or 
subacute,  the  upper  often  unicalcarate-dcntate  at  basal  margin,  the 
teeth  otherwise  rather  smaller  and  less  numerous  than  in  the  ven- 
tral lobes  :  marginal  and  median  leaf-cells  quadrate-oval,  i  5-24  //, 
oblong-hexagonal  near  the  base,  30-56  (l  x  16-18 trigones  in- 
conspicuous, cuticle  plainly  roughened  :  sometimes  gemmiferous 
at  stem-apex  or  on  margins  of  the  upper  leaves,  gemmae  mostly 


*  liemet,  Cat.  Hep.  du  Sud-Ouest  de  la  Suisse  et  de  la  Ilaute-Savoie,  42.  1888. 


SCAPANIA 


151 


unicellular,  spherical  or  oval,  i  5-20  ft.,  occasionally  in  branching 
moniliform  threads :  dioicous  :  $  and  9  plants  mingled  or  in 
separate  tufts,  $  plants  more  slender :  androecium  terminal, 
$  bracts  smaller,  approximate  or  rather  distant,  more  rarely 
subimbricate,  ventricose,  lobes  nearly  equal,  the  dorsal  often  cal- 
carate-dentate  at  base,  antheridia  2-6,  accompanied  usually  by  a 
very  few  short  capillary  or  sometimes  leaf-like  paraphyses  :  perianth 
oblong  or  obovate,  truncate,  denticulate  or  subentire  at  the  mouth. 

On  moist  banks.  Mendocino  (590,  near  Eureka  [945, 
946  {c.  pej\y\  and  Kneeland  Prairie  Road  (1025),  Humboldt  Co. 

Scapania  Oakesii,  as  represented  in  Austin's  Hep,  Bor.-Am., 
no.  14,  and  in  two  other  specimens  from  herb.  Austin  in  herb. 
Underwood  (all  from  the  White  Mountains  of  New  Hampshire)  is 
a  large  plant  evidently  veiy  closely  related  to  Scapania  iindiilata 
purpurea.  Our  plants  are  usualty  smaller,  more  copiously  sub- 
ciliate-dentate  on  the  carina,  and  have  more  resemblance  super- 
ficially to  S.  neinorosa,  yet  are  certainly  distinct  from  either  of  the 
species  named.  They  accord  veiy  closely  with  Dr.  Scouler's 
specimens  from  Observatory  Inlet,  in  herb.  Torrey,  which  Austin 
made  var.  &  of  Scapania  Oakesii.  In  some  of  the  Eureka  plants 
the  carina  is  richly  provided  with  teeth  as  far  as  the  sev^enth  or 
eighth  pair  of  leaves  from  the  stem-apex. 

Plates  108  and  109.    Scapania  Oakesii. 

1.  Plants,  natural  size. 

2.  Perianth  and  bracts,  ventral  view,  X  ^2. 

3  and  6.   ^  plants,  X  ^2,  the  former  gemmiferous  at  apex. 

4.  Apical  portion  of  a  9  plant,  dorsal  view,  X  12. 

5.  Perianth  and  bracts,  dorsal  view,  X  ^2. 

7.  $  bract  and  antheridia,  X  23. 

8.  Antheridium,  X  53- 

9.  Antheridial  paraphyses,  X  ^o^- 

10  and  II.  Leaves  from  near  the  stem-apex  of  a  sterile  9  plant,  X  23. 

12.  Leaf,  ventral  view,  X  23. 

13.  Leaf  from  lower  part  of  stem,  carina  entire,  X  23. 

14.  9  bract,  X  23. 

15.  Leaf-cells  from  near  apex  of  ventral  lobe,  X  3^5 • 

16.  Portion  of  perianth-mouth,  X  53- 

17.  Margin  of  apex  of  ventral  lobe,  X  ^o^. 

f  igs-  I»  3,  4,  6-13,  15,  and  17,  from  no.  946  (near  Eureka);  2,  5,  and  14,  from 
no.  945  (near  Eureka);  16,  from  no.  1025  (Kneeland  Prairie  Road). 


152 


JUNGERMANNIACEAE 


4.  ScAPAxiA  uxDULATA  (L.)  Dumort.  Rec.  d'Obs.  Jung.  14.  1835, 
JiLUgennajuiia  ttiidiLlata  L.  Sp.  PI.  2:  1132,     1753.  Hook, 

Brit.  Jung. //.  18 16. 

Green,  rose-red,  or  dark  purple,  rarely  brownish,  in  compact  or 
loose  tufts  :  stems  rather  rigid,  usually  erect,  sometimes  floating, 
I- 10  cm.  long,  sparingly  branched,  with  very  few  root-hairs, 
denudate  and  brittle  below:  leaves  larger  and  imbricate  at  apex, 
approximate  or  distant  below,  mostly  soft  and  flaccid,  usually  un- 
dulate-crisped or  crumpled  in  drying,  entire,  denticulate,  or  ciliate- 
dentate,  slightly  alate-carinate,  carina  entire  ;  ventral  lobes  round- 
trapezoidal,  mostly  twice  the  size  of  the  dorsal,  subequal  toward 
the  stem-apex,  sometimes  broadly  pointed,  decurrent,  convex  or 
nearly  flat ;  the  dorsal  lobes  equally  broad,  obliquely  and  broadly 
ovate,  with  a  rather  obtuse  point,  loosely  incumbent  or  somewhat 
bent  away  from  the  stem  :  leaf-cells  near  the  margin  quadrate  or 
oval,  1 5-20  oblong-hexagonal  in  the  middle,  45—60  fi  x  1 5-30  fi^ 
mostly  thin-walled,  cuticle  more  or  less  distinctly  hyaline-rough- 
ened :  dioicous  :  perianth  oblong  or  more  rarely  obovate,  slightly 
narrowed  at  the  entire,  repand,  or  subdentate  mouth. 

On  stones  in  streams  and  springs  or  in  very  moist  places^ 
especially  in  mountainous  regions. 

North  Fork  of  the  Little  River,  Mendocino  Co.,  on  submerged 
rocks  (600).  No.  649,  from  the  same  locality  but  growing  on  a 
log  just  above  the  water-line  appears  to  be  a  variety  approaching 

5.  nemorosa  in  the  subciliate-dentate  upper  leaves  and  in  the  form 
and  relative  size  of  the  lobes,  having,  however,  the  subentire  peri- 
anth mouth  of  5.  luidulata.  No.  639,  sterile,  from  wet  rocks  in 
stream-bed  seems  to  agree  with  the  latter,  as  does  also  a  specimen 
collected  in  the  region  of  the  Yosemite  Valley  by  C.  M.  Cooke, 
Jr.,  in  1896  (ex  dono  A.  W.  Evans).  Also  collected  in  California 
by  Dr.  Bolander,  the  exact  locality  unknown. 

Scapania  uliginosa  (Sw.)  Dumort.,  so  far  as  we  know,  has  not 
yet  been  collected  in  California,  but  its  discovery  there  may  be 
expected.  It  differs  from  Scapania  nndulata  chiefly  in  the  smaller, 
reniform,  strongly  convex  dorsal  lobes,  much  narrower  than  the 
ventral  lobes  and  their  size,  and  in  the  always  entire  leaves 

decurrent  on  both  sides,  especially  long-decurrent  on  the  ventral 
margin.  » 

Scapania  irrigua  (Nees)  Dumort.  differs  from  .V.  nndulata  in 
the  softer,  weaker  stems,  bearing  root-hairs  to  the  apex,  in  the 


SCAPANIA 


15B 


thinner,  softer,  less  decurrent  leaves,  with  nearly  orbicular  lobes, 
of  which  the  convex  dorsal  is  the  size  of  the  ventral  and 

sharply  pointed,  the  ventral  commonly  with  a  short  point. 

5.  ScAPANiA  UMBROSA  (Schrad.)  Dumort.  Rec.  d'Obs.  Jung.  14. 

1835- 

Jimgennannia  lunbrosa  Schrad.  Syst.  Samml.  Krypt.  Gew.  2  : 
5.  1797.  Schrad.  Jour.  Bot.  1801  :  67.  1803.  Hook.  Brit. 
Jung.  //.  2^.     1 8 16. 

Jungennannia  convexa  Scop.  (?)  Fl.  Carn.  2:  349.  1772. 
[2d  ed.] 

Martinellia  convexa  Lindb.  Muse.  Scand.  6.  1879. 

Scapania  convexa  Pearson,  List  Can.  Hep.  15.  1890. 

Yellowish-green  to  brown,  sometimes  tinged  with  purple,  in  an 
extended,  closely  appressed,  usually  compact  stratum  :  stems  5—1  5 
mm.  long,  branching  or  subsimple,  ascending,  decurved  at  apex  es- 
pecially when  dry  :  leaves  increasing  somewhat  in  size  upward,  not 
decurrent,  bilobed  their  length,  the  fold  rounded  or  acute, 

sometimes  with  a  trace  of  a  winged  keel ;  ventral  lobes  2-3  times 
larger  than  the  dorsal,  oblong-ovate,  acute  or  subacuminate,  ir- 
regularly serrate-dentate  toward  apex,  deflexed  and  often  some- 
what secund ;  doi'sal  lobes  ovate,  acute,  sometimes  narrowly 
pointed,  dentate,  neaily  parallel  to  the  stem,  appressed-imbricate, 
except  in  slender  sterile  conditions :  leaf-cells  small,  roundish-oval, 
12-27  fL,  more  elongated  toward  base,  thick-walled,  trigones  dis- 
tinct, cuticle  minutely  roughened  :  gemmae  in  dark-brown  clusters 
at  stem- apex,  mostly  uniseptate,  oblong-elliptical,  18x9/^.:  dioi- 
cous :  androecium  terminal,  antheridia  1-3  in  axils  of  smaller, 
saccate,  nearly  equal-lobed  leaves,  accompanied  by  a  few  short, 
usually  septate  hairs  :  perianth  oblong,  from  an  obconic  base,  1.2- 
2  mm.x.'6-i  mm.,  twice  as  long  as  the  involucral  leaves,  often 
purple  at  base,  compressed,  mouth  entire  or  repand :  capsule  oval- 
oblong,  dark-brown,  long-exserted  ;  spores  brown,  punctulate, 
10  fj.\  elaters  125-165  /iX7-8  /v.. 

Exsicc.  Hep.  Am..  191. 

On  old  logs  beside  shaded  streams.  North  Fork  of  the  Little 
River,  Mendocino  Co.  (647,  683)  and  Eureka,  Humboldt  Co. 
(961)  ;  also  on  compact  argillaceous  soil  in  the  first-named  locality 
(686).  No.  711,  from  a  log  in  Russian  Gulch,  near  Mendocino, 
which  we  formerly  (Erythea  5  :  89.  1897)  referred  to  Scapania 
glaucocepJiala,  we  now  believe  to  be  an  abortive,  gemmiferous  con- 


154 


JUNGERMANNIACEAE 


dition  of  Scqpania  tunbrosa.  Collected  by  Professor  Macoun  on 
Mt.  Mark,  Vancouver  Island. 

Our  plants  appear  to  agree  with  the  European  in  all  the  more 
important  details,  yet  the  marginal  teeth  of  the  leaves  are  rather 
larger  and  more  numerous  than  in  most  of  the  European  speci- 
mens examined,  the  cuticle  less  distinctly  roughened,  the  dorsal 
lobes  a  little  less  closely  imbricated,  and  the  perianth  is  sometimes 
longer. 

It  is  very  probable,  as  was  first  pointed  out  by  Lindberg,"^  that 
Jungennannia  convexa  Scop,  is  identical  with  J.  tunbrosa  Schrad., 
though  the  perianths  thecae'')  were  described  by  Scopoli  as 
oval,  and  dentate  at  the  apex,  while  those  of  Scapania  umbrosa 
are  oblong,  with  an  entire  or  repand  mouth.  We  have  been  in- 
formed by  Prof  Briosi  that  the  Scopoli  collection  is  not  preserved 
at  Pavia  and  as  we  have  been  unable  to  locate  it,  we  prefer  to 
retain  for  this  species  a  name  concerning  the  correct  application  of 
which  we  can  have  no  doubts. 

6.  ScAPANiA  CURTA  (Mart.)  Dumort.  Rec.  d'Obs.  Jung.  14.  1835. 

Jiuigcrinannia  airta  Mart.  Fl.  Crypt.  Erlang.  148.//.  ^. /. 
1817. 

Bright-  to  dark-green,  sometimes  brownish,  loosely  gregarious 
or  in  thin  mostly  inconspicuous  mats  :  stems  ascending,  4-15  mm. 
long,  .1-.35  mm.  thick,  simple  or  innovating  above  :  leaves  rather 
soft,  usually  increasing  in  size  toward  the  stem-apex,  bilobed  to 
about  the  middle,  the  fold  outwardly  rounded  or  acute,  rarely  with 
a  trace  of  a  wing ;  ventral  lobes  2-3  times  larger  than  the  dorsal, 
orbicular-ovate  to  obovate,  acute,  apiculate,  or  obtuse,  entire  or 
sparingly  dentate,  patent-horizontal,  slightly  or  not  at  all  decurrent, 
a  little  concave  ventrally  ;  dorsal  lobes  deltoid-  or  quadrate-ovate, 
crecto-patent  to  patent-horizontal,  acute,  entire  or  slightly  dentate, 
ascending,  never  appressed-imbricate  ;  leaf-cells  translucent,  round- 
ish-hexagonal, 16-36//,  becoming  oblong  toward  the  base,  the 
walls  with  small  trigones,  cuticle  nearly  smooth ;  gemmae  in  a 
dense  cluster  at  the  stem-apex  or  borne  on  the  margins  of  the 
upper  leaves,  brownish  or  colorless,  ovoid  to  oblong-ellipsoidal, 
24  X  i6/>«,  simple  or  uniseptate  :  dioicous  :  androecium  terminal, 
antheridia  single  or  in  groups  of  2  or  3  :  perianth  long-obovate, 
1.8-3.5  X  1-1.8  mm.,  the  mouth  shortly  ciliate-dentate,  rarely 
repand. 


*I\lusc.  Scand.  6.  1897. 


SCAPANIA 


155 


On  a  log  by  a  stream  near  Sisson,  Siskiyou  Co.  (38),  in  com- 
pany with  Riccardia  iatifrons,  CJiiloscypJius  polyanthus,  and  CepJia- 
lozia  media.  Only  a  few  scattered  gemmiferous  shoots  were  found, 
yet  we  feel  little  hesitancy  in  referring  them  to  Scapaina  curta, 
concerning  the  distribution  of  which  in  America  little  is  at  present 
known.  The  specimens  agree  closely  with  Rab.  Hep.  Eur.  93 
«  communis,  i.  forma  sterilis")  though  the  leaf-cells  are  some- 
what larger.  The  gemmae  are  broader  than  is  usual  in  European 
forms,  judging  from  the  specimens  accessible  ;  only  in  G.  &  R. 
Hep.  Eur.  382,  have  we  found  anything  to  equal  the  measure- 
ments given  above. 

7.  ScAPAXiA  HETEROPHYLLA  M.  A.  Howe,  BulL  Torr.  Bot.  Club, 
25:  iS^.pLjjd.  1898. 

Plants  obscurely  complanate,  dark  green,  often  bleaching  at 
apices  on  drying,  erect  or  ascending,  forming  compact  cushions  : 
stems  4-6  cm.  high  from  a  rhizomatous  base,  rigid,  fastigiately  and 
subdichotomously  branched,  brown,  becoming  almost  black,  nearly 
or  wholly  destitute  of  root-hairs,  denudate  below,  20-30  cells  in 
thickness :  leaves  scarcely  increasing  in  size  upward,  sometimes 
smaller  at  the  stem-apex,  the  upper  erecto-patent,  subimbricate, 
the  lower  approximate,  more  spreading,  often  with  squarrose  tips, 
all  strongly  undulate-crisped  both  when  moist  and  when  dry, 
bilobed  to  the  middle  or  bipartite,  complicate,  the  carina  acute 
or  somewhat  rounded,  but  never  winged,  commonly  bistratose 
toward  the  base  near  the  fold,  the  lobes  sometimes  almost  dis- 
united ;  now  and  then  with  an  unlobed  leaf  irregularly  interpo- 
lated, most  frequently  in  the  position  of  an  underleaf ;  margins  of 
the  lower  leaves  commonly  erose,  of  the  upper  entire  or  sparingly 
denticulate  ;  ventral  lobes  twice  as  large  as  the  dorsal  or  subequal, 
1.7-2.5  mm.  in  length,  .85-1.6  mm.  in  maximum  width,  broadly 
obovate,  elliptical,  or  obovate-oblong,  usually  decurrent,  mostly 
rounded-obtuse  at  apex  ;  dorsal  lobes  obtuse,  obHquely  and 
broadly  ovate  or  elliptical,  not  decurrent,  sometimes  appressed  at 
stem-apex,  especially  on  the  younger  shoots,  but  mostly  ascend- 
ing or  slightly  squarrose-reflexed  :  leaf-cells  generally  opaque,  with 
smooth  or  slightly  roughened  cuticle,  near  the  margin  subquadrate 
or  roundish-hexagonal,  16-28 near  the  base  oblong,  60— 90 /i  x 
25-30//;  trigones  indistinct  or  wanting:  remaining  parts  un- 
known. 

Exsicc.  Hep.  Am.  192. 

On  submerged  stones  in  a  cold  mountain  stream  (alt.  3000  ft.) 


156 


JUNGERMANNIACEAE 


in  company  with  CJdloscypJins  polyantJios  rividans  and  Pore  Ha 
rivnlaris,  Sisson,  Siskiyou  Co.,  July,  1894  (no.  34). 

The  stream,  which  is  formed  by  a  great  spring  beside  the  rail- 
way track  about  three  fourths  of  a  mile  north  of  the  village  of 
Sisson,  is  said  to  maintain  nearly  a  uniform  volume  throughout  the 
year,  and  as  plants  were  found  wholly  under  water  in  the  last  of 
July  and  the  first  of  August,  it  is  probable  that  their  submersion  is 
permanent. 

The  above  species  was  referred  at  first,  uncritically,  to  Scapania 
iindidata  and  listed  under  that  name  in  Erythea  (4  :  49.  1896). 
From  5*.  undidata,  however,  it  is  certainly  very  distinct  in  the  ob- 
scurely complanate  branches,  in  the  sometimes  3 -ranked,  often 
more  deeply  lobed,  erecto-patent,  never  alate.-carinate,  leaves,  with 
more  or  less  squarrose  tips,  and  in  the  obovate  rather  than  round- 
trapezoidal  ventral  lobes.  The  interpolated  unlobed  leaves  stand 
sometimes  in  about  the  general  position  of  dorsal  or  ventral  lobes,  but 
more  often  squarely  subtend  the  ventral  surface  of  the  stem.  They 
can  doubtless  be  explained  in  some  cases,  from  the  point  of  view 
of  ontogeny,  by  the  separation  of  the  normally  united  lobes,  but  in 
other  cases  this  hypothesis  seems  to  find  little  justification.  We 
have  noticed  one  or  two  ////w-lobed  leaves  out  of  hundreds  exam- 
ined and  in  such  the  complete  disjunction  of  the  most  ventral  lobe 
would  have  thrown  it  nearly  into  the  place  of  an  underleaf. 
When  the  leaf-lobes  are  subequal  it  is  often  difficult  to  distinguish 
between  the  dorsal  and  ventral  aspects  of  the  stem,  especially  if 
further  confused  by  the  presence  of  the  supernumerary  entire 
leaves.  From  stems  of  such  a  character  as  this,  however,  may 
spring  young  shoots  in  which  the  leaves  are  regularly  distichous 
and  acutely  complicate,  in  the  ordinary  Scapania  fashion,  with  the 
ventral  lobes  twice  the  size  of  the  dorsal.  In  the  axils  of  the  upper 
leaves  are  sometimes  to  be  found  numerous  short  clavate  para- 
physes,  unicellular  or  of  two  or  three  oblong  cells  in  a  lineal  series. 

We  are  indebted  to  Herr  K.  Loitlesberger  for  pointing  out  (/;/ 
lit.)  that  the  leaves  of  our  plant  are  two  cells  thick  in  the  middle,'* 
a  texture  which  he  has  never  found  in  submerged  Scapania  nndulata. 

It  is  possible  that  the  species  deserves  to  be  separated  gener- 
ically  from  Scapania,  but  in  absence  of  perianth  and  sporogonium, 
we  can  do  no  better  than  refer  it  to  a  genus  with  which  it  surely 
has  very  much  in  common. 


Radula 


157 


Plate  iio.    Scapania  heteroi'Hylla. 

I.  An  entire  plant,  i  the  natural  size. 

2  and  3.  Opposite  views  of  a  portion  of  the  stem,  showing  the  often  three-ranked 
leaves,  X  I5- 

4.  Dorsal  view  of  stem  and  leaves,  X  I5- 

5.  Ventral  view  of  the  same  object,  showing  an  unlobed  underleaf,  X  ^5- 

6.  The  leaf  with  lobes     a  "  and  "  a' already  shown  in  figures  4  and  5. 

7.  A  typical  underleaf,  X  15- 

8.  Outline  of  a  deeply  lobed  leaf,  X  I5- 

9.  Apex  of  a  branch  of  the  same  plant  from  which  figures  4  and  5  were  drawn  ; 
leaves  here  distichous,  X  ^S- 

10.  The  leaf  *'d"  from  the  foregoing,  X  ^5- 

II.  Ventral  view  of  a  leaf,  X  ^S- 

12.  Cells  from  near  base  of  leaf,  X  244. 

13.  Cells  from  the  apical  margin  of  one  of  the  upper  leaves,  X  244. 

14.  Paraphyses  from  axils  of  leaves,  X  244. 

15.  Cross-section  of  the  stem,  X  3^- 

33.  RADULA  Dumort././.  Comm.  Bot.    112.  1822. 

Marfme//ii/s  S.  F.  Gray,  p.p.  Nat.  Arr.  Brit.  PL  i  :  690.  1821. 
Stephanina  O.  Kuntze,*  Rev.  Gen.  PL  839.     1891.    Schiffn.  ; 
Eng.  &  Prantl,  Nat.  Pflanzenfam.  i'^-  113.  1895. 

*  Dr.  Otto  Kuntze  evidently  errs  somewhat  in  asserting  that  the  original  Radula 
of  Dumortier  ( 1822)  was  an  equivalent  of  the  Martinellius  of  S.  F.  Gray  (1821), 
inasmuch  as  Gray's  Alartinellius  contained  a  fourth  generic  element,  viz.,  the  Adelan- 
thus  of  Mitten  {^Martinellius  decipiens  S.  F.  Gray),  which  did  not  appear  in  the 
original  Radtcla,  even  though  it  was  given  a  place  in  the  Radula  of  183 1.  Now,  by 
the  "method  of  residues  " — which  seems  to  us  a  natural  and  logical  mode  of  settling 
the  descent  of  names  originally  applied  to  generic  composites — it  is  clear  that  after 
Dumortier,  under  the  name  of  Radula,  had  removed  the  modem  Radula,  Scapania, 
and  Plagiochila  elements  from  Gray's  tetrameric  genus  Martinellius,  the  Adelanthtts 
component  was  left,  which  would  thus  fall  heir  to  the  name  imposed  by  Gray.  This 
would  seem  to  us  a  particularly  happy  circumstance  in  view  of  the  fact  that  the  la'er 
name  Adelanthus  was  earlier  used  in  a  different  sense  by  Endlicher.  Again,  by  the 
''method  of  residues,"  the  name  Radula,  after  the  excision  in  1833  and  1835  of  the 
Scapania  and  Plagiochila  elements  from  the  Radula  of  1822,  descends  to  the  genus 
which  has  borne  the  name  Radula  with  nearly  every  hepaticologist  since  1833.  By 
''priority  of  place,"  too,  the  name  Radula  would  descend  in  the  same  way  inasmuch 
as  R.  complanata  was  always  the  first  species  mentioned  by  Dumortier  and  was  appar- 
ently always  more  or  less  clearly  in  his  mind  as  the  type  of  the  genus.  But  the  gen- 
eral application  of  the  "priority  of  place"  principle  would  result  in  attaching  S.  F. 
Gray's  name  Alartinellius  to  this  same  species  as  has  already  been  pointed  out  by  Dr. 
Carrington  and  others. 

The  use  of  Radulum  'by  Fries  in  1825  for  a  genus  of  fungi  cannot  disqualify 
Dumortier's  Radula  of  1822.  If  either  name  should  disappear,  it  is  the  Radulum  of 
Fries. 


158 


JUNGERMANNIACEAE 


Plants  moderately  large,  rarely  small  and  slender,,  green  or 
yellowish-green,  tinging  the  water  a  yellowish-green  when  soaked 
out,  in  most  species  creeping  and  forming  at  length  wide  de- 
pressed mats.  Stems  rather  loosely  pinnate  or  subbipinnate,  very 
rarely  dichotomous  ;  the  branches  infrafoliar  in  origin.  Leaves 
alternate,  incubous,  conduplicate-bilobed,  the  lobes  mostly  very 
entire,  the  ventral  lobe  smaller,  often  somewhat  inflated  near  the 
fold,  its  free  margin  nearly  always  appressed  to  the  dorsal  lobe  ; 
leaf-cells  small,  chlorophyllose,  often  containing  "oil-bodies." 
Root-hairs  always  springing  from  a  mammilliform  outpocketing  of 
the  ventral  lobe  near  its  base.  Underleaves  everywhere  wholly 
wanting.  Mostly  dioicous,  in  a  few  species  paroicous  or  autoicous. 
Androecia  in  dioicous  species  amentiform,  usually  terminal  on 
main  stem  or  the  principal  branches  ;  the  $  bracts  3-3  5  pairs, 
equitant,  with  subequal  lobes ;  antheridia  single  or  less  com- 
monly in  2's  or  3's.  Archegonia  several,  acrogenous,  very  rarely 
cladogenous,  the  matured  perianth  sometimes  pseudolateral 
through  the  development  of  a  "  subfloral  "  innovation.  5  bracts 
a  single  pair,  often  a  little  smaller  than  the  leaves,  the  somewhat 
enlarged  ventral  lobe  without  root-hairs.  Perianth  in  most  species 
strongly  compressed  dorso-ventrally,  in  a  few  subterete,  very  rarely 
carinate  or  plicate,  the  mouth  broad  from  the  beginning,  some- 
what bilabiate,  the  truncate  lips  entire  or  repand-crenate.  Calyptra 
narrowly  obovoid  or  elongate-pyriform,  rather  firm  and  subopaque, 
its  walls  composed  of  2  or  3  layers  of  cells. 

Capsule  mostly  oval-cylindrical,  2-3  times  as  long  as  broad, 
4-valved  to  the  base,  the  valves  bistratose,  the  longitudinal  walls 
of  the  exterior  cells  with  moniliform  or  nodular  thickenings,  the 
much  thinner  interior  cells  usually  with  delicate  transverse  striae. 
Seta  short  and  stout.  Spores  large,  subglobose  or  ellipsoidal, 
minutely  granulate-papillate  or  subechinulate.  Elaters  long  and 
slender,  obtuse,  closely  spiral. 

Key  to  tlie  Species. 

Dioicous ;  androecia  linear,  amentiform  ;  dorsal  leaf-lobes  obovate,  the  inner  margin 
adnate  to  stem,  not  surpassing  it,  leaf-cells  9-16  //  ;  spores  subglobose  or  ellipsoidal, 
48-60  ^  in  maximum  diameter.  I.  J\.  Bolamicri. 

Paroicous ;  dorsal  leaf-lobes  quadrate-orbicular,  the  inner  margin  surpassing  the  stem, 
leaf-cells  16-24// ;  spores  subglobose,  30-38 //.  2.  R.  complatiata. 

I.    Radula  Bolaxderi  Gottsche ;  Steph.   Hedwigia,   23:  145. 

1884. 

Radiila  spicata  Aust.  Bull.  Torr.  Bot.  Club,  6:  19.  1875. 
Pearson,  List  Can.  Hep.  4.  7.  1890.  Not  Radiila  spicata  Mitt. 
Bonplandia,  10  :  19.  1862. 


Radula 


159 


Light-green,  slender,  creeping,  becoming  intricately  caespitose: 
stems  1—2  cm.  long,  pinnately  or  somewhat  bipinnately  branched, 
the  branches  mostly  short :  leaves  contiguous  or  laxly  imbricate, 
patent  or  erecto-patent,  subascending,  the  obtuse  carina  arcuate 
and  decurrent ;  dorsal  lobes  obovate,  .35-85  mm.  long,  .27-. 6 5 
mm.  wide,  nearly  plane  or  slightly  concave,  almost  longitudinally 
adnate  to  the  stem,  which  the  inner  margin  does  not  surpass,  apex 
very  obtusely  rounded  ;  ventral  lobes  of  the  leaves  on  the  younger 
ramuli  subequal  to  the  dorsal  lobes  or  one  half  as  large,  those  of 
the  adult  leaves  one  third  the  size  of  the  dorsal,  inflated,  rhom- 
boidal -ovate,  the  obtuse  or  subacute  apex  more  or  less  appressed 
to  the  dorsal  lobe,  the  inner  margin  almost  wholly  adnate  to  the 
stem  and  scarcely  incumbent,  the  outer  margin  obliquely  truncate; 
leaf-cells  9- 16 /v.,  the  walls  scarcely  thickened  at  the  angles: 
dioicous  :  androecia  terminal  on  main  stem  or  principal  branches^ 
numerous,  linear,  3-7  mm.  long,  .5-6  mm.  wide,  usually  directed 
downward;  ^  bracts  10—35  pairs,  very  closely  imbricated, 
strongly  ventricose,  the  lobes  equal  :  9  bracts  of  about  the  same 
size  as  the  leaves,  the  lobes  subequal:  perianth  obconic,  1.2-2 
mm.  long,  .75—1.2  in  greatest  width,  bilabiate,  the  lips  entire  :  cap- 
sule prolate-ellipsoidal,  exserted  1-2  mm.,  the  valves  about  i  mm. 
long,  transverse  striae  of  the  inner  cells  very  obscure  or  wanting  ; 
spores  subglobose  or  ellipsoidal,  48-60  (i  in  maximum  diameter, 
minutely  and  densely  echinulate-papillate  ;  elaters  150-200 /v.  x 
6-8  (1. 

On  the  trunks  of  trees  {Alnus,  Umbelhilaria,  etc.).  Com- 
mon in  the  Coast  Range  Mountains — at  least,  from  San  Francisco 
northward  ;  Mill  Valley,  Marin  Co.;  Cazadero,  Sonoma  Co.;  Men- 
docino (549)  ;  Eureka  (918)  and  near  McBride's,  Mad  River 
(105  i),  Humboldt  Co.    Vancouver  Island  (Macoun).* 

2.  Radula  complanata  (L.  )  Dumort.  Comm.  Bot.  112.  1822. 

Jungerniannia  complanata  L.  Sp.  PI.  1133.  1753- 
Mostly  yellowish-green,  rather  flaccid,  closely  and  radially 
creeping,  finally  forming  depressed  mats  :  stems  1-6  cm.  long,  ir- 
regularly pinnate  :  leaves  closely  imbricate,  patent,  the  margins 
occasionally  gemmiferous;  dorsal  lobes  quadrate-orbicular,  .5- 


*  There  seems  to  be  little  or  nothing  in  the  description  and  figure  of  Herr  Stephani's 
apparently  wholly  sterile  Radula  a?rfica  (from  Chlowak  and  Chilcoot,  Alaska. — 
Bot.  Jahrb.  8  :  98.  //.  j>.  /.  //.  1886)  to  distinguish  it  from  the  juvenile  condition  of 
Radula  Bolanderi. 


160 


JUXGERMANNIACEAE 


1.6  mm.  long  by  .4-1.5  mm.  wide,"^  concave  ventrally  with  the 
obtuse  apex  more  or  less  deflexed,  or  nearly  plane,  the  inner 
margin  surpassing  the  stem  ;  carina  slightly  arcuate  ;  ventral  lobe 
about  one  fourth  the  size  of  the  dorsal,  subquadrate,  now  and  then 
a  little  elongated  in  a  direction  parallel  to  the  carina,  appressed  to 
the  dorsal  lobe  throughout  or  lightly  inflated  toward  the  base, 
subobtuse  or  acute,  the  inner  superior  angle  free  and  somewhat 
incumbent  upon  the  stem  ;  leaf-cells  16-24 /v.,  the  walls  distinctly 
thickened  at  the  angles  :  paroicous  :  perigonial  leaves  2-4  pairs, 
immediately  subtending  the  9  bracts,  ventricose  at  base,  the  ven- 
tral lobes  obtuse,  Y^-yi  the  size  of  the  dorsal  :  ventral  lobes  of 
the  9  bracts  subequal  to  the  dorsal  or  one  half  as  large,  obtuse  : 
perianth  obconic  or  elongate-obconic,  1.8-3  by  .7-1.2  mm. 

in  maximum  width,  strongly  compressed,  bilabiate,  the  lips  entire 
or  slightly  repand  :  capsule  prolate-ellipsoidal  or  obovoid,  .9—1.3 
mm.  X  .45-55  mm.,  exserted  about  i  mm.;  spores  subglobose, 
30-38  fj.,  finely  granulate-papillate;  elaters  160— 210  u.  x  6—9/7.. 

On  branches  and  trunks  of  trees  in  moist  or  deeply  shaded 
places  (the  species  inhabits  rocks  also,  but  has  not  yet  been  found 
in  such  situations  in  California);  Berkeley,  Fruit  Vale  (Miss  Byx- 
bee);  near  Olema,  Marin  Co.;  near  Mendocino  (707);  Eureka 
(917)  and  Blue  Lake  (1015),  Humboldt  Co.;  collected  in  California 
by  Dr.  Kellogg  and  probably  also  by  Dr.  Bolander,  though  we 
have  seen  no  specimens  of  the  latter's  collecting. 

It  seems  to  us  to  be  doubtful  whether  Radnla  Krausei  Steph. 
(Bot.  Jahrb.  8:  97.  pL  3.  /.  10.  1886) — at  least  as  represented 
in  Hep.  Am.  149,  a  portion  of  which,  we  understand,  was  sub- 
mitted to  Herr  Stephani  before  its  distribution — can  be  satisfac- 
torily distinguished  from  R.  complanata.  If  we  are  not  mistaken, 
the  supposed  distinctive  characters  of  Radnla  Krausei  pass  quite 
imperceptibly  into  those  of  the  Pacific  coast  R.  complanata,  which, 
we  believe,  cannot  be  well  separated  from  the  European  plant. 

34.  PORELLA  L.  Sp.  PI.  1106.     1753.     Ex  Dill.  Hist.  Muse. 
459.  //.  68.    1741.   Lindb.  Act.  Soc.  Sci.  Fenn.  9:  335.  1869. 

Raddi,  Atti  Soc.  Ital.  Sci.  Mod.  18:— (7):  1818. 
Mem.  Mat.  e  Fis.  Soc.  Ital.  Sci,  Mod.  18:  18.  1820. 

*The  dorsal  leaf-lobes  of  Radnla  complanata  are  said  by  Herr  Jack  (Flora,  64  : 
356.  1881 )  to  be  always  broader  than  long,  but  we  do  not  find  them  always  so  even  in 
a  specimen  collected  near  Salem  by  Jack  and  Leiner.  In  the  Californian  specimens 
here  referred  to  K.  complanata — as  well  as  in  the  North  Ameritan  plant  in  general — the 
leaves  are  ordinarily  a  little  longer  than  broad. 


PORELLA 


161 


Antoiria  Raddi,  Atti  Soc.  Ital.  Sci.  Mod.  i8:— (8):  1818. 
Mem.  Mat.  e  Fis.  Soc.  Ital.  Sci.  Mod.  18:  19.  1820. 

Cavendishia  S.  F.  Gray,  Nat.  Arr.  Brit.  PI.  i  :  689.     182 1. 

Madothcca  Dumort.  Comm.  Bot.  iii.  1822.  Nees,  Natur- 
gesch.  Eur.  Leberm.  3:  157.  1838. 

Plants  large,  dark-green  to  yellowish-brown,  mostly  somewhat 
regularly  bi-  or  tri-pinnate,  rarely  subsimple  ;  rqot-hairs  in  tufts  at 
the  base  of  the  underleaves,  usually  sparingly  developed.  Leaves 
very  deeply  2-parted ;  the  dorsal  lobes  large,  incubous,  obliquely 
orbicular-ovate  to  oblong,  entire,  repand  or  somewhat  dentate ; 
ventral  lobes  much  smaller  than  the  dorsal,  sometimes  nearly  dis- 
crete, ovate,  lingulate,  oblong,  linear,  or  lanceolate,  nearly  parallel 
with  the  stem,  entire  or  toothed,  margins  plane  or  revolute.  Un- 
derleaves large,  somewhat  similar  in  form  to  the  ventral  lobes  but 
usually  broader,  entire  or  dentate,  often  long  decurrent  on  both 
sides.  Antheridia  spherical,  very  short-stalked,  single  in  the  axils 
of  saccate,  densely  imbricate,  nearly  equally  bilobed  opposite  leaves, 
these  connate  with  the  underleaves  and  forming  short,  lateral,  oval, 
to  linear-oblong  spikes.  Archegonia  generally  numerous,  terminal 
on  very  short  (most  rarely  a  little  elongated)  lateral  branches. 
Perianth  oval  to  obovate,  flattened  dorso-ventrally  toward  the 
mouth,  from  a  more  or  less  obconical  base,  much  longer  than  the 
bracts,  two-lipped  after  elevation  of  the  capsule  or  sometimes  irreg- 
ularly torn,  mouth  ciliate,  dentate,  or  subentire.  O  bracts  usually 
a  single  pair  with  a  single  bracteole  in  addition  to  the  underleaf  at 
the  base  of  the  branch,  the  latter  underleaf  united  with  the  sub- 
tending cauline  leaf  and  functioning  as  its  ventral  lobe,  or  free, 
leaving  the  cauline  leaf  unlobed.  Capsule  spherical  to  ovoid-oblong, 
on  a  short  seta,  yellowish-brown,  opening,  usually  not  quite  to  the 
base,  by  four  often  irregularly  split  valves  ;  cell -walls  of  the  valves 
mostly  with  irregular  nodulose  thickenings.  Elaters  commonly  2- 
(1-3-)  spiral  ;  spores  several  times  broader,  more  or  less  echinulate. 

Key  to  the  Species. 

Stems  subsimple  or  somewhat  fasciculately  branching,  short,  tumid  ;  underleaves  cau- 

date-lacinulate  at  base  ;  perianth-mouth  ciliate.  i.  P.  Bolanderi. 

Stems  more  or  less  regularly  1-3-pinnate. 
Usually  shining. 

Ventral  lobes  more  or  less  spurred  outwardly  at  base,  mostly  linguiform  or 

ovate-oblong,  margins  plane  or  lightly  recurved.  3.  P.  Roellii. 

Ventral  lobes  regularly  rounded  outwardly  at  base,  ovate,  margin  recurved. 

4.  P.  naviadaris. 

Usually  dull  ;  rather  flaccid,  dorsal  lobes  subimbricate,  ventral  lobes  long-decurrent, 
underleaves  distant.  2.  P.  rivularis. 


162 


JUNGERMANNIACEAE 


I.  PoRELLA  BoLANDERi  (Aust.)  Peai'soD,  List.  Can.  Hep.  7.  1890. 
[Excluding  specimens  cited  (?)] 

Madotheca  Bolaiideri  Aust.  Bull.  Torr.  Bot.  Club,  3  :  14.  1872. 

Dark-  or  yellowish-green,  dull:  stems  1.5-6  cm.  long,  sub- 
simple  or  with  a  few  somewhat  fasciculately  disposed  obtuse  tumid 
branches,  often  subpendulous,  more  or  less  vaulted  or  flexuous 
when  dry  :  dorsal  lobes  of  leaves  densely  imbricate,  appressed  or 
subsquarrose,  dimidiate-ovate  to  oblong,  1.5-2.9  mm.  x. 67-1. 8 
mm.,  sometimes  considerably  narrowed  toward  the  obtuse  apex, 
slightly  decurved  when  moist,  circumvolute-deflexed  in  drying, 
rather  distinctly  margined  by  somewhat  inflated  subrectangular 
cells,  the  inferior  margin  more  or  less  undulate,  often  narrowly  in- 
flexed,  the  superior  repand  or  here  and  there  caudate-dentate,  the 
base  long  drawn  out  and  projecting  beyond  the  stem,  trigones 
small ;  ventral  lobes  and  underleaves  approximate  or  more  often 
imbricate,  sometimes  entirely  conceahng  the  stem  ;  ventral  lobes 
ovate-lanceolate  to  linear-lanceolate,  rarely  almost  subulate,  acute 
or  occasionally  somewhat  obtuse,  subfalcate,  canaliculate,  sHghtly 
twisted,  long-decurrent,  nearly  discrete,  .4-1  mm.  X.08-.4  mm., 
about  ^  as  wide  as  the  underleaves,  length  -|— |  the  width  of  the 
dorsal,  undulate-repand  above,  sparingly  caudate-lacinulate  on  in- 
ner side  at  base,  often  also  on  the  outer:  underleaves  ovate-lingu- 
late  to  oblong,  a  little  wider  than  the  stem,  apex  obtuse,  subacute, 
rarely  emarginate  or  slightly  cleft,  often  deflexed,  margins  plane 
or  recurved,  undulate-repand,  long-decurrent,  the  wings  with  com- 
monly 2  or  3  cauda-like  laciniae  on  either  side  toward  the  base  : 
dioicous  :  $  spikes  oblong  to  almost  linear,  2-4  mm.  in  length  :  9 
branch  short ;  dorsal  lobes  of  9  bracts  subobtuse  or  acute,  the  ven- 
tral acute  or  often  subulate -pointed,  margins  of  both  subentire  or 
denticulate  above,  ciliate-caudate  at  base,  bracteole  large,  ovate, 
usually  acute,  denticulate  above,  caudate-lacinulate  below  ;  under- 
leaf  subtending  9  branch  acutely  emarginate  or  bifid  ;  perianth 
broadly  ovate  from  a  shortly  obconic  base,  somewhat  compressed, 
lightly  undulate -plicate  dorsally,  often  furnished  ventrally  with  2 
or  three  rarely  winged  carinae,  narrowed  at  the  ciliate,  subtruncate, 
deeply  bilabiate  mouth  :  capsule  oval  or  oval-oblong,  exserted  by 
about  its  own  length  ;  spores  29-40  fi,  minutely  echinulate  ;  elaters 
180-310  nx  10-12  fj.,  2-  (3-)  spiral. 

Exsicc.  Hep.  Am.  31. 

On  stones,  under  shelving  rocks,  and  on  the  bark  of  living 
trees,  especially  of  Quercns  agrifolia. 

First  collected  by  Dr.  Bolander.  Berkeley  (7,  1189,  1190, 
1 197),  Fruit  Vale  (1219),  San  Lcaindro  (Underwood);  Mitchell 


PORELLA 


163 


Canon,  Mt.  Diablo  (1195,  1196,  12 18);  Ukiah  (756),  Pieta  (797), 
Mendocino  Co.;  near  Hay  Fork  (i  108),  Trinity  Co.;  Oak  Run  and 
Cow  Creek,  Shasta  Co.  (M.  S.  Baker  and  F.  P.  Nutting)  ;  Nord- 
hoff,  Ventura  Co.  (Miss  Jacqueline  K.  Newton) ;  Pasadena  and 
Elsinore  (A.  J.  McClatchie). 

Type  in  herb.  Pearson,  Knutsford,  Cheshire,  England.  We 
have  seen  no  specimens  of  Porclla  Bolanderi  from  any  station  out- 
side of  California.  Can.  Hep.  no.  10  (distributed  as  P.  Bolanderi) 
is,  so  far  as  we  have  seen  it,  Porella  rividaris — in  one  pocket 
mixed  with  P.  naviadaris.  Can.  Hep.  no.  1 1  (issued  as  PorcUa 
Bolanderi  var.),  from  Mt.  Benson,  Vancouver  Island,  is  Porella 
Roellii  Stephani. 

In  moist,  densely  shaded  places,  Porella  Bolanderi  assumes  a 
more  lax,  flaccid  habit,  resembling  certain  forms  of  P.  rividaris. 
Such  conditions,  however,  when  the  characteristic  ciliate  perianths 
are  wanting,  can  usually  be  distinguished  from  any  state  of  P. 
rividaris  by  the  more  oblong,  more  distinctly  marginate,  dorsal 
leaf-lobes,  and  by  the  longer  and  narrower  ventral  lobes  and  un- 
derleaves,  which  are  more  pronouncedly  caudate  at  the  base. 

Plates  hi  and  112.    Porella  Bolanderi. 

I -  3.  Plants,  9  1  natural  size. 

4.  Plant,  $ ,  natural  size. 

5.  Portion  of  plant,  ventral  view,  X  12. 

6.  Mature  perianth  and  part  of  stem,  ventral  view,  X  12. 

7.  Young  perianth,  bracteole,  bracts,  etc.,  ventral  view,  X  ^2. 

8.  Half  of  mouth  of  young  perianth,  X  53" 

9  and  10.  Transverse  sections  of  perianth,  the  lower  side  ventral,  X  23. 

II-  13.  Cauline  leaves,  showing  the  ventral  lobe,  X  23. 
14.  Leaf  and  ventral  lobe,  X  12. 

15-17.  Leaves,  dorsal  view,  X  ^2.  ' 
18  and  19.  Underleaves,  X  23. 
20.  Leaf-cells,  X  305- 

Figs.  I  and  2  from  specimen  collected  in  Fruit  Vale  by  Miss  Byxbee  ;  3,  5»  ^4? 
16,  17,  and  20,  from  no.  1190  (Berkeley)  ;  4,  from  no.  1189  (Berkeley)  ;  6  and  12, 
from  no.  1196  (Mt.  Diablo)  ;  7-10,  13,  15,  and  19,  from  no.  7  (Berkeley)  ;  li  and  18, 
from  no.  1 195  (Mt.  Diablo). 

2.  Porella  rivularis  (Nees)  Trevis.  Mem.  r.  1st.  Lomb.  III.  4  : 

407.  1877. 

Madotheca  rividaris  Nees,  Naturgesch.  Eur.  Leberm.  3  :  196. 
1838. 


164 


JUNGERMANNIACEAE 


Porclla  dcntata  Lindb.  Act.  Soc.  Sci.  Fenn.  9  :  342.  1869, 
Dull,  or  very  rarely  a  little  polished,  opaque,  commonly  dark 
green,  sometimes  yellowish,  mostly  soft  and  flaccid  :  stems  irregu- 
larly pinnate  or  subdichotomous,  3-10  cm.  long,  prostrate  or  as- 
cending, forming  loose  or  more  dense  mats,  branches  somewhat 
obtuse,  scarcely  diminishing  in  width  toward  apex  :  dorsal  lobes  of 
leaves  usuallysubimbricate  or  approximate,  sometimes  distant,  rarely 
closely  incubous,  obliquely  ovate  to  orbicular-ovate,  rounded-ob- 
tuse, 1-2  mm.  X  .8-2  mm.,  entire  or  subdenticulate,  flat  or  slightly 
concave  beneath,  only  a  little  decurved  at  the  apex,  trigones  mostly 
small  ;  ventral  lobes  small,  obliquely  ovate,  acute,  .35—7  mm.  x 
.12-.4  mm.,  length  -i- 1-  the  width  of  the  dorsal,  about  one  half  as 
broad  as  the  underleaves,  margins,  especially  the  outer,  for  the 
most  part  broadly  revolute,  often  giving  the  lobe  a  twisted  appear- 
ance, long  decurrent,  usually  dentate  or  subciliate  at  base  inter- 
nally and  sometimes  unindentate  externally  but  scarcely  spurred  : 
underleaves  distant,  quadrate,  orbicular  to  broadly  ovate,  about 
twice  the  width  of  the  stem,  apex  rounded-obtuse,  sometimes  re- 
flexed,  margins  repand-undulate,  very  long  decurrent,  the  wing 
sometimes  exceeding  the  free  portion  in  length  and  usually  acutely 
dentate  or  subciliate  :  dioicous  :  $  spikes  oval  to  oblong,  1.5—2.5 
mm.  in  length  :  9  branch  short  :  ventral  lobes  of  the  single  pair  of 
9  bracts  acute  or  subobtuse,  entire  or  repand,  the  dorsal  obtuse, 
bracteole  ovate-linguiform,  repand  ;  perianth  ovate,  with  lateral 
margins  deflexed,  deeply  bilabiate,  the  lips  subentire  or  repand- 
dentate,  usually  plane  :  spores  27—45  papillate-echinulate  ;  ela- 
ters  180-290  11  X  8-10/^,  rather  obtuse,  2(3-)  spiral. 

Exsicc.  Hep.  Am.  150  (as  Porclla  Rocllii,  wdiY.),  194A,  and 
194B. 

On  moist  rocks,  stones  in  streams,  and  bases  of  trees  in  densely 
shaded  places.  Widely  distributed  and  extremely  variable  accord- 
ing to  habitat. 

Near  Berkeley  (7d,  7e,  19,  1175,  1182,  12 14);  Mill  Valley  and 
Olema  (1178);  Cazadero  ( 1 1 79) ;  Mendocino  (714,  718);  Ukiah 
(745);  Blue  Lake  (1177);  Russ  &  Graham's  Ranch  (1084) 
Humboldt  Co.;  Sisson,  Siskiyou  Co.,  (1176);  Cow  Creek  and 
Burney  Falls,  Shasta  Co.  (M.  S.  Baker  &  F.  P.  Nutting)  ;  Yose- 
mite  Valley  region  (C.  M.  Cooke,  Jr.).  Collected  in  California 
by  Dr.  Bolander  also,  the  exact  localities  unknown,  and  by  Mr. 
S.  B.  Parish  in  the  southern  part  of  the  state. 

Our  determination  is  based  upon  the  drawing  (accompanying 
G.  &  R.  Hep.  Eur.  no.  371)  of  the  original  plant  from  the  bed  of 


PORELLA 


165 


the  river  Bober,  near  Hirschberg,  Silesia,  and  upon  European 
specimens  and  the  descriptions  of  authors. 

The  name  dcntata,  appHed  by  Hartman  in  1832  (Skand.  Fl. 
354,  2d  ed.)  to  what  he  considered  a  variety  of  Jiingerniannia 
platypJiylla  was  taken  up  for  the  above  species  by  Lindberg  in 
1869,  but  was  abandoned  by  him  ten  years  later  without  explana- 
tions. Hartman' s  description  of  his  variety  dentata  is  quite  inade- 
quate for  its  proper  identification  and  as  we  have  failed  in  our 
efforts  to  see  his  original  specimens,  if  such  exist,  we  prefer  to  ad- 
here to  the  first  name  which  was  accompanied  by  an  intelligible 
diagnosis. 

3.  PoRELLA  RoELLii  Stcph.  Bot.  Centralb.  45:  203.  1891. 

Green  or  yellowish-brown,  usually  shining  :  stems  procumi- 
bent,  rather  flaccid,  densely  depressed-caespitose,  4-8  cm.  long, 
subdichotomous  below,  the  primary  branches  for  the  most  part 
simply  and  remotely  pinnate,  branchlets  short,  often  attenuate- 
deflexed  :  dorsal  lobes  of  leaves  densely  imbricate,  appressed,  ob- 
'liquely  ovate,  patent-divergent,  .85-T.7  mm.  x  .65-1.5  mm.,  apex 
narrowed,  subtruncate,  obtuse,  or  most  rarely  apiculate,  inferior 
margin  more  or  less  undulate-crisped,  the  superior  repand,  cells  at 
basal  angles  slightly  smaller,  all  with  evident  trigones  ;  ventral 
lobes  ovate,  linguiform  or  ovate-oblong,  usually  much  narrowed 
toward  the  obtuse  or  subacute  apex,  suberect  or  patent,  scarcely 
connate  with  the  dorsal,  commonly  about  ^  as  wide  as  the  un- 
derleaves,  length  ^  the  width  of  the  dorsal,  somewhat  concave 
ventrally,  margins  plane  or  slightly  recurved,  a  little  decurrent, 
very  rarely  dentate  above,  more  or  less  strongly  spurred  at  the 
base,  especially  at  the  outer  angle,  the  spur  entire  or  dentate, 
rarely  subciliatc :  underleaves  approximate,  ovate-linguiform, 
rounded-obtuse,  the  margins  recurved,  long-decurrent,  entire  or 
most  rarely  subdentate,  the  wings  sometimes  crisped  :  dioicous  : 
^  spikes  1-2  mm.  long  :  9  branch  somewhat  elongated,  bearing 
3-8  leaves  (or  "bracts'')  (usually  2  pairs)  nearly  similar  to  the 
cauline  ;  inmost  bracts  a  little  larger,  the  lobes  subacute,  entire, 
repand-dentate,  or  sparingly  denticulate,  bracteole  ovate,  subentire 
or  denticulate  ;  perianth  large,  somewhat  goblet-shaped  or  broadly 
obovate,  undulate-concave  ventrally,  here  and  there  inflated, 
scarcely  narrowed  or  lobed  at  the  wide  truncate  dentate  mouth, 
the  oral  margins  plane  or  slightly  deflexed  at  the  sides. 

Under  shelving  rocks  and  on  moist  shaded  cliffs,  rarely  on 
tree-trunks.   Mill  Valley  (1168)  and  ]\It.  Tamalpais  (1171),  Marin 


166 


JUXGERMANXIACEAE 


Co.;  near  Cazadero  (i  i/o),  Sonoma  Co.;  Ukiah  (759)  and  Navarro 
(Miss  Edith  S.  Byxbee),  Mendocino  Co.;  Blue  Lake  (994)  and 
Deer  Creek  (1068,  1073,  1231),  Humboldt  Co.;  Hay  Fork  (1109). 
Trinity  Co. 

Type  from  Kitchelos  Lake,  Washington  (Roell,  June  12,  1888) 
— in  herb.  Stephani,  Leipsic.  Type  duplicate  in  herb.  Under- 
wood. 

Porella  RocUii  is  closely  related  to  the  forms  of  the  European 
P.  larrigata  with  obtuse  dorsal  lobes  and  subentire  ventral  lobes 
and  underleaves,  as  represented,  for  example,  in  Carrington  and 
Pearson's  no.  275  Hep.  Brit.  Exsicc,  from  Scotland.  This  is  es- 
pecially true  of  British  Columbia  specimens  (Macoun  :  Cascades, 
Yale,  May,  1875  ;  no.  63,  near  Victoria,  May  30,  1893),  and  of 
our  no.  1068.  These  latter  differ,  however,  from  P.  laevigata  in 
the  smaller,  narrower,  more  pointed,  and  more  strongly  calcarate 
ventral  lobes,  and  in  the  usually  more  slender  fronds.  They  are 
forms  like  these,  we  take  it,  that  have  been  referred  by  IMr.  Pear- 
son *  to  P.  laevigata,  but  in  the  predominating  forms  in  Washing- 
ton, Oregon,  and  California  the  plant  is  much  less  suggestive  of  P. 
laevigata,  and  we  prefer  to  maintain  Stephani' s  species,  and  to  as- 
sociate with  it  the  British  Columbia  specimens  alluded  to  and  our 
no.  1068,  even  though,  as  must  be  admitted,  they  make  a  near 
approach  to  certain  conditions  of  the  European  plant.  Perianths 
occur  only  in  our  nos.  994  and  1231  ;  they  appear  to  differ  but 
slightly  from  those  borne  by  a  specimen  of  P.  laevigata  from 
Teneriffe,  which  we  owe  to  the  kindness  of  Matthew  B.  Slater, 
Esq.,  of  Malton,  England. 

Plates  113  and  114.    Porella  Roellu. 
I.   9  plaint,  natural  size. 
2  and  3.  Sterile  plants,  natural  size. 

4.  $  plant,  natural  size. 

5.  Portion  of  plant,  ventral  view,  X  12. 

6-8.  Cauline  leaves,  showing  ventral  lobes,  X  23. 
9.  Rameal  leaf,  ventral  view,  X  23. 
10  and  II.  Underleaves,  X  23. 

12  and  13.     9  bi'anches,  with  bracts  and  mature  perianths,  X  ^2. 

14.  Outline  of  cauline  leaves,  dorsal  view,  X  ^2. 

15.  Leaf-cells,  X  305- 

16  and  17.  Leaves,  viewed  dorsally,  X  12. 


*  List  of  Canadian  Ilepiticae,  7.  1890. 


PORELLA 


167 


l8.  About  one  fourth  of  perianth-mouth,  X  53- 

Figs.  I,  12,  13,  and  18,  drawn  from  no.  994  (Blue  Lake,  Humboldt  Co.)  ;  2, 
from  no.  759  (Ukiah)  ;  3,  from  no.  1109  (Hay  Fork,  Trinity  Co.)  ;  4,  from  no.  I168 
(Mill  Valley)  ;  5,  6,  8-10  and  14-17,  from  a  portion  of  the  original  material  collected 
by  Roell  at  Kitchelos  Lake,  Washington,  and  communicated  to  Professor  Underwood 
by  Herr  Stephani  ;  7  and  li,  from  no.  1068  (Deer  Creek,  Humboldt  Co.). 

4.  PoRELLA  XAVicuLARis  (Lehm.  &  Lindenb.)  Lindb.  Act.  Soc. 
Sci.  Fenn.  9:  337.  1869. 

Jtingcrniannia  navicularis  Lehm,  &  Lindenb.  ;  Lehm,  Pugill,  6  : 
38.  1834. 

Madotlieca  navicularis  Xees,  /.  /.  G.  L.  N,  Syn.  Hep.  277. 
1845. 

Madotheca  Doiiglasii  Tayl.  Lond.  Jour.  Bot.  5  :  379,  1846. 

Madoihcca  Californica  Hampe,  in  herb,  Sulhvant,  fide  Austin, 
Hep,  Bor.-x-\m,  91, 

Shining  or  more  rarely  dull,  brownish-yellow,  large  :  stems 
somewhat  regularly  bipinnate,  4-12  cm,  long,  procumbent-caespi- 
tose,  or  subpendulous  with  apices  ascending  when  moist,  branch- 
lets  gradually  narrowed,  subacute,  convex  on  drying  and  slightly 
deflexed  :  dorsal  lobes  of  leaves  densely  imbricate,  appressed,  for 
the  most  part  closely  wrapped  about  the  stem  when  dry,  obliquely 
orbicular-ovate  to  oblong-ovate,  1,2-2.5  mm.  x  1-1.8  mm., 
rounded-obtuse,  apex  decurved,  subcucullate,  inferior  margin 
slightly  undulate-crisped,  the  superior  decurrent,  its  wing  some- 
times laciniate-crispate,  otherwise  very  entire,  the  inferior  basal 
angle  composed  of  numerous  small  thick-walled  cells  the 
diameter  of  the  remainder,  trigones  conspicuous  throughout ;  ven- 
tral lobes  ovate,  obtuse,  most  rarely  subacute,  regularly  rounded 
outwardly  at  base,  .5-1  mm.  x  .33-75  mm.,  a  little  smaller  than 
the  underleaves,  length  somewhat  more  than  the  width  of  the 
dorsal,  margins  entire,  recurved,  decurrent,  apex  often  deflexed  : 
underleaves  approximate,  quadrate-oblong,  rounded-obtuse,  mar- 
gins entire,  recurved,  long-decurrent,  apex  occasionally  deflexed  : 
dioicous  :  ^  branches  oval  to  oblong,  iy2—2y2  mm.  in  length  : 
9  branch  short,  lobes  of  the  single  pair  of  bracts  minutely  dentic- 
ulate or  entire,  the  ventral  usually  acute,  the  dorsal  obtuse,  brac- 
teole  wide  ;  perianth  broadly  obovate  from  an  obconic  somewhat 
inflated  base,  scarcely  narrowed  above,  deeply  bilabiate,  the  lips  at 
first  subciliate-denticulate,  later  obsoletely  crenulate-dentate  or 
nearly  entire,  strongly  revolute,  giving  perianth  the  appearance  of 
being  squarely  or  obliquely  truncate  :  capsule  ovoid,  yellowish- 
brown,  exserted  by  about  its  own  length  ;  spores  50-85  /v.,  echinu- 


168 


JUNGERMAXNIACEAE 


late,  bright  yellowish-green;  elaters  275-325  a  x  g-io  ji,  obtuse, 
2-  (3-)  spiral. 

Exsicc.  Hep.  Bor.-Am.  91. 
Hep.  Am.  30. 

Common  on  trunks  and  branches  of  trees  and  more  rarely  on 
rocks  throughout  the  Pacific  Coast  fi-om  California  to  Alaska  ;  as 
far  east  as  Idaho  (Leiberg) ;  Mexico  {fide  Gottsche). 

Mill  Valley  (1215),  Olema  (1216,  1217);  Duncan's  Mills 
(1180),  Cazadero  (1181),  Sonoma  Co.;  Mendocino  (563,  564), 
Navarro  (Miss  Edith  S.  Byxbee),  Ukiah  (776) ;  Mission  Hills,  San 
Francisco  (Dr.  Kellogg)  ;  Lake  Pilarcitos,  San  Mateo  Co.  (W.  C. 
Blasdale)  ;  Santa  Cruz  Mts.  (Underwood).  Collected  in  California 
also  by  the  botanists  of  Beechey's  expedition  (in  herb.  Taylor),  by 
Dr.  Bolander  and  others. 

The  original  specimens  were  collected  by  Menzies  on  the  wes- 
tern coast  of  North  America,  exact  locality  unknown.  Linden- 
berg's  specimen  of  Jnngermannia  navicularis  n.  sp."  is  preserved 
in  the  Naturhistorisches  Museum  at  Vienna.  We  are  indebted  to 
Prof  Dr.  G.  von  Beck  for  the  privilege  of  seeing  a  portion  of  this 
type.  The  European  plants  referred  by  some  authors  to  Porella 
navicularis  probably  all  belong  with  P.  platyphylla  or  P.  laevigata. 
This  species  in  a  fertile  condition  can  always  be  very  easily  distin- 
guished from  any  simulating  form  of  P.  platypJiylla  by  the  broadly 
obovate  perianth,  scarcely  narrowed  at  the  mouth,  and  by  the  twice 
larger  diameter  of  the  spores.  When  sterile,  if  more  obv^ious 
characters  fail,  the  numerous,  small,  thick-walled  cells  filling  the 
inferior  basal  wing  of  the  dorsal  lobe  are  of  importance  ;  these  are 
yi-y2  the  diameter  of  the  cells  in  the  middle  of  the  lobe  while  in 
P.  platyphylla  the  cells  are  nearly  of  a  uniform  size  throughout  the 
lobe  or  slightly  smaller  at  the  basal  margin. 

In  two  cases,  we  have  observed  a  peculiar  modification  of  Por- 
ella navicularis  due,  we  believe,  to  the  influence  of  an  epiphytic 
fungus.  The  leaves  were  here  bilobed  rather  than  bipartite,  the 
union  of  the  lobes  resembling  that  of  Lcjcunia  or  Radida.  The 
ventral  lobes  were  shortened  so  that  their  longer  axis  was  nearly 
parallel  to  that  of  the  dorsal  lobe  and  were  conspicuously  inflated 
as  in  most  of  the  Lejeuniae.  The  underleaves  were  also  shortened 
and  often  ventricose.     In  fact,  when,  as  sometimes  happened,  a 


Frullania 


169 


sterile  plant  was  thus  transformed  throughout,  there  was  Httle,  out- 
side of  the  axillary  branching,  to  distinguish  it  from  the  holostipous 
Lejetiniae.  The  inflated  cavities  of  the  ventral  lobes  and  sometimes 
also  those  of  the  underleaves  were  mostly  occupied  by  one  or 
more  yellowish  or  colorless  subspherical  bodies,  40-120  //.  in 
diameter,  probably  sporangia  of  a  fungus  belonging  to  the  order 
CJiytridiales,  perhaps  to  the  genus  RJiizophidiiini  Schenk.  At 
maturity,  the  contents  of  some  of  the  globose  bodies  become  re- 
solved into  numerous  minute  ovoid  colorless  masses,  4—6  //  in 
diameter.  In  both  of  the  cases  where  the  Porclla  was  thus  affected 
it  was  growing  in  company  with  species  of  Radula  {Radida  coni- 
plaiiata,  near  Mendocino,  no.  707,  and  Radida  Bolandcri,  near 
McBride's,  Mad  River,  Humboldt  Co..  no.  105  i). 

Plates  115  and  116.    Porella  navicularis. 
I  and  2.    Plants,  $  and  9  »  natural  size. 
3.  Portion  of  9  plant,  viewed  ventral ly,  X 
4-7.  Leaves,  showing  ventral  lobes,  X  ^2. 
8  and  9.  Underleaves,  X  23. 

10.  Outline  of  cauline  leaves,  dorsal  view,  X  ^2. 

11.  9  branch,  with  young  perianth,  bracts,  bracteole,  etc.,  ventral  view,  X  21. 

12.  Mature  perianth,  ventral  aspect,  and  sporogonium,  X  12. 

13.  Inferior  basal  margin  of  dorsal  lobe,  X  25. 
14  and  15.  Leaves,  dorsal  aspect,  X  12. 

16.  One  half  of  mouth  of  young  perianth,  X  53- 

17.  Leaf-cells,  X  305 • 

Figs.  I,  7,  and  15  drawn  from  no.  776  (Ukiah);  2,  3,  6,  8,  ii,  12,  14,  and  16, 
from  no.  564  (Mendocino);  4,  5,  9,  and  lo,  from  no.  1215  (iNIill  Valley). 

35.  FRULLANIA  Raddi,  Atti  Soc.  Ital.  Sci.  Mod.  i8:— (9). 

1818.    Mem.  Mat.  e  Fis.  Soc.  Ital.  Sci.  Mod.  18:  20.  1S20. 

Spruce,  Trans,  and  Proc.  Bot.  Soc.  Edinb.  15  :  3.  1884. 

Plants  large  or  small,  depressed-caespitose,  mostly  reddish- 
brown,  less  commonly  green  or  nearly  black.  Stems  moderately 
stout,  opaque,  composed  of  several  layers  of  small  cells,  pinnately 
ramose,  the  branches  all  lateral,  each  arising  exactly  in  the  axil 
of  a  cauline  leaf,  from  which  it  is  always  free  ;  root-hairs  springing 
in  tufts  from  the  bases  of  the  underleaves.  Leaves  alternate,  some- 
what obliquely  or  almost  transversely  inserted,  complicate-bilobed  ; 
the  dorsal  lobes  incubous,  obliquely  ovate  to  suborbicular,  nearly 
always  entire  ;  ventral  lobes  (lobules)  usually  developed  as  inflated 
galeate,  cucullate,  cylindrical-clavate,  or  rarely  digitiform  Avater- 
sacs,  or  sometimes  evolute,  more  or  less  distant  from  the  stem, 


170 


JUXGERM  ANNI ACEA  E 


commonly  bearing  near  the  base  on  the  side  toward  the  stem  a 
small  or  minute,  subulate  or  triangular  process  (stylus) ;  lobule  of 
the  leaf  subtending  a  branch  mostly  explanate  and  almost  equally 
bifid  ;  leaf-cells  mostly  small  to  medium-sized,  with  more  or  less 
conspicuous  trigones  at  the  angles  and  sometimes  also  with  inter- 
mediate lateral  thickenings.  Underleaves  always  present,  consider- 
ably smaller  than  the  leaves,  bifid  (very  rarely  entire).  Dioicous  or 
autoicous  (rarely  paroicous).  Androecia  usually  occupying  short 
lateral  branches,  globose  to  oblong ;  ^  bracts  saccate,  closely  im- 
bricate, about  equally  2-lobed  ;  antheridia  2-4.  Archegonia  usu- 
ally 2-4,  very  rarely  as  many  as  12,  acrogenous  or  cladogenous, 
subfloral  "  innovations  always  wanting.  9  bracts  2-5  pairs,  larger 
than  the  leaves,  free  from  the  perianth,  the  two  of  the  inmost 
pair  (in  extra-limital  species)  occasionally  connate  with  the  bracte- 
ole  and  with  each  other  to  form  a  gamophyllous  exciple  ;  lobules 
evolute,  subentire  or  more  or  less  dentate,  ciliate,  or  laciniate  ; 
bracteole  often  connate  with  one  or  both  of  the  inmost  bracts. 
Perianth  free,  commonly  emersed,  more  or  less  compressed  dorso- 
ventrally,  trigonous  in  cross-section  with  the  third  angle  ventral  or 
sometimes  tetragonous  with  two  ventral  angles,  often  bearing  in 
addition  several  shorter  supplementary  dorsal  or  ventral  folds  or 
ridges,  otherwise  smooth  or  in  a  few  species  roughened  by  tuber- 
cles, contracted  at  the  'apex  to  a  tubular  rostellum,  irregularly 
ruptured  by  the  exsertion  of  the  capsule.  Calptra  free,  included, 
pyriform  or  obovoid,  fleshy.  Capsule  globose,  exserted  on  a  short 
seta,  dehiscing  by  four  valves  for  about  ^  the  distance  from  the 
apex  to  the  base,  the  valves  composed  of  two  layers  of  cells,  those 
of  the  outer  layer  rather  large  and  thick,  with  strong  columnar 
thickenings  (appearing  nodular  in  surface  view)  at  the  angles  and 
in  the  lateral  walls,  those  of  the  inner  layer  smaller  and  thinner, 
more  or  less  papilliform,  their  boundaries  obscured  by  irregular 
thickenings  ;  the  basal  uncleft  portion  of  the  capsule-wall  made  up 
of  several  layers  of  thin-walled  colorless  cells.  Seta  scarcely 
exceeding  the  perianth  or  2-3  times  longer,  stout,  8  or  9  cells  in 
diameter,  with  cells  alternate  (/.  c,  seta  not  appearing  articulate  in 
surface  view),  abruptly  dilated  above  to  form  the  base  of  the  cap- 
sule. Elaters  rather  few,  stout,  unispiral,  persistently  affixed  to 
the  upper  half  of  the  capsule-valves,  the  free  ends  truncate 
or  with  a  trumpet-like  expansion.  Spores  large,  papillate  or  ver- 
ruculose.* 


*  The  North  American  species  of  Frullania  have  been  admirably  figm-ed  and  de- 
scribed by  Dr.  Alexander  \V.  Evans  (Trans.  Conn.  Acad.  10  :  1-39.  //.  1-15- 
May,  1897).  The  following  descriptions  of  the  Californian  species  are  taken  with  bu*^ 
slight  modifications  from  Dr.  Evans'  paper. 


Frullania 


171 


Key  to  ttie  Species. 

Lobules  galeate  (when  inflated),  about  as  broad  as  long;  perianth  terminal  on  the 
main  stem  or  a  leading  branch. 
Dioicous,  flagelliferous  ;  lobules  (excepting  those  of  bracts  and  of  leaves  subtend- 
ing branches)  always  inflated,  slightly  longer  than  broad.     I.  F.  Bolanderi. 
Autoicous,  flagella  wanting  ;  lobules,  when  inflated,  broader  than  long,  very  often 
evolute.  2.  F.  Catalinae. 

Lobules  oval- cylindrical  to  short-clavate,  i]^-2y2.  times  as  long  as  broad;  dioicous; 
perianth  terminal  on  a  rather  short  lateral  branch.^ 
Dorsal  lobes  of  the  leaves  rounded-obtuse  to  acuminate-apiculate,  usually  (but 
not  always)  marked  with  a  short  median  line  of  slightly  enlarged  discolored 
cells  ;  lobules  separated  from  the  stem  by  about  their  own  width  ;  underleaves 
mostly  rhombic-obovate,  gradually  narrowed  to  the  base  and  without  a  trace  of 
an  auricle,  the  margins  plane,  often  bluntly  unidentate  on  either  side. 

4.  F.  Franciscana . 

Dorsal  lobes  of  the  leaves  rounded-obtuse,  discolored  cells  scattered  or  wanting ; 
lobules  separated  from  the  stem  by  less  than  half  their  width  ;  underleaves  sub- 
quadrate-orbicular,  more  or  less  auriculate  at  base,  the  margins  plane  or  slightly 
recurved  at  the  sides.  3.  F.  Californica. 

Dorsal  lobes  of  the  leaves  acute  or  more  commonly  acuminate-apiculate,  discolored 
cells  scattered  or  wanting,  rarely  in  a  short  median  line  ;  lobules  separated  from 
the  stem  by  about  their  own  width ;  underleaves  suborbicular  or  reniform, 
auriculate  at  base,  the  margins  strongly  recurved  at  least  toward  the  apices. 

5.  F.  Alsq^iallensis. 

1.  Subgenus  Trachvcolea  Spruce. 
I.  Frullania  Bolanderi  Aust.  Proc.  Acad.  Nat.  Sci.  Philad. 
1869:  226.    1869.    Evans,  Trans.   Conn.  Acad.  10  :  8.  //. 
2.  1897. 

FniUania  Pctalinncnsis  Gottsche ;  Bolander,  Cal.  Med.  Gaz. 
1870:  184  (zi-o)  (name  only).  1870. 

Frullania  Hallii  Aust.  Bull.  Torn  Bot.  Club,  6:  20.  1875. 

Dark  green,  varying  to  reddish-brown,  closely  appressed,  dif- 
fusely repent  or  caespitose  :  stems  6-20  mm.  long,  .07-.  14  mm. 
in  diameter,  irregularly  pinnate,  the  branches  often  prolonged  into 
erect  or  ascending  flagella,  these  crowned  at  the  apex  with  a  few 
smallwell-developed  leaves  but  furnished  otherwise  only  with  patent- 
horizontal  or  subsquarrose  underleaves  and  the  merest  rudiments 
of  lateral  leaves  :  dorsal  lobes  of  leaves  mostly  subimbricate,  as- 
cending when  moist,  obliquely  ovate  or  semiorbicular,  .35-5  mm. 
X  .27-.45  mm.,  arching  over  the  stem  and  subtruncate  at  base, 
entire,  the  slightly  decurved  apex  rounded-obtuse  ;  lobules  large, 
galeate,  .23-32  mm.  x  .2-27  mm.,  truncate  at  base,  close  to  the 

*  While  the  three  Californian  species  of  this  section  are  closely  allied  and  have  long 
been  confused,  they  are  nevertheless  entirely  distinct.  The  credit  of  finally  disen- 
tangling and  defining  them  belongs  largely  to  Dr.  Evans. 


172 


JUNGERMANNIACEAE 


stem  ;  stylus  small,  lanceolate  or  subulate  :  underleaves  distant, 
rhombic-obovate  or  somewhat  orbicular,  slightly  exceeding  the 
stem  in  width,  bifid  about  one  third  their  length  with  mostly  sub- 
acute lobes  and  sinus,  margins  plane,  entire  or  bearing  one  or  two 
teeth  at  the  sides :  superior  and  median  cells  of  the  dorsal  lobes 
i8-30/><,  all  rather  thick-walled,  trigones  small  or  conspicuous,  in- 
termediate thickenings  very  rarely  developed  :  dioicous  :  androecia 
ovoid  to  oblong,  ^  bracts  in  4-15  pairs  :  archegonia  3  or  4,  ter- 
minal on  the  main  stem  or  a  leading  branch  :  9  bracts  in  two  or 
three  pairs,  gradually  two  or  three  times  larger  than  the  leaves, 
unequally  bifid,  the  dorsal  lobes  ovate,  rounded-obtuse,  entire  or 
slightly  repand  ;  lobules  ovate  or  lanceolate,  obtuse  or  subacute, 
plane  or  lightly  concave  ventrally,  entire  except  for  a  small  tooth 
(stylus)  near  the  middle  of  the  inner  margin  :  bracteole  connate 
on  one  or  both  sides,  ovate  or  oblong-ovate,  commonly  bifid 
its  length  with  acute  lobes  and  sinus  but  sometimes  rounded 
at  apex  or  merely  emarginate,  rarely  3-  or  4-dentate  :  perianth 
about  half-emersed,  obovoid,  1.2- 1. 6  mm.  x  .8-1.3  mm.,  with  a 
rather  broad,  usually  two-angled  ventral  keel  and  one  or  more 
supplementary  ventral  and  dorsal  ridges,  very  rarely  with  a  few 
scattered  tubercles,  the  beak  short  and  broad  and  usually  flaring 
at  the  mouth  :  capsule  exserted  by  about  its  own  diameter  ;  spores 
variable  in  form  and  size,  mostly  40-60//  in  greatest  diameter. 
Exsicc.  Hep.  Bor.-Am.  105b. 
Hep.  Am.  28. 

On  the  bark  of  living  trees  {Qucrcus,  Alnus,  Uinbcllularia, 
Acsciihis,  etc.).  Very  common,  especially  in  the  Coast  Range 
counties.  Berkeley,  Fruit  Vale,  and  San  Leandro  (Prof.  L.  M. 
Underwood,  June,  1888),  Alameda  Co.;  Mission  Dolores  (Bolan- 
der),  San  Francisco  ;  Tomales  (Bolander)  and  Mill  Valley,  Marin 
Co.;  Kenwood  (Mr.  F.  T.  Bioletti),  Sonoma  Co.;  Mendocino 
(Bolander) ;  Blue  Lake  (1004),  Humboldt  Co.;  Mt.  Diablo,  Contra 
Costa  Co.;  Monterey  (Prof.  W.  G.  Farlow,  May,  1885)  and  Pa- 
cific Grove,  Monterey  Co.;  San  Gabriel  Mountains,  near  Pasadena 
(Prof.  A.  J.  McClatchie,  June  i,  1893).  The  species  extends 
northward  to  British  Columbia. 

2.  Frullania  Catalinae  Evans,  Trans.  Conn.  Acad.   10:  11. 

//.  4.  1897. 

Reddish-brown,  sometimes  tinged  with  green,  depressed-caespi- 
tose  :  stems  5-15  mm.  long,  .1-.  18  mm.  in  diameter,  irregularly 
pinnate  :  dorsal  lobes  of  leaves  imbricate,  squarrose  or  semivertical 


Frullania 


173 


when  moist,  obliquely  ovate,  .45-./  mm.  x  .4-. 62  mm.,  arching 
over  the  stem  and  subtruncate  at  base,  entire,  the  slightly  decurved 
apex  rounded-obtuse;  lobules  inflated  and  broadly  galeate,  .16— 
.18  mm.  long,  .2-27  mm.  wide,  separated  from  the  stem  by  about 
one  fourth  their  width,  or  very  often  imperfectly  inflated  or  wholly 
explanate,  then  ovate-linguiform,  .18-.3  mm.  long,  .1-.  18  mm. 
wide;  stylus  subulate,  minute  :  underleaves  distant  or  contiguous, 
plane,  broadly  rhombic  or  rhombic-obovate,  mostly  1-2  times  the 
width  of  the  stem,  bifid  ■^j-)^  their  length,  with  obtuse  or  sub- 
acute lobes  and  sinus,  entire  or  more  commonly  unidentate  on  the 
sides:  superior  and  median  cells  of  the  dorsal  lobes  16— 30 all 
with  slightly  thickened  walls,  small  trigones,  and  no  intermediate 
thickenings  :  autoicous  :  androecium  small,  ovoid,  occupying  a 
short  branch  springing  from  near  the  involucre,  S  bracts  1—3 
pairs  :  archegonia  about  6,  terminal  on  the  stem  or  the  principal 
branches  ;  9  bracts  in  two  or  three  pairs,  gradually  larger  than 
the  leaves,  unequally  bifid,  the  dorsal  lobes  ovate,  entire  or  sub- 
crenulate  toward  the  base,  obtuse  ;  lobules  shorter  and  narrower, 
ovate,  obtuse  or  subacute,  bearing  a  single  distinct  tooth  or  seg- 
ment slightly  above  the  middle  of  the  inner  edge,  otherwise  en- 
tire :  bracteole  oblong  or  narrowly  ovate,  connate  on  one  side  with 
bract,  bifid  for  ^  or  more  of  its  length,  with  lanceolate  parallel 
or  somewhat  divergent  lobes,  otherwise  entire  or  sometimes  uni- 
dentate toward  the  base  of  the  free  margin  :  perianth  about  half- 
exserted,  broadly  fusiform  to  obovoid-pyriform,  1.4-1.8  mm.  x 
.75-1.1  mm.,  with  a  distinct  two-angled  ventral  keel  and  one  or 
more  less  pronounced  dorsal  and  ventral  ridges,  gradually  nar- 
rowed to  a  short  broad  beak  :  spores  30-40  //  in  greatest  diam- 
eter, thickly  and  minutely  granulate-papillate. 

On  rocks  in  a  canon,  Santa  Catalina  Island  (Professor  A.  J. 
McClatchie,  Sept.  8,  1893). 

II.  Subgenus  Thyopsiella  Spruce. 
3.  Frullania  Californica  (Aust.)  Evans,  Trans.  Conn.  Acad.  10  : 
25.       12.  f.  11-22.  1897. 
Frullania  Grayana  Californica  Aust.  /.  /.  ;  Underw.  Bull.  111. 
State  Lab.  Nat.  Hist.  2  :  67.  1884. 

Frullania  Asagrayana  Californica  Aust.  {cJi.  eme?id.),  M.  A. 
Howe,  Erythea,  2:  98.  1894. 

Frullania  Asagrayana  alsophila  M.  A.  Howe,  Er}^thea,  2  :  99. 
1894. 

"  Frullania  Tamarisci  Nees,  ?"  ;  Bolander, /. Cal.  Med.  Gaz. 
1870:  184  (40).     1870.     y^ot  Frullania  Tamarisci  (1^.)  Dumort. 


174 


JUNGERMANNIACEAE 


Green,  reddish-brown,  or  fuscous,  densely  depressed-caespi- 
tose  or  rather  laxly  repent  :  stems  light  brown  or  greenish,  1-3 
cm.  long,  .1-.22  mm.  in  diameter,  once  or  twice  pinnate:  dorsal 
lobes  of  leaves  more  or  less  imbricate,  suborbicular,  those  of  the 
main  stem  .45-8  mm.  x  .35-.8  mm.,  all  arching  over  the  stem 
and  cordate  at  base,  entire,  rounded-obtuse  and  somewhat  de- 
curved  at  apex,  discolored  cells  scattered  or  wanting  ;  lobules 
oval-cyHndrical  to  short-clavate,  .13-22  mm.  x  .09-13  mm., 
separated  from  the  stem  by  less  than  half  their  width  ;  stylus  mi- 
nute, subulate  or  disc-like  :  underleaves  distant  or  approximate, 
subquadrate-orbicular,  1—3  times  the  width  of  the  stem,  more  or 
less  auriculate  at  base,  bifid  about  ^'  their  length  with  mostly  sub- 
acute lobes  and  narrow  or  broad  sinus,  margins  entire,  plane 
or  slightly  reflexed  at  the  sides  :  superior  and  median  cells  of  the 
dorsal  lobes  mostly  10-20//,  all  rather  thick-walled,  trigones  in- 
conspicuous and  intermediate  thickenings  scanty  :  dioicous  :  androe- 
cia  subglobose,  bracts  2-5  pairs  :  archegonia  terminal  on  a  short 
branch  :  9  bracts  in  two  or  three  pairs,  unequally  bifid,  the  dor- 
sal lobes  ovate  or  ovate-lanceolate,  acute,  entire  ;  lobules  lanceo- 
late or  subulate,  acuminate,  often  canaliculate  or  tubulose  by  the 
recurving  of  the  margins,  sometimes  falcate,  bearing  on  the  inner 
margin  a  laciniate  lobe-like  segment,  otherwise  entire  :  bracteole 
connate  with  the  bract  on  one  side,  ovate,  bifid  to  about  the  mid- 
dle with  subulate-acuminate  or  lanceolate-acuminate  lobes  and 
bearing  on  either  side  at  the  base  a  distinct  usually  laciniate  segment, 
otherwise  entire  :  perianth  ovoid,  1.6-2  mm.  x  .9-1.2  mm.,  some- 
what gradually  or  rather  abruptly  narrowed  to  the  short  beak, 
strongly  unicarinate  ventrally,  smooth  :  mature  sporogonium  un- 
known. 

Exsicc.    Hep.  Bor,-Am.  108,/./.  (distributed  as  F.  Nisqiial- 
i ens  is). 
Hep.  Am.  197. 

On  rocks,  logs,  and  trunks  of  living  trees.  Mission  Dolores 
(Bolander)  and  Clarendon  Heights  (Howe),  San  Francisco  ;  Mill 
Valley  and  Olema,  Marin  Co.;  Duncan's  Mills  (1209)  and  "Tur- 
ner's Cafion,"  Sonoma  Co.;  Wildcat  Cafion,"  near  Berkeley  ; 
near  Lake  Pilarcitos,  San  Mateo  Co.  (Blasdale)  ;  Pacific  Grove^ 
Monterey  Co.  There  are  in  existence  several  additional  speci- 
mens of  Dr.  Bolander's  collecting,  attributed  simply  to  "  Cali- 
fornia." The  species  extends  as  far  north  as  Victoria,  B.  C,  where 
it  has  been  collected  by  Professor  Macoun. 


Frullaxia 


175 


4.  Frullaxia  Fraxciscaxa  M.  A.  Howe,  Erythea,  2  :  99.  2. 
1894.    Evans,  Trans.  Conn.  Acad.  10:  26.   pL  ij. 
f.  i-S.  1897. 

Fndlania  Gray  ana  Califoriiica  Aust.  p.  p.\  Underw.  Bull.  111. 
State  Lab.  Nat  Hist.  2:  67.  1884. 

Fndlania  nnciflora  Calif ornica  Gottsche  ;  Bolander,  Cal.  Med. 
Gaz.  1870:  184  (40)  (name  only).  1870. 

Fndlania  Tainarisci  Nees  ?  "  ;  Bolander,  p.  p.  Cal.  Med.  Gaz. 
1890:  184(40).     1870.     Not  Fridlania  Taniarisci  {\^}jT)\imox\., 

Reddish-brown,  varying  to  greenish,  forming  wide  depressed 
mats:  stems  1.5-4  cm.  long,  .1-2  mm.  in  diameter,  light  brown 
or  greenish,  mostly  bipinnate  ;  dorsal  lobes  of  leaves  imbricate, 
obliquely  ovate,  those  of  the  main  stem  and  principal  branches 
.6-1  mm.  X  ■45--8  mm.  all  arching  over  the  stem  and  cordate  at 
base,  usually  marked  with  a  short  median  line  of  discolored  slightly 
enlarged  cells,  entire,  the  decurrent  apex  from  rounded-obtuse  to  acu- 
minate-apiculate  ;  lobules  oval-cylindrical  to  short-clavate,  .18-23 
mm.  X  .  I-.  14  mm.,  separated  from  the  stem  by  about  their  own  width; 
stylus  minute:  underleaves  distant,  subquadrate-rhombic  to  obovate, 
mostly  rhombic-obovate,  usually  2-2  times  the  width  of  the 
the  stem,  gradually  narrowed  to  the  neither  auriculate  nor  appen- 
diculate  base,  bifid  about  3^  their  length  with  narrow  sinus  and 
obtuse  lobes,  the  margins  plane,  usually  bluntly  unidentate  near 
the  middle  :  superior  and  median  cells  of  the  dorsal  lobes  mostly 
16-25  /J?,  all  rather  thick-walled,  trigones  and  intermediate  thick- 
enings becoming  more  conspicuous  toward  the  middle  and  the 
base  :  dioicous  :  androecia  subglobose  or  oval,  ,  $  bracts  3-7 
pairs  ;  antheridia  in  3's  :  archegonia  terminal  on  a  short  branch  : 
9  bracts  in  commonly  three  pairs,  unequally  bifid,  the  dorsal  lobes 
of  the  inmost  pair  ovate,  acute  or  acuminate-apiculate  or  rarely 
obtuse,  cili  ite  at  the  dorsal  base,  otherwise,  entire  ;  lobules  ovate 
or  lanceolate,  acute,  slightly  concave  ventrally,  ciliate  at  the  \^en- 
tral  basal  margin,  otherwise  entire  :  bracteoles  connate  with  the 
bract  on  one  side,  ovate,  bifid  to  about  the  middle  with  a  narrow 
sinus  and  lanceolate  acuminate  lobes,  ciliate  at  base,  otherwise  en- 
tire :  perianth  oblong-obovate,  1.5-2.3  mm.  x  .9-1.1  mm.,  ab- 
ruptly short-rostrate,  strongly  unicarinate  ventrally,  smooth  :  cap- 
sule exserted  by  about  twice  its  own  diameter  of  .7-. 8  mm.,  spores 
60-80  a  in  greatest  diameter,  the  verruculae  mostly  aggregated  in 
small  circular  patches. 

Exsicc.  Hep.  Bor.-Am.  108,/./.  (distributed  as Nisqnal- 
Icnsis). 


176 


JUXGERMANNIACEAE 


On  tree-trunks  and  old  fences,  rarely  on  rocks.  Common  in 
the  Coast  Range  Mountains  from  San  Francisco  northward. 
Mission  Dolores  (Bolander),  on  metamorphic  sandstone,  San 
Francisco"  (Bolander,  Apr.  4,  1864),  and  Clarendon  Heights 
(Howe),  San  Francisco  ;  Mill  Valley  and  Olema,  Marin  Co.; 
Mendocino  (Bolander  ;  Howe,  547,  548,  574,  579,  1210,  1211); 
Eureka  (Bolander ;  Howe,  970).  Collected  by  Dr.  A.  Kellogg 
also  and  by  Dr.  Bolander,  without  definite  locality. 

5.  Frullania  Nisquallensis  Sulliv.  Mem.  Amer.  Acad.  H.  4  : 
175.  1849.  Evans,  Trans.  Conn.  Acad.  10  :  21.  pi.  10. 
1897. 

Reddish-brown,  varying  to  yellowish-green,  depressed-caespi- 
tose :  stems  1.5-3  ^m.  long,  .12-27  ^^"^  diameter,  usually 

dark-brown,  mostly  bipinnate  :  dorsal  lobes  of  the  leaves  imbri- 
cate, obliquely  ovate,  those  of  the  main  stem  and  principal  branches 
.63-1.3  mm.  X  -5 5-9  mm.,  all  arching  over  the  stem  and  cor- 
date at  base,  entire,  the  strongly  decurved  apex  acute  or  more 
commonly  acuminate-apiculate,  discolored  cells  scattered  or 
wanting  or  rarely  in  a  short  median  line  ;  lobules  oval-cylindrical 
to  short-clavate,  .16-.24  mm.  x  .09— .14  mm.,  separated  from  the 
stem  by  about  their  own  width  ;  stylus  usually  minute  and  sub- 
ulate :  underleaves  distant  or  contiguous,  suborbicular  or  reniform, 
I— 3  ^  times  the  width  of  the  stem,  auriculate  at  base,  bifid  about 

their  length  with  obtuse  or  sometimes  apiculate  lobes,  the  sinus 
rather  broad  and  obtuse  or  subacute,  margins  entire,  strongly  re- 
curv^ed  at  least  toward  the  apices  of  the  mostly  reflexed  and  often 
subcucullate  lobes  :  superior  and  median  cells  of  the  dorsal  lobes 
mostly  16-30//,  all  rather  thick-walled,  trigones  and  often  inter- 
mediate thickenings  becoming  conspicuous  toward  the  middle  : 
dioicous  :  androecium  ovoid,  $  bracts  4-9  pairs  :  archegonia  ter- 
minal on  a  short  branch  :  9  bracts  in  about  three  pairs,  deeply 
and  unequally  bifid,  the  dorsal  lobes  ovate  or  ovate-lanceolate, 
acuminate,  entire,  sinuous,  or  very  sparingly  dentate  ;  lobules 
mostly  subulate-acuminate,  canaliculate  or  subtubulose  by  the 
recurving  of  the  margins,  often  falcate  or  distorted,  bearing  on  the 
inner  side  toward  the  base  a  small  laciniate  segment  or  a 
cluster  of  cilia  :  inmost  bracteole  connate  with  bracts  on  both 
sides  or  sometimes  on  one  side  only,  ovate,  bifid  Yi-Yz  its 
length  with  lobes  similar  to  the  lobules  of  the  bracts,  the  margins 
ciliate  or  dentate  toward  the  base  :  perianth  long-ovoid,  2-2.5 
mm.  X  .9-1.2  mm.,  rather  gradually  narrowed  to  the  short  beak, 
strongly  unicarinate  ventrally,  smooth  :  capsule  exserted  by  about 


Frullania 


177 


its  own  diameter  of  .7-8  mm.;  spores  60-100/^?  in  maximum 
diameter,  verruculose. 

On  rocks  and  trunks  of  trees. 

Clarendon  Heights,  San  Francisco  (Jan.  23,  1895);  Duncan's 
Mills  (1209,  in  part),  Sonoma  Co.;  Mendocino  (1223)  ;  Kneeland 
Prairie  Road  (1042),  Humboldt  Co.,  near  Crescent  City,  Del 
Norte  Co.  (Thomas  Howell,  April,  1892).  The  species  extends 
as  far  north  as  Vancouver  Island  (Macoun). 


178 


Anthocerotaceae 


Class  ANTHOCEROTES. 

The  class  AntJwcerotes  (see  page  9)  comprises  the  single 
order  Anthocerotales  and  the  single  family  Anthocerotaceae. 

ANTHOCEROTACEAE. 

GametophOre  a  wholly  leafless  thallus,  commonly  flat,  orbic- 
ular, or  semiorbicular,  more  or  less  lobed  or  radiately  dissected, 
sometimes  subpinnate  (Dendroceros),  or  ribbon-shaped  and  sub- 
dichotomously  branched,  with,  or  more  frequently  without,  a  well- 
defined  costa,  usually  of  several  layers  of  cells,  often  with  interior 
mucilage  cavities,  more  rarely  with  intercellular  air-spaces,  without 
proper  epidermis,  but  provided  on  the  ventral  surface,  sometimes 
also  on  the  dorsal,  with  inconspicuous  clefts  ("  stomata,"  "muci- 
lage-slits"), the  thallus  becoming  through  these  infected  with 
Nostoc  colonies  ;  cells  with  a  single  large  chloroplast  which  com- 
monly encloses  the  nucleus. 

Sexual  organs  embedded  in  the  thallus  ;  antheridia  arising 
endogenously,  short-stalked,  single  or  in  groups  of  2—4  (rarely 
more),  occupying  cavities  separated  from  the  dorsal  surface  by  two 
layers  of  cells,  the  covering  ruptured  at  maturity ;  walls  of  arche- 
gonium  confluent  with  surrounding  cells  of  the  thallus,  the  neck- 
canal  communicating  with  dorsal  surface  at  maturity  of  the  egg- 
cell.  Special  calyptra  not  differentiated.  "  Involucre  "  tubular, 
of  several  layers  of  cells,  formed  from  archegonium-wall  and  ad- 
jacent cells  of  thallus,  usually  soon  broken  through  at  the  apex 
by  the  elongating  capsule  and  remaining  as  a  sheath  about  its 
base,  or  (in  Nototliylas)  irregularly  torn  by  external  agencies. 

Sporogonium  consisting  of  a  pod-like,  usually  erect  and  much 
elongated  capsule,  a  bulbous  foot,  and  a  short  intervening  zone 
occupied  by  a  long-active  meristematic  tissue  ;  capsule  dehiscing, 
with  rare  exceptions,  from  the  apex  downward  by  two  vah^es,  its 
walls  containing  chlorophyll  and  in  most  species  of  Anthoccros 
bearing  stomata,  each  with  two  crescentic  guard-cells  ;  a  slender 
thread-like  columella,  surrounded  throughout  and  covered  at  the 
apex  by  the  spore -forming  layer,  probably  always  present,  but 
sometimes  early  disintegrated  and  obscure..  Spores  more  or  less 
plainly  tetrahedral,  long  adhering  in  fours,  smooth,  verrucose, 
papillate,  or  echinulatc,  ripening  successively  from  apex  of  cap- 
sule downward  ;  sterile  cells  from  the  archesporium  single  and 
subcubical  at  maturity,  or,  more  often,  elongated  and  forming 
hcteromorphic,  variously  contorted,  occasionally  branched  fila- 


Anthoceros 


179 


ments  of  2-4  cells,  sometimes  (^AntJioccros  sp.  and  Dcndroceros — 
in  tropics  and  southern  hemisphere)  with  one  or  more  well  devel- 
oped spiral  bands,  or,  more  commonly  (pseudo-elaters),  with  spiral 
thickenings  rudimentary  or  wanting. 

Three  genera,  Antlioceros,  Dendroceros,  and  Notothylas,  are  or- 
dinarily recognized,  of  which  only  the  first  is  known  to  occur 
within  our  limits. 

ANTHOCEROS  L.  Sp.  PL     1139.     1753.    Ex  Mich.  Nov.  PI. 
Gen.  \o,  pi.  7./.  /,  2.  1729. 

Thallus  suborbicular,  variously  lobed  or  irregularly  dissected, 
now  and  then  somewhat  ribbon-shaped  and  subdichotomous,  of 
more  than  one  layer  of  cells  throughout,  costa  wanting,  indistinct, 
or  rarely  prominent ;  monoicous,  proterandrous,  or  sometimes  di- 
oicous  (P)"^;  capsule  erect,  bivalved,  much  exceeding  the  involu- 
cre in  length,  usually  very  long,  with  stomata  (in  the  North  Amer- 
ican species).  Spores  granulose-papillate,  tuberculate,  or  echinu- 
late,  rarely  nearly  smooth  ;  sterile  cells  (in  all  our  species)  without 
spiral  thickenings. 

Key  to  the  Species. 

Thallus  with  peduncled  tubers  from  ventral  surface.      i.  A.  phymatodes. 
Thallus  without  ventral  peduncled  tubers. 

Spores  fuscous  or  black.  4.  A.  fiisifoi-mis. 

Spores  yellow. 

Spores  with  a  few  (8-15)  crescentic  verrucae  on  convex  (outer)  surface,  or 
nearly  smooth ;  thallus  usually  with  marginal,  sometimes  peduncled,  gland- 
ular-thickenings. 2.  A.  Pearsoni. 

Spores  thickly  granulose-papillate.  3.  A.  Caroliiiianns  occidentalis. 

I.  AxTHOCEROS  PHYMATODES  ]\L  A.  Howe,  Bull  Torn  Bot.  Club, 
25:  12.  pL         j2j.  1898. 

Thallus  dark  green,  blackening  on  drying,  dissected  ;  the  seg- 
ments dichotomous,  oblong  or  linear  in  sterile  plants,  broader  in  the 
fertile,  4-10  mm.  x.5— 2.5  mm.,  prostrate  or  very  slightly  ascending 
at  the  apices,  at  first  radiate,  becoming  later  intricately  entangled, 
narrowly  canaliculate  dorsally,  with  ascending  sinuate  or  repand 
margins,  obtuse,  carinate  with  a  broad,  thick,  very  distinct  or  some- 
times obsolescent,  naked  or  sparingly  radiculose  costa,  this  occupy- 
ing nearly  the  whole  of  the  narrower  segments  and  sending  down 
here  and  there  a  fleshy  or  elongated  and  slender  process  (.1-4 

*  Certain  species  of  Anthoceros  have  been  described  as  dioicous,  yet  in  a  genus  of 
which  the  best  known  representatives  are  proterandrous,  and  in  which  the  archegonia 
are  observed  with  so  much  difificulty,  it  may  be  suspected  that  careful  investigation,  by, 
modern  methods,  would  show  proterandry  to  obtain  in  all. 


180 


Anthocerotaceae 


mm.  in  thickness),  terminated  by  a  globose  or  ellipsoidal  tuber, 
the  latter  .25-1  mm.  in  diameter,  pale  when  living,  yellowish- 
brown  on  drying,  becoming  finally  closely  covered  with  root-hairs  ; 
thallus  10—16  cells  thick  in  region  of  costa,  passing  gradually  or 
rather  abruptly  into  the  more  or  less  extended  5-3-stratose  mar- 
ginal lamina;  surface-cells  rhombic  to  rhombic-oblong,  33—66/^ 
X  20—33  /A  indistinct  after  drying  :  involucres  separate,  cylindrical, 
1.7-2  mm.  X  .5  mm.,  smooth,  truncate,  entire  or  slightly  repand 
at  the  mouth. 

On  compact  soil  in  a  nearly  level  open  plot  about  fifty  feet 
north  of  ''The  Old  Mill,"  Mill  Valley,  Marin  Co.,  California 
(Howe:  Mar.  19,  1892,  and  Feb.  22,  1896;  the  latter,  which 
bears  the  involucres,  we  consider  the  type).  Also  collected  by 
Dr.  Bolander  at  "  Oakland,  on  slides  near  the  bay."  The  few 
capsules  that  have  been  seen  are  very  immature,  projecting  only 
about  I  mm.  beyond  the  mouth  of  the  involucre,  and  it  is  quite 
possible  that  the  description  of  the  involucre  given  above  will  need 
some  modification  on  the  discover},^  of  fully  ripened  sporogonia. 

It  is  practically  certain  that  this  species  is,  in  part,  at  least, 
what  Austin  referred  provisionally  to  Anthocei'os  cacspiticiiis  De 
Not.  (Bull.  Torr.  Bot.  Club,  6:  26.  1875),  drawing  his  description, 
however,  from  a  specimen  in  herb.  Torrey  under  the  name  A. 
laevis  Linn.,  fi-om  the  Island  of  Corsica."  We  had  thought  to 
take  up  for  A.  phymatodcs  the  specific  name  Torreyi  Aust.  MS., 
which  appears  in  parenthesis  after  A.  caespiticiiis  in  the  place  cited, 
but  an  examination  of  the  Austin  collection,  so  generously  loaned 
by  Mr.  Pearson,  showed  the  manuscript  diagnosis  of  A.  Torreyi 
affixed  to  a  scrap  of  the  Corsican  plant  (which  we  refer  to  AiitJio- 
ceros  dicJiotomus  Raddi)  and  that  plant  alone  alluded  to  at  the  close 
of  the  description.  A.  Torreyi  Aust.  MS.  is  therefore  a  synonym 
of  A.  dichotomiis  and  not  of  A.  pJiymatodes.  Bolander' s  specimen 
is  accompanied  by  a  slip  of  brown  paper  on  which  is  written — 
evidently  by  himself — Oakland  on  slides  near  the  bay,"  but  it  is 
found  in  the  Austin  collection  glued  to  Austin's  manuscript  de- 
scription of  A.  Lescioii  from  New  Orleans,  the  evident  type  of 
which  appears  in  the  same  collection  under  the  name  of  A.  Liido- 
vicianus. 

Anthoceros  phymatodes  is  a  near  ally  of  A.  dicJiotomns  Raddi, 
specimens  of  which  from  Raddi's  herbarium  we  have  had  the 


Anthoceros 


181 


privilege  of  examining  through  the  kindness  of  Professor  Massa- 
longo,  as  well  as  specimens  from  Portugal  and  Italy,  obligingly- 
communicated  by  Herr  Stephani.  But  A.  dichotomiis  has  a  costa 
}i-y^  the  width  of  the  segment,  6-10  cells  thick,  the  lamina 
mostly  2-stratose,  involucre  .7—1  mm.  wide,  often  enlarged  at  the 
irregularly  2-4-lobed  mouth  ;  while  A.  pJiymatodcs  has  a  costa 
i- 1-  the  width  of  the  segment,  10-16  cells  thick,  the  marginal 
lamina  5-3-stratose,  involucre  .5-6  mm.  wide,  not  expanded  at  the 
entire  or  repand  mouth.  To  facilitate  further  comparison,  when 
the  mature  sporogonia  are  found,  it  may  be  added  that  in  A.  dicho- 
to)niis  ixom  Raddi's  hei'hd.num  A ;it/ioceros  polyinorpJms^2idi<^\  j 
dichotonius  Raddi  "  is  the  legend  the  packet  bears)  the  capsules  are 
erect  or  slightly  curved,  7.5—15  mm.  long,  often  thickened  toward 
the  apex,  spores  yellow,  becoming  yellowish-brown,  rounded- 
tetrahedral,  almost  wholly  smooth  on  all  faces,  42—60  a  in  maxi- 
mum diameter,  pseudo-elaters  yellowish,  of  1-4  irregular  elongated 
cells,  geniculate,  variously  contorted,  often  branched. 

Anthoceros  cacspiticiiis  DeNot.,  the  probable  original  of  which 
(from  the  vicinity  of  Cagliari,  in  southern  Sardinia),  we  have  been 
allowed  to  study  through  the  indulgence  of  Professor  Pirotta,  is  not 
closely  related  to  A.  dicJiotoinns,  with  which  it  was  compared  by 
its  author.  It  is  a  near  ally  of  AntJioceros  pjinctatus  and  may  not 
be  specifically  distinct.  The  spores  incline  to  be  fuscous  and  are 
finely  setose-papillate  on  the  outer  face  and  foveolate-reticulate 
on  the  inner  faces  ;  they  are  40—45  fi  in  maximum  diameter.  The 
pseudo-elaters  are  practically  as  in  A.  punciatus.  The  involucres 
measure  .65—1.1  mm.  x  .25— .5  mm.  It  is  surely  very  different 
from  the  Californian  plant,  even  though  mature  spores  of  the  latter 
are  wanting.  The  thallus  apparently  bears  no  tubers  ;  it  is  thmner 
than  in  A.  pJiymatodcs,  is  quite  indistinctly  costate,  and  has  the  pe- 
culiar narrow  ascending  marginal  shoots  oi  A.  piinctatus.  It  is  de- 
scribed by  DeNotaris  as  being  erect,  while  A.  pJiymatodcs  is  pros- 
trate. 

AntJioceros  tiiberosus  Tayl.,  from  Swan  River,  Australia  (Drum- 
mond,  1873),  have  examined  through  the  courtesy  of  Dr.  B. 
L.  Robinson,  Curator  of  the  Gray  Herbarium,  in  which  the  Tay- 
lor collection  is  incorporated.  It  differs  from  our  plant  in  the 
much  thinner,  broader,  scarcely  costate  thallus,  the  terminal  or 


182 


AXTHOCEROTACEAE 


marginal  tubers,  and  the  shorter  somewhat  obovate  involucres. 
The  spores  are  yellow,  becoming  brownish,  sparingly  w^arty-gran- 
ulose,  35-45  tJ- ;  pseudo-elaters  mostly  of  i  or  2  elongated  genicu- 
late cells. 

AiitJioceros  pJiyinatodcs  differs  from  AntJioceros  Donnellii  Ausi. 
in  the  larger,  less  numerous  tubers  on  stouter  stalks,  also  in  the 
thicker,  more  opaque  thallus,  with  less  distinct  surface  cells,  in  the 
longer  segments,  etc. 

The^tubers  of  A.  pJiyinatodcs  are  formed  at  the  apex  of  the 
costa,  but  their  peduncles  become  strictly  ventral  in  their  attach- 
ment by  the  continued  onward  growth  of  the  segment.  Two  or 
three  tubers  successively  older  and  larger  as  one  passes  backward 
may  often  be  seen  depending  from  a  single  branch.  The  tuber 
consists  of  a  cortex  of  2—4  layers  of  nearly  empty  cells  enclosing 
a  central  mass  of  smaller  cells  so  densely  filled  with  oil  drops  or 
nearly  colorless  granules  that  the  cell  boundaries  in  a  section  are 
rendered  obscure. 

An  interesting  paper,  On  the  Structure  and  Contents  of  the 
Tubers  of  AntJioceros  tubcrosiis  Taylor"  was  published  in  the 
Memoirs  and  Proceedings  of  the  Manchester  [Eng.]  Literary  and 
Philosophical  Society,  vol.  41,  no.  2  (1896)  by  Mr.  J.  H.  Ash- 
worth.  Mr.  Ashworth  finds  oil-drops  and  proteid  granules  in  the 
interior  cells  of  the  tubers  of  this  species.  We  have  stated  (Bull. 
Torn  Bot.  Club,  25:  14.  1898)  that  the  granules  in  the  tubers 
of  AntJioceros  pJiyniatodcs,  in  material  preserved  in  alcohol,  show  a 
starch  reaction  with  iodine,  but  further  tests  upon  the  same  ma- 
terial either  fail  entirely  to  show  the  starch  reaction  or  show  it  so 
obscurely  that  we  must  doubt  the  accuracy  of  our  first  observation. 
The  granules  mostly  contain  one  or  two  more  highly  refringent 
portions,  as  described  by  Mr.  Ashworth  in  A.  tubcrosiis,  and,  on 
the  application  of  iodine,  they  usually  are  stained  a  yellowish- 
brown  or  remain  altogether  unaffected. 

Plates  117  and  118.    Anthoceros  phymatodes. 

1.  Portion  of  the  thallus,  natural  size. 

2.  Ventral  view  of  a  thallus  segment,  showing  costa  and  tuber,  X  23.  * 
3-6.  Lateral  views  of  thallus  segments,  showing  form  and  stages  of  development 

of  tubers,  X  23. 

7  and  8.  Old  tubers,  showing  vegetative  propagation  by  sprouting,  X  23. 
9.  Longitudinal  section  through  costa  and  young  tuber,  X  47- 


Anthoceros 


183 


10.  Longitudinal  section  showing  early  stages  in  the  formation  of  the  tuber, 
X  47-.  The  section  is  somewhat  oblique  and  does  not  exhibit  the  full  thickness  of  the 
costa. 

11.  Section  through  a  well-grown  tuber,  X  53- 

12.  A  single  interior  cell  from  fig.  ii,  X  S^S- 

13.  Longitudinal  section  through  the  costa,  X  53- 

14  and  15.  Transverse  sections  of  thallus  segments,  X  53* 

16.  Outlines  of  two  other  transverse  sections  of  thallus  segments,  X  23. 

17.  Involucres,  X  12. 

Figs.  1-6  and  9-17  from  specimen  collected  in  Mill  Valley,  Marin  Co.,  February 
22,  1896 — mostly  drawn  from  material  fixed  with  I  fo  chromic  acid  and  preserved  in 
alcohol ;  7  and  8,  from  same  locality,  March  19,  1892. 

18.  Involucre  of  Anthoceros  polynno7-phns  Raddi,  }-  dichototmis  Raddi,  ex  herb. 
Raddi,  X  12. 

2.  Anthoceros  Pearsoni  M.  A.  Howe,  Bull.  Torr.  Bot.  Club,  25  : 
^.  pi.  322,  323.  1898. 

Thallus  smooth,  prostrate  or  ascending,  rarely  erect,  mostly 
irregularly  dissected  ;  the  major  segments  linear  or  elongate-flabel- 
liform,  5-20  mm.  x  1-5  mm.,  often  intertangled,  sparingly  lobed 
or  two  or  three  times  subradiately  dichotomous,  plane  or  slightly 
concave,  rarely  costate  ventrally  at  apices,  in  sterile  plants  usually 
6-1 1  cells  thick,  gradually  or  abruptly  reduced  to  three  or  four  at 
the  margin,  in  the  fertile  commonly  expanded  into  a  wide  3-  or  2- 
stratose  lamina  with  a  lightly  crisped  periphery,  the  lobes  in  both 
often  terminating  in  dark  green,  globose  or  difform,  glandular- 
thickenings,  .2-2  mm.  in  diameter,  these  sometimes  long-peduncled, 
rarely  descending  and  radicelliferous  ;  surface-cells  rather  distinct, 
rhombic-trapezoidal  to  oblong,  20-70  fi  x  20-30  //,  the  inner  larger 
but  compact ;  Nostoc  colonies  usually  abundant,  ellipsoidal  or  fusi- 
form :  monoicous  :  involucres  solitary  or  approximate  in  pairs, 
subterminal  in  erect  forms,  somewhat  pellucid,  mostly  light  green, 
elongate-cylindrical,  2.3-3.3  mm.  X  . 5- 85  mm.,  sometimes  decum- 
bent at  base,  the  mouth  entire,  repand-dentate,  often  lacerate  with 
age,  now  and  then  scarious  :  capsule  pale  brown,  yellowish,  or  drab, 
slender,  sulcate,  8-32  (mostly  20-24)  mm.  x  .24-4  mm.,  rather 
thick-walled,  the  valves  finally  thin  and  often  twisted,  stomata  abun- 
dant ;  columella  filiform,  naked  ;  spores  yellow,  the  convex  face  with 
a  few  (8-15)  small,  crescentic,  rarely  obsolete  warts,  otherwise 
smooth  or  most  faintly  granulose,  35-50  p.  in  maximum  diameter ; 
pseudo-elaters  dilute  yellow  or  occasionally  tinged  with  brown,  of 
1-4  cells,  geniculate  and  variously  contorted,  often  branched,  the 
cells  30-80/7.  X  6-\o  fi,  with  wall  of  nearly  uniform  thickness. 

On  moist  banks  and  dripping  rocks.  Mill  Valley  (no.  16, 
May  7,  1892 — the  type — and  1205),  Olema — also  collected  in 


184 


Anthocerotaceae 


Marin  Co.,  by  Dr.  Bolander  and  by  T.  S.  Brandegee  ;  Duncan's 
Mills  (1198,  1200)  and  Cazadero  (1199)  Sonoma  Co.;  Mendocino 
(587,  702)  ;  Deer  Creek  Canon,  Russ  and  Graham's  Ranch 
(1070)  Humboldt  Co.;  near  Douglas  City  (1202),  Trinity  Co.; 
Oak  Run,  Shasta  Co.  (Baker  and  Nutting)  ;  Mitchell's  Canon, 
Mt.  Diablo  (1201);  Berkeley  (i  204) ;  Palo  Alto  (Prof.  D.  H. 
Campbell)  ;  Santa  Monica  (Dr.  H.  E.  Hasse)  ;  San  Bernardino 
(Mr.  S.  B.  Parish,  173  i,  2220). 

A7itJioceros  Pcarsoni\s>  the  commonest  yellow-spored  AntJwccros 
of  California,  and  perhaps  of  the  Pacific  Coast  as  a  whole.  We 
have  seen  as  many  as  twenty  specimens,  ranging  from  southern 
California  to  Washington,  and  they  are  always  very  clearly  dis- 
tinct from  Anthoceros  laevis  L.,  A.  Caroliniamis  Michx.,  and  A. 
Hallii  Aust.  ;  differing  from  the  last-mentioned  in  having  the  cap- 
sule usually  four  times  as  long,  in  the  much  more  perfect  pseudo- 
elaters,  composed  of  1-4  elongated  cells,  in  the  spores  being 
scarcely  otherwise  roughened  than  by  8-1  5  small  crescentic  ver- 
rucae,  etc.;  very  different  from  both  A.  laevis  and  A.  Caroliniamis^ 
in  the  character  of  the  spore-markings,  the  spores  of  the  two  latter 
species  being  always  thickly  granulose-papillate.  Sterile  condi- 
tions can  sometimes  be  only  doubtfully  distinguished  from  A. 
Hallii,  but  can  be  separated  from  glandular-thickened  forms  of  A. 
Caroliniaiius  occidcntalis  by  the  narrower  segments  and  often  pe- 
duncled  glands.  The  species  is  extremely  variable  in  thickness 
and  form  of  thallus,  though  very  constant  in  spore-markings. 
In  its  thicker,  slightly  costate  conditions,  particularly  when  the 
glandular-thickenings  are  long-stalked,  descending,  and  tuber- 
like, the  thallus  bears  some  resemblance  to  that  of  A.  phyinatodes, 
but  in  all  such  cases,  we  believe,  the  attachment  of  the  tubers 
never  becomes  strictly  ventral  ;  their  peduncles  can,  with  care, 
always  be  traced  back  to  the  margins  of  the  frond.  The  glands 
or  tubers  are  10-25  cells  in  thickness  and  consist  of  a  more  or  less 
distinctly  defined  capsule  of  1-3  layers  of  undifferentiated  cells  en- 
closing cells  containing  at  first  an  unusually  large  chloroplast  and 
later  filled  with  oil -drops  or  densely  crowded  with  whitish  gran- 
ules. These  organs  are  evidently  food  reservoirs,  adapting  the 
thallus  to  persistence  through  the  dry  summers  of  the  region  that 
the  plant  inhabits.     In  the  neighborhood  of  San  Francisco  the 


AXTHOCEROS 


185 


thickenings  begin  to  appear  in  January  or  February,  attaining  full 
development  in  May  or  June.  In  our  no.  702,  collected  near 
Mendocino,  California,  these  marginal  thickenings  often  enclose 
numerous  antheiidia. 

The  above  description  of  AntJioceros  Pearsoni  is  based  upon 
various  specimens,  but  our  no.  16,  collected  at  Mill  Valley,  Marin 
County,  May  7,  1892,  from  which  the  figures  of  the  spores,  pesudo- 
elaters  and  capsules  have  been  drawn,  we  consider  the  type  of  the 
species. 

Plates  119  and  120.    Anthoceros  Pearsoni. 

1.  Segment  of  the  thallus,  with  mature  sporogonia,  natural  size. 

2,  3  and  4.  Portions  of  a  sterile  thallus,  showing  its  form,  manner  of  branching 
and  the  beginnings  of  the  marginal  glandular-thickenings,  natural  size. 

5.  Marginal  glandular-thickenings,  fully  developed,  X  12. 

6  and  7.  Thallus  segments  with  younger,  sometimes  peduncled,  glandular  thick- 
enings, X  12. 

8.  Ventral  view,  showing  glandular- thickening  bearing  root-hairs,  X  12. 
9  and  10.  Longitudinal  sections  through  well-developed  glandular-thickenings, 
the  latter  exhibiting  embedded  antheridia,  X  53- 

11.  Transverse  section  of  margin  of  the  thallus,  X  53- 

12.  Median  longitudinal  section  of  the  thallus,  X  53- 
13-15.  Involucres,  X  ^2. 

16.  Pseudo-elaters,  X  225. 

17.  Spores,  X  305- 

Figs.  1,13,  16  and  17  from  no.  16,  Mill  Valley,  INIarin  Co.  (the  type)  ;  2-4, 
near  Dancan's  Mills,  Sonoma  Co.  (no.  1200);  5,  9,  10,  14  and  15,  near  Mendocino 
(no.  702)  ;  6  and  7,  Cazadero,  Sonoma  Co.  (no.  1 199)  ;  8,  Palo  Alto  (Prof.  D.  H. 
Campbell);  ii  and  12,  Mill  Valley,  Feb.  22,  1896. 

3.  Anthoceros  Carolinianus  occidentalis  M.  A.  Howe,  Bull. 
Torn  Bot.  Club,  25  :  8.  //.  321.  1898. 

Thallus  large,  thin,  ecostate,  usually  densely  caespitose,  ascend- 
ing, now  and  then  with  margins  and  lobes  glandular-thickened, 
light-  or  dark-green,  a  little  pellucid  or,  on  drying,  opaque  with  a 
somewhat  fatty  lustre,  dissected  ;  major  segments  oblong  or  obo- 
vate,  1-3  times  subdichotomously  branched,  8-30  mm.  x  2-7  mm., 
more  or  less  widened  at  the  rounded  crenate  apex,  4-8  cells  thick 
in  the  middle,  passing  gradually  into  a  wide  3-  or  2-stratose  mar- 
gin, or  nearly  uniform  throughout ;  surface-cells  distinct  in  terres- 
trial forms,  obscure  in  the  aquatic,  irregularly  rhombic,  becoming 
elongate-hexagonal,  20-1 20 x  18-30  ji:  monoicous  :  involucres 
sometimes  united  in  pairs,  thick-walled,  somewhat  pellucid,  long- 
cylindrical,  2-6.5  mm.  X. 5-1  mm.,  sometimes  contracted  or  now 


186 


AXTHOCEROTACEAE 


a  little  widened  toward  the  thin,  subentire  or  repand-dentate,  rarely 
scarious  mouth,  often  longitudinally  plicate  toward  the  base  :  capsule 
slender,  25-50 mm.  x.25-. 5  mm.,  pale  brown,  short-pedicellate,  the 
valves  twisted  ;  spores  yellow,  thickly  granulose-papillate,  especially 
on  the  convex  (outer)  face,  35-56  (jl  in  maximum  diameter  ;  pseudo- 
elaters  nearly  colorless  to  light  yellowish-brown,  of  1—4  cells, 
geniculate,  often  branched,  the  cells  extremely  variable  in  size  and 
form. 

On  wet  rocks  and  on  the  banks  of  rivulets  near  the  water-line. 

Olema  (Jepson),  Mt.  Tamalpais,  with  Fissidens  graiidifrons 
(Howe,  Setchell);  Howell  Mt.,  Napa  Co.  (Setchell);  North  Fork 
of  Little  River,  Mendocino  Co.;  near  Eureka,  Humboldt  Co.; 
Santa  Cruz  Mts.  (Underwood) ;  San  Bernardino  Mts.  (Parish); 
near  Pasadena  (McClatchie);  Twin  Oaks,  San  Diego  Co.  (Koch). 

A.  Caroliniamis  differs  from  A.  laevis  L.,  chiefly  in  greater  size 
of  the  thallus,  in  the  longer  involucres,  scarcely  expanded,  some- 
times contracted,  at  the  mouth,  and  in  the  (typically)  more  slender 
capsules.  The  species  as  a  whole  is  perhaps  to  be  looked  upon  as 
an  aquatic  or  semi-aquatic  analogue  of  A.  laevis,  from  which  it  is 
probably  not  always  distinct.  It,  however,  departs  so  widely  from 
the  ordinary  conception  of  Anthoceros  laevis  and  is  usually  so 
easily  recognizable  that  we  think  it  has  a  fair  claim  to  specific  rec- 
ognition. 

The  var.  occidentalis  differs  from  the  Anthoceros  Caroliiiianus 
Michx.  of  the  Southern  States  in  the  usually  more  densely  caespitose 
ascending  thallus,  sometimes  forming,  when  growing,  wide  cush- 
ions 3-4  cm.  in  depth,  while  that  of  the  typical  A.  Caroliiiianus 
is  commonly  prostrate ;  in  the  occasionally  glandular-thickened 
margins  and  lobes;' and  in  the  less  slender  capsule.  In  the  glandular- 
thickened  forms  of  this  variety,  involucres  which  seem  abnormally 
shortened  and  incrassate  (2-2.5  X  I  mm.)  are  often  found 

intermingled  with  the  longer.  Such  involucres  commonly  have 
scarious  mouths. 

Plate  121.    Anthoceros  Carolinianus  occidentalis. 
I  and  2.  Portions  of  the  thallus,  natural  size. 
3-5.  Involucres,  yi  12. 

6.  Segment  of  thallus  with  marginal  glandular- thickening,  X  6. 

7.  Transverse  section  of  thallus  segment  (showing  but  one  half  the  width),  X  23 

8.  Spores,  X  305- 

9.  Pseudo-elaters,  X  225. 


Anthoceros 


187 


Figs.  1-3,  8  and  9  from  a  specimen  collected  near  Olema,  Marin  Co.  (Mr.  W.  L. 
Jepson);  4  and  6,  Twin  Oaks,  San  Diego.Co.  (Mr.  F.  W.  Koch);  5,  Howell  Moun- 
tain, Napa  Co.  (Prof.  W.  A.  Setchell);  7,  Mt.  Tamalpais,  Marin  Co. 

4.  Anthoceros  fusiformis  Aust.  Bull.  Torn  Bot.  Club,  6 :  28. 

1875. 

Thallus  in  depressed  rosettes  6—20  mm.  in  diameter,  or  often 
suberect  and  caespitose  in  tufts  5-20  mm.  deep,  thick  and  rather 
rigid,  opaque  and  blackened  when  dry,  undulate-crisped,  deeply 
and  irregularly  dissected,  ecostate,  cavernose  within,  10-25  cells 
thick  in  median  parts,  becoming  rather  abruptly  3-  or  2-stratose  at 
margin,  often  with  large  and  abundant  glandular-thickenings,  more 
or  less  lamellate-cristate,  lacunose,  the  lamellae  sometimes  broad 
and  leaf-like  ;  surface-cells  somewhat  distinct,  subquadrate  to  ob- 
long-hexagonal, 30-75/7.  X  18-40  n,  with  large  chloroplast,  inner 
cells  much  broader  and  longer  in  axile  and  basal  parts,  but  often 
scarcely  larger  toward  the  growing  apices  :  monoicous  :  antheridia 
in  groups  of  2-4  :  involucres  often  numerous  and  crowded,  but 
very  rarely  with  the  bases  united  in  pairs,  subfusiform  or  narrowly 
cylindrical,  often  curved,  2-9 mm.  x  .35-1.2  mm.,  smooth,  furrowed, 
or  rarely  lamellate,  mouth  repand  or  scarious  and  erose-lacerate:  cap- 
sule dusky  brown  to  black,  2-9  cm.  (mostly  3-6  cm.)  x  .2  5-.5  mm., 
rather  long-pedicellate,  valves  commonly  a  little  twisted  on  drying, 
stomata  abundant,  the  guard-cells  nearly  colorless  ;  spores  dark- 
brown  or  black,  45-63  fj.  in  maximum  diameter,  angular,  the  con- 
vex face  with  very  numerous  (125-225)  spines  or  papillae  less  than 
3  11  in  length,  plane  faces  granulose-papillate  or  sinuate-foveolate  ; 
pseudo-elaters  fuscous,  60-250  a  x  8-14  (i,  of  1-4  cells,  genicu- 
late and  variously  contorted,  sometimes  branched. 

Exsicc.   Hep.  Am.  163. 

On  moist  banks.  Common  on  the  Pacific  Coast  from  southern 
California  to  British  Columbia. 

Mill  Valley  (15,  1208),  Ross  Valley  (1207);  Ukiah  (774), 
Pieta  (808),  near  Mendocino  (616  in  part,  630  b,  664);  Eureka 
(935)  ;  Douglas  City,  Trinity  Co.  (1137)  ;  Palo  Alto  (Campbell)  ; 
Felton,  Santa  Cruz  Co.  (Farlow) ;  bank  of  Kaweah  River,  Tulare 
Co.  (Coville  and  Funston,  Death  Valley  Expedition,  no.  1294 — a 
specimen  with  unusually  abundant  and  foliaceous  lamellae)  ;  Devil's 
Canon,  San  Bernardino  Mountains  (Parish,  2305),  near  Rochester 
(Parish);  Pasadena  (McClatchie);  San  Diego  (Orcutt). 

AiitJwceros  fusiformis  is  a  close  ally  of  A.  punctatiis  L.,  differ- 
ing in  the  larger,  thicker,  more  dissected,  and  usually  more  lamel- 


188 


Anthocerotaceae 


late-cristate  thallus,  the  rarely  geminate,  often  longer  involucres, 
the  longer  capsules,  the  rather  larger,  more  minutely  and  thickly 
echinulate  or  papillate  spores,  and  in  the  logner,  narrower  elaters. 
In  the  southern  part  of  California  it  makes  its  nearest  approach 
to  A.  piinctatiis,  but  may  be  distinguished  from  that  species  after  a 
little  experience  by  the  characters  of  the  spores  and  elaters.  The 
involucres  in  the  Californian  plants  are  rarely  more  than  5  mm. 
long.  A.  fusiforinis  ripens  its  spores  in  California  in  April  and 
May,  at  which  time  the  thallus  has  commonly  become  shriveled 
and  inconspicuous. 

Following  Austin's  original  description  of  Anthoceros  fusiformis 
three  specimens  are  cited,  as  follows  :  Observation  [Observa- 
tory] Inlet,  Columbia,  Dr.  Scouler  in  herb.  Torrey ;  also  in  herb. 
Taylor.  California,  Bolander.  Oregon,  E.  Hall."  What  appear 
to  be  these  three  originals  are  preserved  in  herb.  Pearson.  Hall's 
specimen  is  marked  R.  Mts.,  Hall,"  but  this  manner  of  giving 
the  habitat  of  Hall's  Oregon  Hepaticae  is  observed  elsewhere  in 
the  Austin  collection. 

Plate  122.    Anthoceros  fusiformis. 

1.  Plant,  natural  size. 

2.  Portion  of  thallus,  X  23 

3.  Anterior  margin  of  thallus-segment  in  vertical  section,  X  53- 
4  and  5.  Dorsal  lamellae  in  vertical  section,  X  53- 

6.  Nearly  median  longitudinal  section  of  thallus-segment,  X  53- 
7-9.  Involucres,  X  ^2. 

10.  Vertical  longitudinal  section  near  thallus-apex,  showing  antheridia  and  arche- 
gonia,  X  53- 

11.  Spores,  X  305- 

12.  Pseudo-elaters,  X  305- 

Figs.  I  and  9  drawn  from  no.  1207  (Ross  Valley,  Marin  Co.)  ;  2,  from  no.  1208 
(Mill  Valley,  Feb.  22,  1896)  ;  3-6,  and  10,  from  microtome  sections  of  material  col- 
lected in  Ross  Valley  ;  7,  from  no.  808  (Pieta,  Mendocino  Co.) ;  8,  from  a  specimen 
collected  in  California  by  Dr.  Bolander;  II  and  12,  from  no.  15  (Mill  Valley). 


Bibliography 


189 


BIBLIOGRAPHY. 
I. 

List  of  papers,  treatises,  etc.   [including  exsiccatae]  in  which 

THE  HePATICAE  AND  AnTHOCEROTES  OF  CALIFORNIA  ARE  DESCRIBED 
OR  MENTIONED. 

Austin,  C.  F.  Characters  of  some  new  Hepaticae  (mostly  North 
American)  together  with  notes  on  a  few  imperfectly  described 
species.    Proc.  Acad.  Nat.  Sci.  Philad.  1869:  218-234. 

 New  Hepaticae.    Bull.  Torr.  Bot.  Club,  3:  9-18.  1872. 

  Hepaticae  Boreali-Americanae  exsiccatae.    Nos.  1-150.  1873. 

  New  Hepaticae.    Bull.  Torr.  Bot.  Club,  6:  17-21.  1875. 

  Notes  on  the  Anthocerotaceae  of  North  America,  with  de- 
scriptions of  several  new  species.  Bull.  Torr.  Bot.  Club,  6 :  25- 
29.  1875. 

  New  Hepaticae.    Bull.  Torr.  Bot.  Club,  6  :  46,  47.  1875. 

  Notes  and  Criticisms  on  Hepaticae   Americanae  exsiccatae. 

Bull.  Torr.  Bot.  Club,  6  :  85.  1876. 
  Notes  on  Hepaticology.    Bull.  Torr.  Bot.  Club,  6  :  301-306. 

1879. 

Bolander,  H.  N.  A  Catalogue  of  the  Plants  growing  in  the  vicinity 
of  San  Francisco,  pp.  1-43.  San  Francisco.  1870.  (This  ap- 
pears to  have  been  published  originally  in  the  California  Medical 
Gazette.  1870.) 

Campbell,  D.  H.    The  Structure  and  Development  of  the  Mosses  and 

Ferns.    Svo.  pp.  544.  1895. 
  A  new  Californian  Liverwort.  Bot.  Gaz.  21  :   9-13.  //.  2. 

1896. 

  ^oXt?,  on  Sphaerocarpus.    Erythea,  4  :  73-78.  //.  2.  1896. 

  The  Development  of  Geoihallus  tuberosus  Campbell.  Ann. 

Bot.  ID:  489-510.  //.  24,  25.  1896. 
  Recent  Work  upon  the  Development  of  the  Archegonium. 

Bot.  Gaz.  26:  428-431.  1898. 
Coville,  F.  V.    Botany  of  the  Death  Valley  Expedition.  Contrib. 

U.  S.  Nat.  Herb.  4:  1-318.    //.  7-p.  1893. 

Contains  list  of  ten  Hepaticae  on  p.  230. 


190 


Bibliography 


Davidson,  Alice  M.  California  Plants  in  their  Homes :  A  botan- 
ical Reader  for  Children.    8vo.  pp.  216;  Suppl.  pp.  133.  1898. 

Evans,  A.  W.  A  Revision  of  the  North  American  Species  of  Frul- 
la?iia,  a  Genus  of  Hepaticae.  Trans.  Conn.  Acad.  10  :  1-39.  pi. 
1-13.  1897. 

Goebel,  K.  Organographie  der  Pflanzen.  Zweiter  Teil.  Specielle 
Organographie.     i  Heft:  Bryophyten.  1898. 

Refers  to  Geothalliis  tuberosus  Campbell,  pp.  291,  293. 

Howe,  M.  A.  Two  Californian  Cryptogams.  Erythea,  i  :  112,  113. 
1893. 

  Notes  on  Californian  Bryophytes — I.    Erythea,   2  :  97-101. 

//.  I,  2.  1894. 

  Notes  on  Californian  Bryophytes — H.   Erythea,    4 :  48-54. 

1896. 

  Gyrofhyra,  a  new  Genus  of  Hepaticae.    Bull.  Torr.  Bot. 

Club,  24:  201-205.  P^'  302,  joj.  1897. 
  Notes  on  Californian  Bryophytes — HI.      Erythea,  5  :  87- 

94.       3.  1897. 

  The  North  American  Species  of  Fo?'ella.    Bull.  Torr.  Bot. 

Club,  24:  512-527.  1897. 
  The  Anthocerotaceae  of  North  America.     Bull.  Torr.  Bot. 


Club,  25  :  1-2^.  pL  321-326.  li 
  New  American  Hepaticae.    Bull.  Torr.  Bot.  Club,  25:  183- 


1^2.  pi.  336,  337.  1898. 
McClatchie,  A.  J.    Flora  of  Pasadena  and  vicinity.    Reid,  History 

of  Pasadena,  605-649.  1895. 
 Seedless  Plants  of  Southern  California.    Proc.  So.  Calif.  Acad. 

Sci.  I  :  337-395-  1897. 
McFadden,  Effie  B.  The  Development  of  the  Antheridium  of  Ta7'gioiiia 

hypophylla.  Bull.  Torr.  Bot.  Club,  23  :  242-244,  //.  268.  1896. 
Solms-Laubach,  H.  Graf  zu.    Die  Marchantiaceae  Cleveideae  und  ihre 

Verbreitung.    Bot.  Zeit.  57^  15-37.  1899. 
Stephani,  F.   Die  Gattung  Radula.  Hedwigia,  23  :  1 13-1 16;  1 29-137  ; 

145-159;  161-163.  1884. 

Contains  description  of  Raditia  Bolatidcri, 

  Cryptomitrium  tencruvi  Aust.    Bot.  Gaz.  17  :  58-60.  1892. 

 Species  Hepaticarum.    Bull.  Herb.  Boiss.  6:  309-343;  361- 

378;  757-799-    1898;  7:  84-110;  198-225;  381-407.  1899. 

This  work  is  being  continued  and  is  designed  to  include  descriptions  of  all  known 
Hepaticae. 


Bibliography 


191 


Sullivant,  W.  S.  Description  of  the  Mosses  and  Liverworts.  Rep.  Bot. 
[Whipple's]  Exped.;  Pacific  R.  R.  Survey,  4:    185-193.  ^S^j. 

The  seven  Hepaticae  here  enumerated,  without  locality,  were,  doubtless,  in  part 
collected  in  California. 

Underwood,  L.  M.  North  American  Hepaticae.  Bot.  Gaz.  7:  18-21. 
1882. 

  Descriptive  Catalogue  of  the  North  American  Hepaticae,  north 

of  Mexico.    Bull.  111.  State  Lab.  Nat.  Hi;3t.  2:  1-133.  1884. 
  Some  undescribed  Hepaticae  from  California.  Bot.  Gaz.  13: 

1 1 2-1 14.    J>/.  3-6.  1888. 
  The  Distribution  of  the  Hepaticae  of  North  America.  Proc. 

Amer.  Assoc.  Adv.  Sci.  39:  298-304.  1891. 
  A  preliminary  List  of  Pacific  Coast  Hepaticae.     Zoe,  i  :  361- 

367.  1891. 

  A  preliminary  Comparison  of  the  Hepatic  Flora  of  boreal  and 

sub-boreal  regions.    Bot.  Gaz.  17:  305-312.  1892. 

  Notes  on  our  Hepaticae. — II.   The  genus  Riccia.    Bot.  Gaz. 

19:  273-278.  1894. 
  Notes  on  our  Hepaticae. — III.    The  Distribution  of  the  North 

American  Marchantiaceae.    Bot.  Gaz.  20  :  59-71.  1895. 

  Notes  on  our  Hepaticae. — IV.  Fossomb7'onia.  Bot. 

Gaz.  21  :  67-71.  1896. 
  The  genus  Cephalozia  in  North  America.    Bull.  Torr.  Bot. 

Club,  23  :  381-394.  1896. 
Underwood,  L.  M.,  &  Cook,  0.  F.   Hepaticae  Americanae  [Exsiccatae] . 

Nos.  1-200.  1887-1899. 

II. 

List  of  the  principal  papers,  treatises,  etc.  (including  exsic- 
catae), WHICH  have  been  CONSULTED,  IN  ADDITION  TO  THE  FORE- 
GOING. 

o 

Angstrom,  J.    Primae  lineae  muscorum  cognoscendorum,  qui  ad  Cal- 

das  Brasiliae  sunt  collectae.    II.    Hepaticae.    Ofversigt  af  Kongl. 

Vetensk.-Akad.  Forhandl.  1876":  77-92. 
Arnell,  H.  W.    Cm  ndgra  Jungermania  venfricosa  Dicks,  narstdende 

lefvermossarter.    Bot.  Notis.  1890:  97-104.  1890. 
 Lebermoosstudien  im  nordlichen  Norwegen.  4to.    pp.  x,  44. 

1892. 


192 


Bibliography 


Arnell,  H.  W.    Om  slaktnamnet  Porella  Dill.,  Lindb.    Bot.  Notis. 

1893:  127-132.  1893. 
 S.  F.  Gray's  levermoss-slakten.    Bot.  Xotis.  1893:  137-151. 

1893. 

Ashworth,  J.  H.  On  the  Structure  and  Contents  of  the  Tubers  of 
Anthoceros  tiiberosus  Taylor.  Mem.  and  Proc.  Manchester  Lit. 
and  Phil.  Soc.  41  :  — (1-6). //.  2.  1896. 

Austin,  C.  F.    Notes  on  Hepaticology.    Bot.   Bulletin  (now  Bot. 

Gazette),  i  :  35,  36.  1876. 
 New  Hepaticae.    Bull.  Torr.  Bot.  Club,  6:  157,  158.  1877. 

Bellardi,  C.  A.  L.    Appendix  ad  Floram  Pedemontanam.    Mem.  Acad. 
Roy.  Sci.  Turin,  5:  209-286.  1793. 
This  paper,  it  would  appear,  was  published  as  a  separate  in  1792. 

Berggren,  S.  Musci  et  Hepaticae  Spetsbergenses.  Kongl.  Sv.  Vetensk. - 

Akad.  Handl.  13'  :  1-103.  1875. 
Bescherelle,  E.    Enumeration  des  Hepatiques  connues  jusqu'ici  aux 

Antilles  Fran^aises  (Guadeloupe  et  Martinique).  Jour,  de  Bot.  7: 

174-180;  183-194.  1893. 
Bischoff,  G.W.    Beobachtungen  iiber  Sphaerocarpus  terrestris  Michel. 

Acta  Acad.  Caes.  Leop. -Carol.  Nat.  Cur,  13:  829-838.  //.  44. 

1827. 

 Bemerkungen  iiber  die  Lebermoose,  vorziiglich  aus  den  Grup- 

pen  der  Marchantieen  und  Riccieen.  Acta  Acad.  Caes.  Leop.- 
Carol.  Nat.  Cur.  17:  911-1088.  //.  ^57-77.  1835. 

Campbell,  D.  H.  On  the  relationships  of  the  Archegoniata.  Bot. 
Gaz.  16:  323-333.  1891. 

 On  the  Structure  and  Development  of  Dendroceros  Nees. 

Jour.  Linn.  Soc.  33:  467-478.  //.  27,  22.  1898. 

Camus,  F.  Sur  le  Riccia  nigrella.  Bull.  Soc.  Bot.  France,  39 : 
212-230.  1892. 

Cardot,  J.    Sur  les  Riccia  Bischoffii  Hub.  et  R.  nodosa,  Bouch.  Rev. 

Bryol.  19  :  49-53-  1892. 
Carrington,  B.  Irish  Hepaticae.  Trans.  Bot.  Soc.  Edinb.  7:  441-458. 

//.  JO,  II.  1863. 
 Dr.  Gray's  Arrangement  of  the  Hepaticae.    Trans.  Bot.  Sci. 

Edinb.  10  :  305-309.  1870. 
 On  two  new  British  Hepaticae.    Trans.  Bot.  Soc.  Edinb.  10  : 

378-382.  1870. 


Bibliography 


193 


Carrington,  B.    New  British  Hepaticae.    Grevillea,  2  :  85-88. //.  7(5*. 

 British  Hepaticae,  containing  descriptions  of  the  native  species 

of  Jungermannia,  Marchantia,  and  Anthoceros.  Pp.  88.  //.  16^ 
1874-1876. 

 Notes  on  new  British  Hepaticae.    Trans.  Bot.  Soc.  Edinb.  13  : 

461-470.  //.  77,  18.  1879. 

Carrington,  B.,  and  Pearson,  W.  H.    Hepaticae  Britannicae  exsiccatae. 

Nos.  1-290.  1878-1890. 
  List  of  Hepaticae  collected  by  Mr.  Thomas  Whitelegge  in  New 

South  Wales,  1884-5.    Proc.  Linn.  Soc.  of  New  South  Wales,  H. 

2:  1035-1060.  //.  22-3J.  1887. 
Carruthers,  W.    On  the  nomenclature  of  the  British  Hepaticae.  Jour. 

Bot.  3  :  297-302.  1865. 
Corda,  A.  K.  J.    Genera  Hepaticarum  :  Die  Gattungen  der  Leber- 

moose.    Opiz,  Beitrage  zur  Naturgeschichte,  643-655.    1829. -j^ 

(We  have  seen  only  a  pencil  copy  of  this  paper. ) 

  Monographia  Rhizospermarum  et  Hepaticarum.    4to.  pp.  16. 

//.  6.    Prag,  1829. 
DeCandolle,  A.  P.,  &  Lamarck,  J.  B.  de.    Flore  Frangaise.    6  vols. 

1815.  (3ded.) 

De  Notaris,  G.    Primitiae  Hepaticologiae  Italicae.    Mem.  Accad.  Sci. 

Torino,  IL  i  :  287-354.  //.  1839. 
  Appunti  per  un  nuovo  censimento  delle  Epatiche  Italiane. 

Mem.  Accad.  Sci.  Torino,  K.  18:  457-498.  //.  i-io.     1859;  22: 

353-389-  pi-  1-28.  1865. 
Dickson,  J.    Fasciculi  [IV]  Plantarum  Cryptogamicarum  Britanniae. 

4to.  pp.  28,  31,  24,  28.  //.  72.  1785-1801. 
  Observations   on  the  Genus  of  Porella  and  the  Phase  urn 

caitlescens  of  Linnaeus.    Trans.  Linn.  Soc.  3  :  238-239.  //.  20. 

1797. 

Dillenius,  J.  J.  Historia  Muscorum,  in  qua  circiter  sexcentae  species 
veteres  et  novae  ad  sua  genera  relatae  describuntur,  et  iconibus 
genuinis  illustrantur  :  cum  appendice  et  indice  synonymorum.  4to. 
pp.  xvi,  576.  pi.  8j.  1 741.  (Only  the  Edinburgh  reprint  of 
1811  and  the  abridged  English  edition  of  1763  have  been  seen.) 

Dumortier,  B.  C.    Commentationes  Botanicae.    8vo.  pp.  116.  1822. 


*  See  note  under  Ricciocarpus,  page  33 . 


194 


Bibliography 


Dumortier,  'B.  C.    Sylloge  Jungermannidearam  Europae  indigenarum, 
earum  genera  et  species  systematice  complectens.     8vo.  pp.  loo. 
//.  2.     I 83 1. 

  Recueil  d'  Observations  sur  les  Jungermanniacees.     8vo.  pp. 

27-  1835. 

  Hepaticae  Europae.    Jungermannideae  Europae  post  semise- 

culum  recensitae,  adjunctis  Hepaticis.    Bull.  Soc.  Roy.  de  Bot.  de 

Belgique,  13  :  5-203.//./-^.     1874.     (Also  published  separately 

with  the  same  pagination. ) 
Ehrhart,  F.    Beitrage  zur  Naturkunde,  etc.    7  vols.   8vo.  1787-1792. 
Ekart,  T.  P.    Synopsis  Jungermanniarum  in  Germania  vicinisque  terris 

hucusque  cognitarum.    4to.  pp.  72,  //.  ij.  1832. 
Evans,  A.  W.    An  Arrangement  of  the  Genera  of  Hepaticae.  Trans. 

Conn.  Acad.  8:  262-280.     1892.     (Sep.  pp.  2-20.) 
  Two  new  American  Hepaticae.    Bull.  Torr.  Bot.  Club,  20  : 

307-310.  //.  162,  163.  1893. 
 Notes  on  the  North  American  Species  of  Plagiochila.  Bot. 

Gaz.  21  :  185-194.  //.  75,  16,  1896. 
Gayet,  L.  A.    Recherches  sur  le  Developpement  de  I'Archegone  chez 

les  Muscinees.    Ann.  Sci.  Nat.  Bot.  VHI.  3  :  161-258.  //.  7-13. 

1897. 

Goebel,  K.    Zur  vergleichenden  Anatomic  der  Marchantieen.  Arb. 

Bot.  Inst.  Wiirzburg,  2  :  529-535.  1882. 
 Die  Muscineen.    Schenk,  Handbuch  derBotanik,  2  :  315-401. 

1882. 

Gottsche,  C.  M.    Ueber  die  Fructification  der  Jungermanniae  Geocaly- 

ceae.    Acta  Acad.  Caes.  Leop. -Carol.  Nat.  Cur.  21  :  419-466. 

//.  30-32.  1845. 
 Uebersicht  und  kritische  Wiirdigung  der  seit  dem  Erscheinen 

der  Synopsis  Hepaticarum  bekannt  gewordenen  Leistungen  in  der 

Hepaticologie.    Bot.  Zeit.  16:  Beilage,  1-49.  1858. 
 De  Mexikanske  Levermosser.    Efter  Prof.   Fr.  Liebmanns 

Samling  beskrevne.    Dansk.  Vid.  Selsk.  Skrift.  V.  6:  97-380.  //. 

1-20.     1863.     (Sep.  pp.  284.) 
 Ueber  die  Cuticula  der  Scapania-Arten.    Hedwigia,  5:  17-23. 

1866. 

 Einige  Bemerkungen  zu  Thorn.  Jensen  :  Conspectus  Hepatica- 
rum Daniae,  eller  Beskrevelse  af  de  Danske  Halvmosser.  Hed- 
wigia, 6  :  49-59;  65-77.  1867. 


Bibliography 


195 


Gottsche,  C.  M.    Neuere  Untersuchungen  liber  die  Jungermanniae  Geo- 
calyceae.    Abhandl.  des  Naturw.  Vereins  Hamburg,  7  :  39-66. 
1880. 

Gottsche,  C.  M.,  Lindenberg,  J.  B.  G.,  &  Nees  ab  Esenbeck,  C.  G. 

Synopsis  Hepaticarum.     8vo.  pp.  834.  1844-1847. 
Gottsche,  C.  M.,  &  Rabenhorst,  L.    Hepaticae  Europae  [exiccatae]. 

Nos.  1-660.  1862-1879. 
Gray,  S.  F.    A  Natural  Arrangement  of  British  Plants.     2  vols.,  8vo. 

1821. 

Griffith,  W.    Notulae  ad  Plantas  Asiaticas.    Part  II.    On  the  higher 

Cryptogamous  Plants.     8vo.  pp.  257-628.  1849. 
Hartman,  C.  J.    Plandbok  i  Skandinaviens  Flora,  innefattande  Sveriges 

och  Norriges  Vexter,  till  och  med  Mossorna.    8vo.   pp.  xxviii, 
cliv,  408.//.  2.     1832  [ed.  2]. 
Heeg,  M.     Mittheilungen  liber   einige  Arten  der  Gattung  Riccia. 

Bot.  Notis.  1898:  15-24;  107-116.  1898. 
Hooker,  J.  D.    Handbook  of  the  New  Zealand  Flora.     8vo.  pp. 

Ixviii,  798.  1864-1867. 
Hooker,  W.  J.    British  Jungermanniae  :  being  a  history  and  description 

with  colored  figures  of  each  species  of  the  genus  and  microscopical 

analyses  of  the  parts.    Yo\\o,  pi.  88.     18 16. 
Hudson,  W.    Flora  Anglica.    8vo.  pp.  viii,  506.  1762. 
Hubener,  J.  W.  P.    Hepaticologia  Germanica,  oder  Beschreibung  de 

Deutschen  Lebermoose.    8vo.  pp.  314.  1834. 
Husnot,  T.    Hepaticologia  Gallica,  flora  analytique  et  descriptive  des 

Hepatiques  de  France  et  de  Belgique,  accompagnee  de  13  planches 

representant  chaque  espece  de  grandeur  naturelle  et  ses  principaux 

caracteres  grossis.    8vo.  pp.  102.  //.  ij.     187 5-1 881. 

 Hepaticae  Galliae  (exsiccatae).    Nos.  1-200. 

Jack,  J.  B.    Die  Lebermoose  Badens.    Bericht.  Naturf.  Gesellsch. 

Freiburg,   .     1870.     (Sep.  pp.  92.) 

 Hepaticae  Europae.    Jungermannideae  Europae  post  semisaecu- 

lum  recensitae,  adjunctis  Hepaticis,  auctore  B.  C.  Du  Mortier. 

Bot.  Zeit.  35:  49-59;  65-74;  81-93;  97-1 1 1.  //.  I-  1877. 
 Die  Europaischen  Radula-Arten.    Flora,  64  :  353-362.  //. 

8.     1881.    (Sep.  pp.  26.) 
 Beitrage  zur  Kenntniss  der  Pellia-Arten.     Flora,   81  :  1-16. 

//.  I.  1895. 

Jack,  J.  B.,  Leiner,  L.,  &  Stizenberger,  E.    Kryptogamen  Badens 
[Exsicc] . 


196 


Bibliography 


Kaalaas,  B.    Levermosernes  udbredelse  i  Norge.    Nyt  Mag.  Natur- 

vidensk.  33  :  1-490.  1893. 
  Beitrage    zur   Lebermoosflora    Norwegens.  Videnskabssels- 

kabets  Skrifter.  I.  Math.-nat.  Kl.  1898.  Xo.  9.  (Sep.  pp.  28.) 
Kunth,  C.  S.    Synopsis  Plantarum  quas  in  itinera  ad  plagam  aequi- 

noctialem  Orbis  Novi  coUegerunt  Alexander  de  Humboldt  et  Aime 

Bonpland.    4  vols.,  8vo.     182 2-1 8 2 5. 

Kuntze,  0.  Revisio  Generum  Plantarum,  etc.  Vols.  I-Iir\  8vo. 
1891-1898. 

Lamarck,  J.  B.  A.  P.  M.  de,  &  Poiret,  J.  L.  M.  Encyclopedic  me- 
thodique  Botanique.     13  vols.,  4to.     1 783-1 8 17. 

Lehmann,  J.  G.  C.  Novarum  et'  minus  cognitarum  stirpium  Pugillus. 
I-X.    4to.  1828-1857. 

Leitgeb,  H.  Untersuchungen  liber  die  Lebermoose.  I-VI.  4to. 
1874-1881. 

Le  Jolis,  A.  Du  nom  de  genre  Porella.  Rev.  Bryol.  19:  97-10 1. 
1892. 

 Les  genres  d'Hepatiques  de  S.  F.  Gray.     Mem.  Soc.  nation. 

Sci.  nat.  et  math.  Cherbourg,  29:  1-36.  1893. 
  La  nomenclature  des  Hepatiques.     Rev.  Bryol.  21  :  65-68. 

1894. 

 Remarques  sur  la  nomenclature  hepaticologique.    Mem.  Soc. 

nation.  Sci.  nat.  et  math.  Cherbourg,  29:  105-182.  1894. 

 Porella  once  more.    Bull.  Torr.  Bot.  Club,  25:  95,  96.  1898. 

  Encore  sur  Porella.'^-    Rev.  Bryol.  25:  43-49.  1898. 

Levier,  E.    Sur  le  Riccia  miniina  L.  {=  R.  uigrella  DC.  et  R.  soro- 

carpaV>\'5>(z\io'ii^.     Rev.  Bryol.  20  :  101-105.  1893. 

 Riccia  Michelii.    Bull.  Herb.  Boiss.  2  :  229-240.  //.  5.  1894. 

  Riccia  Henriquesii.    Bull.  Herb.  Boiss.  2  :  649,  650.  //.  jo, 

31.  1894. 

  La  pseudopriorite  et  les  noms  a  bequilles.    Bull.  Herb.  Boiss. 

4 :  369-406.  1896. 
  Nachtriigliches  liber  falsche  Prioritat  und  Krlickennamen.  Bot. 

Centralb.  71  :  13-24.  1897. 
  La  pseudopriorita  di  Porella.    Bull.  Soc.  Bot.  Ital.     1898  : 

99-106.  1898. 


*  See  also  :  Howe,  M.  A.  Porella  once  more.  Bull.  Torr.  Bot.  Club,  25  :  96- 
103.     1898.    The  Porella  Question.     Rev.  Bryol.  25 :  76-78.  1898. 


Bibliography 


197 


Levier,  E.  Porella  annule  par  le  Code  de  Rochester.  Bull.  Herb. 
Boiss.  6:  496-502.  1898. 

Lightfoot,  J.    Flora  Scotica.     2  vols.,  8vo.  1777. 

Limpricht,  K.  G.  Lebermoose.  Cohn,  Kryptogamen- Flora  von 
Schlesien,  l  :  227-352  ;  429-444.  1877. 

 Die  deutschen  Saiiteiia-Y oxvci^rv.    Flora,  63  :  90-93.  1880. 

  Neue  Arten  imd  Formen  der  Gattung  Sarcoscypltus  Corda. 

Jahresb.  Schles.  Gesellsch.  58:  179,  184.  1881. 

  Einige  neue  Arten  und  Formen  bei  den  Laub-  und  Lebermoosen. 

Jahresb.  Schles.  Gesellsch.  61  :  204-226.  1884. 

Lindberg,  S.  0.    Musci  Novi  Scandinavici.    Not.  Sallsk.  pro  Fauna  et 

Flora  Fenn.  9:  255-299.  1868. 
  Utredning  af  Skandinaviens  Po7'ella-Y oxm^x.    Acta  Soc.  Sci. 

Fenn.  9  :    329-345.  1869. 
  Contributio  ad  Floram  Cryptogamam  Asiae  Boreali-Orientalis. 

Acta  Soc.  Sci.  Fenn.  lO:  223-280.  1872. 
  Manipulus  Muscorum  Secundus.    Not.  Sallsk.  pro  Fauna  et 

Flora  Fenn.  13:  353-418.  //.  i.  1874. 
 Hepaticae  in  Hibernia  mense  Julii  1873  lectae.    Acta  Soc. 

Sci.  Fenn.  10  :  467-559-  i875- 

  Hepaticologiens  Utveckling  fran  aldsta  tider  till  och  med 

Linne.    4to.  pp.  51.  1877. 
  Om  Jungermauia  julacea       /.  nivalis  Sw.,  och  /.  setiformis 

Ehrh.     Med.  Soc.  pro  Fauna  et  Fl.  Fenn.  3:  172,  173.  1878. 

 Musci  Scandinavici  in  systemate  novo  naturali  dispositi.  8vo. 

pp.  50.  1879. 

 Monographia  praecursoria  Peltolepidis,  Sauteriae,  et  Cleveae. 

Acta  Soc.  pro  Fauna  et  Flora  Fenn.  2  :  — (1-15).  1882. 

Lindberg,  S.  0.,  &  Arnell,  H.  W.  Musci  Asiae  Borealis.  I.,  Leber- 
moose.   Kongl.  Sv.  Vetensk.-Akad.   Handl.  23^:  1-69.  1889. 

Lindenberg,  J.  B.  W.  Synopsis  Hepaticarum  Europaearum.  Nova 
Acta  Acad.  Caes.  Leop.-Car.  Nat.  Cur.  14  (Suppl.)  :  i-i 33. 
//.  7,  2.  1829. 

 L'eber  die  Lebermoos-Gattungen    Coisinia  und  Grivialdia. 

Flora,  16  :  161-176.  1833. 

 Monographie  der  Riccieen.  Nova  Acta  Acad.  Caes.  Leop.- 
Car.  Nat.  Cur.  18:  361-504,//.  ig-37.  1836. 


198 


Bibliography 


Lindenberg,  J.  B.  W.,  &  Gottsche,  C.  M.  Species  Hepaticarum.  4to. 
Fasc.  I-XL,//.  67.  1836-185T. 

Linneaus,  C.    Species  Plantarum,  etc.     2  vols.,  8vo.  1753. 

Martelli,  U.    Una  nuova  specie  di  Riccia.    Xuovo  Giorn.  Bot.  Ital. 

21  :  290-292.  1889. 
Martius,  K.  F.  P.  von.    Flora  cryptogamica  Erlangensis.    8vo.  pp. 

Ixxviii,  508.  //.  6.     181 7. 
Massalongo,  C.     Epatiche  rare  e  critiche  delle  provincie  Venete. 

Atti  Soc.  Yen. -Trent.  Sci.  Xat.  5:  —  (1-16).  //.  i,  2.  1877. 
 Hepaticologia  Veneta,  ossia  monografie  delle  Epatiche  cono- 

sciute  nelle  provincie  Venete.    Atti  Soc.  Yen. -Trent.  Sci.  Xat.  6: 

—  (1-68).  pi.  1-3.  1879. 

 Hepaticae  Italiae-Yenetae   Exsiccatae.    Xos.    1-120.  1878- 

1881. 

 Epatiche  raccolte  alia  della  Terra  del  Fuoco  dal  Dott.  C.  Spe- 

gazzini  nell'anno  1882.     X^uovo  Giorn.  Bot.  Ital.  17:  201-277. 

//.  12-2S.  1885. 
 Repertorio  della  Hepaticologia  Italica.    Ann.  R.  1st.  Bot.  di 

Roma,  2:  87-155.  //.  7-p.  1886. 
 Osservazioni  critiche  sulle  specie  e  varieta  di  Epatiche  Italiane 

create  dal  De  X'otaris.    Ann.  R.  1st.  Bot.  di  Roma,  3:  157-169. 

pL  ig.  18S8. 

 Le  specie  Italiane  del  genere  Jungermannia.    Atti  Soc.  Yen.- 

Trent.  Sci.  Xat.  II.  2  :  —  (1-43).  1895. 
 Hepaticae  in  provincia  Schen-si,  Chinae  interioris,  a  Rev. 

Patre  Josepho  Giraldi  collectae,  etc.    Accad.  di  Yerona,  III.  73  : 

—  (1-63)-  pi-  1897- 

Massalongo,  C,  &Carestia,  A.  Epatiche  delle  Alpi  Pennine.  Xuovo 
Giorn.  Bot.  Ital.  12  :  306-366.  //.  8-11.  1880. 

  Epatiche  delle  Alpi  Pennine.     Xuovo  Giorn.  Bot.  Ital.  14  : 

212-258.  //.  10-14.  1882. 

Micheli,  P.  A.  Xova  plantarum  genera,  etc.  4to.  pp.  234.  //.  108. 
1729. 

Milde,  J.  Zur  Kenntniss  von  Anthoceros  und  Blasia.  Bot.  Zeit.  9  : 
629,  630.  1851. 

  Chaniaeceros  ferfi/is  Milde,  novum  Genus  e  Familia  Anthocero- 

teum.  Xova  Acta  Acad.  Caes.  Leop.-Car.  Xat.  Cur.  26:  167- 
174.  //.  12.  1857. 


Bibliography 


199 


Mitten,  W.    Musci  et  Hepaticae  Vitienses.    Bonplandia,  9  :  365-367. 

1861  ;  10  :  19.  1862. 
  Hepaticae  Indiae  Orientalis.    Jour.  Linn.  Soc.  5  :  89-128. 

1861. 

  The  "  Bryologia  "  of  the  Survey  of  the  49th  Parallel  of  Lati- 
tude.    Jour.  Linn.  Soc.  8  :  12-55.  1S65. 

  List  of  Hepaticae  collected  by  the  Rev.  A.  E.  Eaton  at  the 

Cape  of  Good  Hope  (August  and  September,  1874).  Jour.  Linn. 
Soc.  16:  187-196.//.  4,  J.  1878. 

 An  Enumeration  of  all  the  species  of  Musci  and  Hepaticae  re- 
corded from  Japan.  Trans.  Linn.  Soc.  H.  3:  153-206.  pL  57. 
1891. 

Montagne,  J.  F.  C.  Cinquieme  Centurie  de  Plantes  cellulaires  exo- 
tiques  nouvelles ;  Hepaticae.    Ann.  [^Sci.  Nat.  Bot.  HL  4:  346- 

355-  1845- 

Moore,  D.    Report  on  Irish  Hepaticae.    Proc.  Roy.  Irish  Acad.  II.  2  : 

591-672.  //.  43-45-  1S76. 
Necker,  N.  J.  de.    Methodus  Muscorum  per  classes,  ordines,  genera, 

ac  species,  etc.    8vo.  pp.  xvii,  296.     177 J. 

  Elementa  Botanica,  etc.     3  vols.,  8vo.  1790. 

Nees  von  Esenbeck,  C.  G.    Sylloge  observationum  botanicarum,  c.  tab. 

VII.  aeneis.    Horae  physicae  Berolinenses,  etc.,  39-54.  //•  5-1 1- 

1820. 

  Xaturgeschichte  der  Europaischen  Lebermoose.    4  vols.,  8vo. 

1833-1838. 

Pearson,  W.  H.     Cephalozia  Tiinieri  Hook,  in  North  Wales.  Jour. 

Bot.  21  :   no.  1883. 
  Hepaticae  Natalenses.    Christiania  Vidensk.-Selsk.  Forhandl. 

1886.  No.  3.  1-20.  pL  1-12.  1886. 

  Hepaticae  Knysnanae.     Christiania  Vidensk.-Selsk.  Forhandl. 

1887.  No.  9.  1-16.  //.  1-6.  1887. 

  Blepharostoma  palmatuni  Lindb.     Jour.   Bot.    25:  193-195. 

//.  275.  1887. 

  List  of  Canadian  Hepaticae.     8vo.  pp.  31.  //.  13.     16  June, 

1890. 

 A  new  British  Hepatic.    Jour.  Bot.  30  :  353,  354.  //.  J2p. 

1892. 

Petounikow,  A.  Sur  les  organes  reproducteurs  du  Sphaeroca7pus  ter- 
restris  Mich.    Bull.  Soc.  Bot.  de  France,  14  :  137-142.  //.  j.  1867. 


200 


Bibliography 


Pfeffer,  W.    Die  Oelkorper  der  Lebermoose.     Flora,  57:  2-6;  17- 

27  ;  33-43-    P'  I'  1874. 
Rabenhorst,  L.    Deutschlands  Kryptogamen  Flora,   2^:  1-69.  1848. 
Raddi,  G.    Jimgermanniografia  Etrusca.    Atti  Soc.  Ital.  Sci.  Modena, 

18: — (1-45).//.  7-7.     1818.    iSIem.  Mat.  e.  Fis.  Soc.  Ital.  Sci. 

Modena,  18:  14-56.  pi.  1-7.  1820. 
Roll,  J.    Vorlaufige  Mittheilung  iiber  die  von  mir  in  Jahre  1888  in 

Nord  Amerika  gesammelten  neuen  Arten  der  Lebermoose.  Bot. 

Centralb.  45  :  203,  204.  1891. 
Rossetti,  C.    Epaticologia  della  Toscana  Nord-ovest.    Nuovo  Giorn. 

Bot.  Ital.  22:  305-346.  1890. 
Ruppius,  H.  B.    Flora  Jenensis,  etc.    8vo.  pp.  376.    (Only  the  third 

edition — 1745,  edited  by  Haller — has  been  seen.) 
Schiffner,  V.    De  Junoermannia  Hornscliiichiana  N.  ab  E.    Bot.  Cen- 
tralb. 30:  22-25.  //.  I.  1887. 
 Lebermoose  (Hepaticae).  Forschungsreise  S.  M.  S.  "Gazelle." 

IV.  Theil:  Botanik.    4to.  pp.  48.//.  8.  1890. 
 Ueber  exotische  Hepaticae.    Nova  Acta  Caes.   Leop. -Carol. 

Nat.  Cur.  60  :  221-288.  pL  6-ig.  1893. 
 Hepaticae  (Lebermoose).     Engler  &  Prantl,  Die  natiirlichen 

Pflanzenfamilien,     :  3-141.  1893-1895. 
 Expositio  Plantarum  in  itinere  suo  Indico  annis  1893-94  sus- 

cepto  coUectarum,  etc.     Series  prima  Hepaticarum  partem  conti- 

nens.     Denksch.  Math. -Nat.  CI.  Kais.  Akad.  Wissensch.  67:  153- 

263.  1898. 

Schrader,  H.  A.  Systematische  Sammlung  Kryptogamischer  Gewiichse. 
Erste  Lieferung,  pp.  20.  1796.  Zweyte  Lieferung,  pp.  t6. 
1797. 

 Plantae  cryptogamicae  novae,  rariores  aut    minus  cognitae. 

Schrad.  Jour,  fiir  die  Botanik,  1801  :  66-80.  1803. 
Schwaegrichen,  C.  F.    Historia  Muscorum  hepaticorum  Prodromus. 

8vo.  pp.  39.  //.  J.  1814. 
 Ueber  Porella  piimata  L.  oder  Jiuigermannia  Porella  Dicks., 

Madotheca  Porella  Nees.  Linnaea,  13:  113-117.  //.  ij.  1839. 
Schweinitz,  L.  D.,  von.    Specimen  Florae  Americanae  Septentrionalis 

Cryptogamicae;  sistens  :  Muscos  Hepaticos  hucusque  in  Am  Sept. 

observatos.     8vo.  pp.  27.  1821. 
 On  two  remarkable  Hepatic  Mosses  found  in  North  Carolina. 

Jour.  Acad.  Nat.  Sci.  Philad.  2  :  361-370.  //.  — .  1822. 


Bibliography  201 

Scopoli,  J.  A.    Flora  Carniolica,  etc.     2  vols.,   8vo.     1772  [ed.  2]. 

Sowerby,  J.    English  Botany,  etc.    36  vols.,  Svo.  1790-1814. 

Spruce,  R.    The  Musci  and  Hepaticae  of  Teesdale.    Trans.  Bot.  Soc. 
Edinb.  2  :  65-89.  1846. 

'  The  ^Nlusci  and  Hepaticae  of  the  Pyrenees.    Ann.  and  Mag. 

Nat.  His.  n.  3  :     81-106  ;  269-293  ;  358-380  ;    478-503.  p/. 

1- j ;  4  :  104-120.  1849.  Trans.  Bot.  Soc.  Edinb.  3  :  103-216. 
//.  12-14.  1850. 

 On  Anomoclada ,  a  new  genus  of  Hepaticae,  and  on  its  allied 

genera,  Odontoschisma  and  Adelanthiis.  Jour.  Bot.  14:  129- 
136  ;  161-170;  193-203;  230-235.  //.  lyS,  lyg.  1876. 
[Sep.  pp.  1-33.] 

  Musci  Praeteriti  :   sive  de  Muscis  nonnullis  adhuc  neglectis, 

praetervisis  vel  confusis,  nunc  recognitis.  Jour.  Bot.  19  :  33-40. 
i88i. 

 On  MarsKpella  Stableii  n.   sp.   and  some  allied  species  of 

European  Hepaticae.     Rev.  Bryol.  8:  89-104.  1881. 
 On  Cephalozia,  its  sub-genera  and  some  allied  genera.  8vo. 

pp.  vi,  96.  1882. 
  Hepaticae  of  the  Amazon  and  of  the  Andes  of  Peru  and  Ecuador. 

Trans,  and  Proc.  Bot.  Soc.  Edinb.   15  :  i-xi,  1-588.  //.  1-22. 

1885. 

  Hepaticae  Spruceanae  :  Amazonicae  et  Andinae,  annis  1849- 

1860  lectae  [exsiccatae] .  1892. 
Stephani,  F.    Deutschlands  Jungermannien  in  Abbildungen  nach  der 

Natur  gezeichnet,  nebst  Text.     Bericht.  Bot.  Verein.  Landshut, 

7:  93-164.    Fig.  1-131.  1879. 
  Xeue  und  kritische  Arten  der  Gattung  Riccia.    Hedwigia,  24  : 

2-  7.  //.  I.  1885. 

  Hepaticae  Africanae.    Bot.  Jahrb.  8  :  79-95-        3-  f-  1-9 • 

1887. 

  Hepaticae  von  der  Halbinsel  Alaska.  Bot.  Jahrb.  8:  96-99. 

//.  3.  f.  g-ii.  1887. 
 Aiithoceros  Hiisjioti  Stephani,   n.   sp.  Rev.  Bryol.    15 :  49, 

50.  1888. 

  Hepaticae  Africanae.    Hedwigia,  30 :  201-217,  //.  26-30; 

265-272,  pi.  31-34.  1891  ;  31  :  120-130,  //.  3-7  ;  165-174,  //. 
zo-Zj";  198-214, //.  7^-27.  1892. 


202 


Bibliography 


Petounikow,  A.  Hepaticarum  species  novae.  I-IX.  Hedwigia,  32  : 
17-29;  137-147;  204-214;  315-327.  1893;  33:  i-io;  137- 
169.  1894;  34:  43-65  ;  232-253.     1895;    35:  73-140.  1896. 

  Hepaticae  Africanae.    Engl.  Bot.  Jahrb.  20  :  299-321.  1895. 

  Hepaticae  Japonicae.    Bull.  Herb.  Boiss.  5:  76-108.  1897. 

 Hepaticae  Sandvicenses.  Bull.  Herb.  Boiss.  5:  840-849.  1897. 

SuUivant,  W.  S.  Musci  Alleghanienses,  siv^e  spicelegia  Muscorum 
atque  Hepaticarum  quas  in  Itinere  a  Marylandia  usque  ad  Georgiam 
per  tractus  montium,  A.  D.  ^NIDCCCXLIH,  decerpserunt  Asa 
Gray  et  W.  S.  Sullivant.  [Exsicc]  Nos.  1-292.  1845.  Id. 
[Reprint  of  tickets,  with  ''enumeratio"  substituted  for  ''spicelegia"" 
in  the  general  title],  8vo.  pp.  87.  1846. 

  Contributions  to  the  Bryology  and  Hepaticology  of  North 

America.  Part  I.  Mem.  Am.  Acad.  H.  3:  57-66.  //.  7-5. 
1846.     Part  I L     Ibid.  ^\  169-176.  pL  i-j.  1849. 

  Hepaticae.    A.  Gray,  Man.  Bot.  682-702.  //.  6-8.  1856 

[ed.  2]. 

Szyszylowicz,  I.  "Hepaticae  Tatrenses:"  O  rozmieszczeniu  watro- 
bowcow  w  Tatrach.  Sprawozd.  Kom.  Fizyjogr.  Akad.  Umiej-  19  r 
25-125.  pi.  2-S-  1885. 

Taylor,  T.  DeMarchantieis.  Trans.  Linn.  Soc.  17  :  375-395.  //•  12— 
^S-  1837. 

  On  two  new  species  of  British  Jungermanniae.    Ann.  and 

Mag.  Nat.  Hist.  12:  88-90.  1843. 
  New  Hepaticae.     Lond.  Jour.  Bot.  5:  258-284;  365-417. 

1846. 

Thedenius,  K.  F.  Anmiirkningarom  Herjedalens  Vegetation.  KongL 
Sv.  Vet.  Acad.  Handl.  1838:  24-76.  pi.  i.  1839. 

Trevisan  de  Saint-Leon,  V.  Schema  di  una  nuova  Classificazione 
delle  Epatiche.  Mem.  R.  1st.  Lomb.  Sci.  e.  Lett.  HL  4  :  383— 
451.  1877. 

Underwood,  L.  M.    Notes  on  our  Hepaticae.  L  Northern  Species. 

Bot.  Gaz.  14:  191-198.  1889. 

 Hepaticae.    A.  Gray,  Man.  Bot.  702-732.     1890  [ed.  6]. 

 Recent  work  in  systematic  Hepaticology.    Bot.  Gaz.  17:  218— 

220.  1892. 

•  ^  Index  Hepaticarum — Part  L    Bibliography.    Mem.  Torr.  Bot. 

Club,  4  :  1-91.  1893. 
 The  Pvvolution  of  the  Hepaticae.    Bot.  Gaz.  19:  347-361.  1894. 


Bibliography 


203 


Voigt,  W.  E.  A.  Beitrag  zur  vergleichenden  Anatomic  der  Marchan- 
tiaceen.    Bot.  Zeit.  37:  728-759.   //.  g.  1879. 

Wahlenberg,  G.    Flora  Suecica,  etc.    2  vols.,  8vo.  1824-1826. 

Weber,  F.  Historiae  Muscorum  Hepaticorum  Prodromus.  8vo.  pp. 
160.  1815. 

Weis,  F.  G.    Plantac  Cryptogamicac  Florae  Gottingensis.    8vo.  pp. 
333-  1770- 

Withering,  W.  A  Botanical  Arrangement  of  all  the  Vegetables  natur- 
ally growing  in  Great  Britain,  with  descriptions  of  the  genera  and 
species.  2  vols.,  8vo.  1776.  [Also  later  editions,  with  slightly 
modified  titles,] 

Wright,  C.    Hepaticae  Cubenses  Wrightianae  [exsiccatae] . 


INDEX 


[Pages  on  which  descriptions  are  to  be  found  are  in  heavy  face  type.  ] 


Adelan/hti<:,  is^j 
Aitoiiia  (see  Aytonia) 
Alictdaria,  93 
Androcryphia  lottgiseia,  80 
Aiieura,  70 

latifrons,  73 

viuliijida  major,  72 

viuitifida  ambrosioides,  73 

■balmata  [  =  Rircardia  palmata,  74) 
pinguisy  71 

pinnatijida,  72 
Anthelia,  83,  142 

Juratzkana,  7,  142 
julacea,  143,  144 

nivalis,  143 

Tiirnei-i,  129 
Anthoceros,  69,  178,  179,  184 

caespiti.ius,  1 80,  18 1 

CaroIi7iianus,  184,  1 86 

Carolinianus  occidentalis,  7,  179,  184, 
185,  186 

dichotovms,  8,  69,  1 80,  1 81 

Donnellii,  69,  182 

fusiformis,  7,  179,  187,  188 

Hallii,  184 

/rt'^TW,  180,  184,  186 

Lescurii,  180 

Ludovicianus,  180 

Pearsoni,  7,  179,  183,  184,  185 

phymatodes,  4,  7,  8,  69,  179,  180 
181,  182,  184 

polyinorphus,  181,  183 

punctatiis,  181,  187,  188 

Torj'eyi,  1 80 

iuberosm,  69,  iSl,  182 
Anthocerotaceae,  178 
Anthocerotales,  9,  178 
Anthocerotes,  9,  12,  178 
Antoiria,  l6l 
Aplozia,  97,  98 
Arac/miopsis^  14 1 


Asterella,  8,  34,  39,  46,  49 

Bolanderi,  5,  48,  51,  53,  54 
Californica,  5,  lo,  47,  48,  49,  50 
fragrans,  56 
gracilis,  5,  48,  56 
heniisphaerica,  40 
I        Litidenbergiana,  50 
'         lUidata,  55 

Palmeri,  5,  47,  54,  55,  56 
violacea,  2,  5,  48,  53,  54 
Aytonia  eiythj'osperma  ( =  Plagiochasnia 
erythrospenua  =  Clevea  hyalina, 
37) 

Bellincinia,  160 
Blasia,  63,  77 

pusilla,  6,  78 
Blepharostoma,  84,  139,  141 

arachnoideum,  7,  139,  140,  141 
sejimctmn,  14I 
trichophyllura,  6,  139,  1 41 
I  Blepharozia,  144 

Calobrynvi,  63 
Calypogeia,  13 1 

Trichomanis,  1 3 2,  1 34 
Calxpogeja,  120,  131 
Calypogia,  1 20 
Cavendishia,  1 61 

Cephalozia,  83,  85,  121,  126,  139 
bicuspidata,  6,  123,  124 
crassijlora,  125,  1 26 
divaricata,  6,  87,  123,  127,  129 
divaricata  scabra,  129 
exlcnsa,  1 23 
Lainmersiana,  123,  124 
media,  6,  123,  124,  126,  155 
miiltijiora,  126,  127 
pleniceps,  6,  123,  125,  126 
Turneri,  6,  8,  87,  123,  129 
Cephaloziella,  122 
di^'aricata,  128 

204 


Index 


205 


C/iaetopsis,  139 

trich  ophy  Ua,  139 
C/ieiloscyp/ios,  117 
Chiloscyphus,  84,  117,  120 

lofl!  ocoleoides,  1 1 8 

pallescens,  1 18 

polyanthos,  6,  118,  155 

polyanthos   rivularis,  84,    iig,  120, 
156 

Ciniiniiiiius,  131 

Ti-ic/ioruanis,  132 
Clevea,  34,  36,  48,  49 

hyalina,  5,  37,  38,  39,  48 

hyalina  Calitornica,  5,  38,  48 

Urn  bat  a,  49 

pedicella/a,  38 
Conocephalum,  34,  57,  59 

conicum,  5,  58 
Corslnioideae,  34 
Cryptomitrium,  35,  43,  44 

tenerum,  5,  45 

Dendroceros,  178,  179 
Diplophylleia,  1 12 

Dicksoni,  ill,  1 12 
Diplophyilum,  1 03 

Dicksoni,  III 
Diimortiei  a,  12,  34 
Ditvalia  brcvipedunciilata,  45 

tenera,  45 

Eucalyx,  93 
Eiiccphalozia,  122 

biciispidata,  1 23 

media,  126 
Eiijiingennannia ,  97 
Eunardia,  93 

Fegatella,  57 

conic  a,  58 
Finibi'aria,  46 
Fimb7-iaria ,  46 

Bolanderi,  51,  52 

Californica,  48,  50 

gracilis,  57 

Lindenbergiana,  50 

midata,  55 

Pabneri,  55 

vio/acea,  53 
Fossombronia,  63,  69,  79 

angtilosa,  81 


oistata,  81 

longiseta,  6,  69,  80,  81,  82 
pus  a  la,  81,  82 
Texana,  8 1 
ttiberifera,  69 
IVondraczeki,  81,  82 
Frullania,  83,  169,  170 

Asagi'ayana  alsophila,  173  . 
Asagrayana  Calif omica,  173 
Bolanderi,  7,  171 
Californica,  7,  17 r,  173 
Catalinae,  7,  171,  172 
Franciscana,  7,  171,  175 
Grayana  Californica,  173,  175 
Hal  Hi,  171 

Nisquallensis,  7,  171,  174,  175,  176 
Petahiniensis,  1 7 1 
taniarisci,  I73,  175 
nncifora  Calif oJ-nica,  175 

Geocalyx,  83,  120 

graveolens,  6,  120,  121 
Geothallus,  3,  63,  68,  69,  70 

tuberosus,  4,  5,  68 
Grimaldia,  10,  35,  40,  41 

androgyna,  41,  42,  43 

Californica,  5,  41,  42,  43 

dicJiotoma,  4 1 

fragrans,  41 

pilosa,  41 
Gymnocolea,  103 

inflata,  lio 
Gyrothyra,  84,  89,  91,  92 

Underwoodiana,  6,  89,  92 

HaplojiiitriujH,  63 
Harpanthus  Flotovianiis ,  1 10 
Hepaticae,  9,  12 
Herpetiuiii,  136 

reptatis,  136,  1 37 
Hypenajitron,  47 

Jungermannia,  85,  93,  97,  98,  103 

aniplcxicaulis,  I02,  103 
aquatica,  1 19,  1 20 
asplenioides,  1 13 
Baniriensis,  109,  iio 
bicuspidata,  1 23 
B  la  si  a,  78 

Bolanderi,  6,  98,  99,  100,  106 
complanata,  159 


206 


Index 


concin7tata,  143 
convcxa,  153,  154 
cordifoliay  120 
cremilata,  94 
cremdata  gmcillinia ,  94 
ctirta,  154 

Danicola,  6,  98,  100,  loi,  102,  i 
106 

Dicksoni,  il  i 
divaricata,  127 
emarginata,  88 
extensa,  123,  1 24 
Jila7>ientosa,  138 
Genthiana,  94,  95 
gracillima,  94,  95 
graveolens,  12 1 
heferocolpa,  1 08,  1 09 
heterophylla^  II6 
Hornschtichiana,  Iio 
incisa^  1 04 
injlata^  IIO 
julacea,  1 43 
Juratzkana,  142 
La?nntersiana,  1 23 
/anceolata,  97 
lurida,  1 03 
Miillen,  109 

MuUeri  Danaensis,  loo,  1 08 
Mulleri  heteroco'pos,  loS 
nana,  I03 

navicularis,  1 67,  1 68 
nemorosa,  149 
7iivalis,  143 
obovata,  96 

III,  112 
pallescens,  1 18 
pallescens  rivularis^  II 9 
palmata,  74 
pingtas,  71 
platyphylla,  1 65 
pleniceps,  125 
polyanthos,  Ii8 
porelloides,  1 13 
porphyroleiica ,  105,  I07 
pumila,  6,  98,  99,  100 
reptans,  137 
;  iparia,  97,  99 
rubra,  94,  95,  1 00 
spJiaerocarpa ,  103 


Sprengelii,  132 

Siarkii,  128 

stipulacea,  I  TO 

/^rirt',  102 

tricho77ianis,  132 

t7'ichophylla,  1 39 

Tu7-77eri,  129 

twibrosa,  153,  1 54 

U7ididata,  152 

ve7itricosa,  I05,  I07 

Wattiana,  108 

Woodsii,  136 
JUNGERMANNIACEAE,  62,  82,  83 
Ju7igc7-77ia7iniaceae  ak7'ogy7iae  (  =Junger- 

manniaceae,  82),  91 
Jungt7-77ia7i7tiaceae  a7iakrogy7iae ,  63 
JUNGERMANNIALES,  9,  62,  63 

Kantia,  84,  92,  131 

^2r^«/^7,  135 

calypogeia,  1 33,  1 35 

Sprengelii,  1 33,  1 34 

trich077ia7iis,  6,  132,  1 33,  1 34,  135 

Lejeu7iia,  168,  169 
Lepidozia,  10,  83,  136,  141 

attenuata,  138 

Calif 07-nica,  1 44 

Jila77ie7itosa,  138 

hu77ulli77ia,  139 

Fearsoni,  137 

reptans,  6,  137,  138 
Liochlaena,  97 

lanceolata,  97 
Lophocolea,  83,  84,  107,  114,  117,  120 

bidentata  cuspidata,  II5 

cuspidata,  6,  115,  119 

heterophylla,  6,  84,  115,  116,  117,  119 

Leibergii,  1 16 
Lophozia,  83,  84,  85,  97,  98,  103,  112 

heterocolpa,  6,  104,  106,  108, 109,  no 

incisa,  6,  104 

inflata,  6,  104,  no 

Miilleri,  no 

ovata,  6,  85,  104,  in 

ventricosa,  6,  104,  105,  106,  107 
Lunularia,  34,  59 

cruciata,  5,  60 

7>ulga7-is  (    -  Z.  cruciata') 


Index 


207 


Madotheca,  l6l 

Bolande7'i,  1 62 

Calif ornica,  1 67 

Doitglasii,  167 

7iavictilaris,  167 

riviilaris,  1 63 
^archantia,  34,  60 

androgynoy  41 

conica,  58 

cruciata,  60 

geminata,  6 1 

giacilis,  56 

hejuisphaerica^  40 

hyalina,  37 

Ludwigii,  57 

Oregonensis,  62 

pilosa,  56 

polymorpha,  5,  59,  60,  61,  62 
tenera,  45,  46 
"Marchantiaceae,  13,  34 

M ARCH ANTI ALES,  9,  12 

IMarsupella,  85,  87,  88 

Bolanderi,  6,  85,  86,  87,  88 

emarginata,  6,  86,  87,  88 

sphacelata  ery/hrorhiza ,  87 
Martinellia,  146 

<r(7;^^'^'x^,  153 
Martinellius,  1 12,  146,  157 

decipiens^  1 57 
Mastigophora^  1 36 

Ca/ifornica,  144 

repta?is,  1 36 
Mesophylla,  93 
JVIetzgeria ,  63,  75 

conjugatay  75 
fur c  at  a  y  75 
Metzgeriaceae,  62,  63,  75 
Mniuni  irichomanis,  132,  135 

-Nardia,  85,  88,  92,  93 

Bolanderi^  86 

crenulata,  6,  87,  93,  94,  95,  130 

crenulata  gracillima,  94 
gracinima,  94 
haematostida^  92 
hyalina y  97 

obovata,  6,  94,  96,  97,  loo 
scalarisy  9' 5  93 
J^ardius,  85,  93 

emarginatus,  88 


Notothylas  ,  1 78,  1 79 
Nowellia,  122 

Pellia,  63,  71,  75,  77 

calycina,  77 

epiphylla,  76,  77 

endiviaefolia,  76,  77 

Neesiana,  6,  76,  77 
Petalophylltwi  lajJicllatum,  69 

Preissii,  69 
Plagiochasma  erythrosperma,  37,  38 

pedicellatiis,  38 
Plagiochila,  84,  85,  112,  113,  157 

asplenioides,  6,  113 

porelloides,  1 13 
Platycoaspis,  43 
Pleuroschisma,  136 

reptans,  137 
Porella,  83,  160,  169 

Bolanderi,  7,  161,  162,  163 

dentata,  1 64,  165 

laevigata,  4,  166,  168 

navicularis,  2,  7,  i6l,  163,  167,  168, 
169 

platyphylla,  1 68 

rivularis,  7,  156,  l6i,  163 

Roellii,  7,  8,  161,  163,  164,  165,  166 
Prionolobus,  122 

Turner  i,  129 
Ptilidium,  84,  144 

Californicum,  7,  144,  145 

ciliare,  1 45 
Radula,  84,  112,  146,  157,  168 
arctic  a,  1 59 

Bolanderi,  7,  158,  159,  169 

compianata,  7,   157,  158,  159,  160, 
169 

ICrausei,  160 

spicafa,  158 
Radulum,  157 
Reboulia,  35,  39,  40 

hemisphaerica,  3,  5>  40 
Riccardia,  63,  70 

latifrons,  6,  71,  73,  74,  75,  155 

major,  6,  71,  72 

multifida  ambrosioides,  6,  71,  72,  73 
palmata,  6,  71,  74,  75 
pinguis,  5,  63,  71 
pinnatifida,  70,  72,  73 


208 


Index 


Riccia,  8,  13,  14,  16,  18,  19,  29,  69 

.?^':^'-rt'^^rtA/,  28,  29,  30 
Americana,  5,  14,  24,  25,  26 
arvensis  hirta,  1 8 
Austini,  25 
Bischoffii,  27 
Brandegei,  32 
bulbifera,  69 

Californica,  5,  14,  15,  16,  20,  21,  22, 

Campbelliana,  3,  5,  14,  15,  26,  27, 

28,  30 
cancellata,  69 
Catalinae,  5,  15,  30,  31 
ciliata^  4,  18,  19 
ciliata  intumescens,  1 8,  1 9 
crinita,  1 9 

crystallina,  5,  14,  15,  31,  32 
Jiiiiians,  33 
7vw/»,  32,  33 

glauca,  5,  15,  17,  21,  22,  23 

Gougetiana,  27 

//zV/^/,  18 

intianescens^  1 8 

laniellosa,  25,  27 

lainellosa  A??iencana,  24,  25 

Lescuriana,  5,  15,  16 

Alichelii  ciliaris,  1 9 

minima,  5,  14,  15,  23,  28,  29 

natans,  33 

nigrella,  5,  14,  15,  25,  27,  28,  29,  30 

palniata,  1 9 
pcrennis,  69 
soj'ocarpa,  23,  24 
stibinerinis,  21,  22 

trichocarpa,  5,  8,  14,  15,  17,  18,  I9, 
20,  25,  27,  29 

timiida,  18,  19 

vesicata^  69 

Wafsoni,  33 
RiCCiACKAR,  12,  13,  62 
Ricciocarpus,  33 

natans,  33 

Saccogyna,  120 

grazcolcns,  1 21 
viticulosOy  120 


Sarcoscyphos,  85 

Ehrharti,  88 
Sai'coscyphus,  87 
Saj'coscyphiis  Bo/anderi,  86 
Sauteria,  39)  48 

^7/^/«r/,  39 

linibata,  38,  48,  49 
Scapania,  84,  146,  156,  157 

albescens,  147 

aspeia,  1 50 

Eolanderi,  7,  138,  147,  148,  I49> 

Californica,  147 

caudata,  147 

convex  a,  153 

curta,  7,  147,  154,  155 

glaiicocephala,  153 

heterophylla,  7,  146,  147,  155,  15 

irrigna,  1 52 

nemorosa,  7,  87,  147,  149,  150,  I 
152 

7,  147,  150,  151 

21 1 ig  in  OS  a,  152 

umbrosa,  7,  147,  153,  1 54 

undulata,  7,  I47,  152,  156 

undulata  piupurea,  151 
Selaginella,  9 
Southbya  obovata,  96 
Sphaerocarpus,  62,  63,  64,  69 

Berleri^  65 

Californicus,  5,  65,  66,  67 
cristatus,  3,  5,  64,  65,  66,  67 
.Donnellii,  64,  65,  67 
Michelii,  66 

JMichelii  Californicus,  65 
Sphaerocarpus,  66,  67 
terresiris,  66 

terrestris  Californicus,  65,  67 
Spheiiolobus,  1 12 
Stephanina ,  157 

Targionia,  8,  34,  35 

hypophylla,  5,  8,  36 

Michdii,  36 

.S}!'// aeroca rpus^  6 6 
Telaranea,  14 1 
Thy  ops  i el  la,  173 
Trachycolea,  17 1 


Mem.  Torr.  Box.  Club. 


Pr  .  88. 


RICCIA  TRICHOCARPA    M.  A.  Howe. 


THE  HELIOTYPE  PRINTING  CO.,  BOSTON 


Mem.  Torr.  Bot.  Club. 


Pl.  89. 


RICCIA  CALIFORNICA  Au 


Mem.  Torr.  Bot.  Club. 


Pl.  90. 


THE  HELIOTYPE  PRINTING  CO..  BOSTON 


Mem.  Torr.  Box.  Club. 


Pl. 


THE  HELIOTYPE  PRINTING  CO..  BOSTON 


Mem.  Torr.  Bot.  Club. 


Mem.  Torr.  Bot.  Club. 


PL.  93. 


CRPTOMITRIUM  TENERUM  (Hook.)  Aust. 


Mem.  Torr,  Bot  Club. 


Pl.  94. 


CRPTOMITRIUM  TENERUM  cHook.)  Aust. 


Mem.  Torr.  Box.  Club. 


PL.  95. 


ASTERELLA  CALIFORNICA  (Hampe  )  Under^v.* 


THE  HELIOTYPE  PRINTING  CO..  BOSTON 


Mem.  Torr.  Bot.  Club. 


Pl.  96. 


ASTERELLA  CALIFORNICA  (Hampe  )  Underw. 


THE  HELIOTYPE  PRINTING  CO..  BOSTON 


Mem.  Torr.  Box.  Club. 


PL.  97. 


ASTERELLA  BOLANDERI   fAust.)  Underw. 


Mem.  Torr.  Box.  Club. 


Pl.  98. 


ASTERELLA  VIOLACEA   (Aust.)  Underw. 


THE  HELIOTYPE  PRINTING  CO..  BOSTON 


Mem.  Torr.  Bot.  Club. 


I -15-  ASTERELLA  PALMERI  (Aust.)  Undei 
16-20.    FOSSOMBRONIA  LONGISETA  Aust. 


THE  HELIOTYPE  PRiNTrNG  CO..  BOSTON 


Mem.  Torr.  Bot.  Club. 


Pl.  ioo. 


1-8.  SPHAEROCARPUS  CRISTATUS  M.  A.  Howe. 
9-12.    SPHAEROCARPUS  CALIFORNICUS  Aust. 


THE  HELIOTYPE  PRINTING  CO..  BOSTON 


Mem.  Tork.  Hot.  Club. 


Pl.  loi 


THE  HELIOTYPE  PRIN-^ING  CO..  BOSTON 


MexM.  Torr.  Bot.  Club. 


Pl.  I02. 


GYROTHYRA  UNDERWOODIANA    M.  A.  H 


THE  MELIOTYPE  c,^|mtinG  CO..  BOSTON 


Mem.  Torr.  Box.  Club. 


Pl.  104. 


CEPHALOZIA  PLENICEPS  Aust. 


THE  HELtOTVPE  PRINTING  CO..  BOgT( 


Mem.  Torr.  Box.  Club. 


Pl.  105. 


PTILIDIUM  CALIFORNICUM  (Aust.)  Underw.  and  Cook. 


THE  HELIOTYPE  PRINTING  CO..  BOSTON 


Mem.  Torr.  Bot.  Club. 


Pl.  107. 


SCAPANIA  BOLANDERI  Aust. 


THE  HELIOTYPE  PRINTING  CO..  BOSTON 


Mem.  Torr.  Bot,  Club, 


Pl.  io8. 


SCAPANIA    OAKESIl  Aust. 


THE  HELIOT-»PE  PRINTING  CO..  BOSTON 


Mem.  Torr.  Bot.  Club. 


Pl.  109. 


SCAPANIA   OAKESII  Aust.. 


THE  HELIOTYPE  PRINTING  CO..  BOSTON 


SCAPANIA  HETEROPHYLLA    M.  A.  Howe. 


THE  HELIOTYPE  PRINTING  CO..  BOSTON 


:em.  Torr.  Bot.  Club. 


Pl.  111. 


PORELLA   BOLANDERT   (Aust.)  Pearson. 


Mem.  Torr.  Box.  Club. 


Pl.  112. 


PORELLA   BOLANDERJ  (Aust.  )  Pearson. 


THE  HELIOTYPE  PRINTING  CO..  BOSTON 


Mem.  Torr.  Bot.  Club. 


Pl.  113 


PORELLA  ROELLII  Steph. 

THE  HELIOTYPE  PRINTING  CO..  BOSTON 


Mem.  Torr.  Bot.  Club. 


Pl.  114. 


PORELLA  ROELLII  Steph. 

THE  HELIOTYPE  PRINTING  CO..  BOSTON 


Mem.  Torr.  Box.  Club. 


Pl.  115. 


THE  HELIOTVPE  PRINTING  CO..  BOSTON 


Mem.  Turk.  Box.  Club. 


Pl.  ii6. 


i 


THE  HELIOTYPE  PRINTING  CO..  BOSTON 


Mem.  Torr.  Bot.  Club. 


Pl. 


ANTHOCEROS  PHYMATODES    M.  A.  Howe. 


THE  HELIOTYPE  "-^IN-^ING  CO..  BOSTON 


Mem.  Torr.  Box.  Club. 


Pl. 


9-17.    ANTHOCEROS  PHYMATODES    M.  A.  Howe. 

18.    ANTHOCEROS  POLYMORPHUS    Raddi,  y   DICHOTOMUS  Raddi 


THE  HELIOTYPE  P«ilNTING  CO..  BOSTON 


Mem.  Torr.  Bot.  Cli  b. 


Pl.  119. 


AXTHOCEROS  PEARSOXI    M.  A.  Howe. 


THE  HELIOTYPE  PRINTING  CO..  BOSTON 


Mem.  Torr.  Bot.  Club. 


Pl.  I20. 


ANTHOCEROS  PEARSONI    M.  A.  Howe. 


THE  HELIOTYPE  PRINTING  CO..  BOSTON 


Mem.  Torr.  Box.  Club. 


Pl.  121. 


ANTHOCEROS  CAROLINIANUS  OCCIDENTALIS    M.  A.  Howe. 


THE  HELIQTYPE  PRINTING  CO..  BOSTON 


WELLESLEY  COLLEGE  LIBRARY 


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