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A     HANDBOOK 


OF 


SYSTEMATIC   BOTANY 


Some  Opinions  of  the  Press  on 
Warming's  "  Systematic  Botany." 

"  A  very  complete,  systematic  classification  of  plants, 
with  all  their  principal  characteristics.  It  is  a  book  well 
worth  reading,  and  is  well  printed  and  illustrated." — Lancet. 

"As  a  text  book  of  Botany  this  will  take  a  high  place, 
and  as  a  handbook  of  reference  it  will  prove  valuable.  The 
volume  is  richly  illustrated,  and  is  altogether  a  serviceable 
text  book  for  students." — Knowledge. 

"  Professor  Warming's  contribution  to  morphology  and 
systematic  botany  are  highly  valued  by  botanical  workers, 
who  will  welcome  this  translation  of  his  Handbook  of 
Systematic  Botany.  We  commend  the  book  to  the  notice 
of  those  who  require  to  obtain  a  good  general  knowledge  of 
the  application  of  morphology  to  purposes  of  classification." — 
ttketuettvtt 

"  It  has  been  well  translated,  and  will  be  found  a  useful 
and  trustworthy  handbook."— -Journal  of  Education. 

"  Mr.  Potter  has  done  English  students  of  botany  a 
great  service  in  giving  them  an  excellent  translation  of  Dr. 
Warming's  important  work." — Educational  Review. 


A    HANDBOOK 


OF 


SYSTEMATIC    BOTANY 


BY 

DR.    E.    WARMING 

Professor  of  Botany    in    the    University   of    Copenhagen 

WITH  A  REVISION  OF  THE  FUNGI  BY 

DR.    E.    KNOBLAUCH, 

Karlsruhe 

TRANSLATED  AND  EDITED  BY 
M.   C.   POTTER,   M.A.   F.L.S. 

Professor  of  Botany  in  the  University  of  Durham 

College  of  Science,  Newcastle-upon-  Tyne 
Author  of  "An  Elementary  Text-book  of  Agricultural  Botany 

WITH    610    ILLUSTRATIONS 


XonDon 

SWAN  SONNENSCHEIN  &  CO.  LIM 

NEW  YORK  :  THE  MACMILLAN  CO 

1904 


FIRST  EDITION,  March  1894  ;  SECOND  EDITION,  July  1904 


PBEFACE. 


THE  present  translation  of  Dr.  E.  Warming's  Haandbog  i  den 
Systematiske  Botanik  is  taken  from  the  text  of  the  3rci  Danish 
Edition  (1892),  and  from  Dr.  Knoblauch's  German  Edition  (1890), 
and  the  book  has  been  further  enriched  by  numerous  additional 
notes  which  have  been  kindly  sent  to  me  by  the  author.  Dr. 
Warming's  work  has  long  been  recognised  as  an  original  and 
important  contribution  to  Systematic  Botanical  Literature,  and  I 
have  only  to  regret  that  the  pressure  of  other  scientific  duties 
has  delayed  its  presentation  to  English  readers.  Dr.  Warming 
desires  me  to  record  his  high  appreciation  of  the  careful  transla- 
tion of  Dr.  Knoblauch,  and  his  obligation  to  him  for  a  number  of 
corrections  and  improvements  of  which  he  has  made  use  in  the  3rd 
Danish  Edition.  In  a  few  instances  I  have  made  slight  additions 
to  the  text ;  these,  however,  appear  as  footnotes,  or  are  enclosed 
in  square  brackets. 

In  the  present  Edition  the  Thallophytes  have  been  revised  and 
rearranged  from  notes  supplied  to  me  by  Dr.  Knoblauch,  to  whom 
I  am  indebted  for  the  Classification  of  the  Fungi,  according  to 
the  more  recent  investigations  of  Brefeld.  The  Bacteria  have 
been  revised  by  Dr.  Migula,  the  Floridese  rearranged  after  Schmitz, 
and  the  Taphrinaceae  after  Sadebeck.  The  main  body  of  the  text 
of  the  Algse  and  Fungi  remains  as  it  was  originally  written  by 
Dr.  Wille  and  Dr.  Rostrup  in  the  Danish  Edition,  though  in  many 
places  considerable  alterations  arid  additions  have  been  made. 
For  the  sake  of  comparison  a  tabular  key  to  the  Classification 
adopted  in  the  Danish  Edition  is  given  in  the  Appendix. 

In  the  Angiosperms  I  have  retained  the  sequence  of  orders  in 
the  Danish  original,  and  have  not  rearranged  them  according  to 

w.  B.  '  * 


VI  PREFACE. 

the  systems  more  familiar  to  English  students.  In  any  rearrange- 
ment much  of  the  significance  of  Dr.  Warming's  valuable  and 
original  observations  would  have  been  lost,  and  also  from  a 
teacher's  point  of  view  I  have  found  this  system  of  great  value. 
Although  at  present  it  may  not  be  completely  satisfactory,  yet  as 
an  attempt  to  explain  the  mutual  relationships,  development  and 
retrogression  of  many  of  the  orders,  it  may  be  considered  to  have 
a  distinct  advantage  over  the  more  artificial  systems  founded 
upon  Jussieu's  Divisions  of  Polypetalae,  Gainopetalse,  and 
Apetalae. 

With  reference  to  the  principles  of  the  systematic  arrangement 
adopted,  I  may  here  insert  the  following  brief  communication 
from  the  author  (dated  March,  1890),  which  he  has  requested  me 
to  quote  from  the  preface  of  Dr.  Knoblauch's  edition  : — "  Each 
form  which,  on  comparative  morphological  considerations,  is 
clearly  less  simple,  or  can  be  shown  to  have  arisen  by  reduction 
or  through  abortion  of  another  type  having  the  same  fundamental 
structure,  or  in  which  a  further  differentiation  and  division  of 
labour  is  found,  will  be  regarded  as  younger,  and  as  far  as 
possible,  and  so  far  as  other  considerations  will  admit,  will  be 
reviewed  later  than  the  '  simpler,'  more  complete,  or  richer  forms. 
For  instance,  to  serve  as  an  illustration  :  EPIGYNY  and  PERIGYNY 
are  less  simple  than  HYPOGNY  ;  the  Epigynous  Sympetalse,  Chori- 
petalse,  Monoctyledones  are,  therefore,  treated  last,  the  Hydro- 
charitacecK  are  considered  last  under  the  Helobiece,  etc.  ZYGOMORPHY 
is  younger  than  ACTINOMORPHY  ;  the  Scitaminece  and  Gynandrce 
therefore  follow  after  the  Liliiflorce,  the  Scrophulariacece  after  the 
Solanacece,  Linaria  after  Verbascum,  etc.  FORMS  WITH  UNITED  LEAVES 
indicate  younger  types  than  those  with  free  leaves  ;  hence  the 
Sympetalce  come  after  the  Choripetalse,  the  Silenece  after  the 
Alsinese,  the  Malvacese  after  the  Sterculiacese  and  Tiliacece,  etc. 

"ACYCLIC  (spiral-leaved)  flowers  are  older  than  cyclic  (verticillate- 
leaved)  with  a  definite  number,  comparing,  of  course,  only  those 
with  the  same  fundamental  structure.  The  Feromca-type  must 
be  considered  as  younger,  for  example,  than  Digitalis  and  Antir- 


PREFACE.  Vii 

rhinum,  these  again  as  younger  than  Scrophularia  ;  Verbascum,  on 
the  contrary,  is  the  least  reduced,  and  therefore  considered  as  the 
oldest  form.  Similarly  the  one-seeded,  nut-fruited  Ranunculacece 
are  considered  as  a  later  type  (with  evident  abortion)  than  the 
many-seeded,  folicular  forms  of  the  Order ;  the  Paronychiece  and 
Chenopodiacece  as  reduced  forms  of  the  Alsinece  type ;  and  the 
occurrence  of  few  seeds  in  an  ovary  as  generally  arising  through 
reduction  of  the  many-seeded  forms.  The  Cyperacece  are  regarded 
as  a  form  derived  from  the  Juncacece  through  reduction,  and 
associated  with  this,  as  is  so  often  the  case,  there  is  a  complication 
of  the  inflorescence  ;  the  Dipsacacece  are  again  regarded  as  a  form 
proceeding  from  the  Valerianacece  by  a  similar  reduction,  and 
these  in  their  turn  as  an  off-shoot  from  the  Caprifoliacece,  etc.  Of 
course  these  principles  of  systematic  arrangement  could  only  be 
applied  very  generally ;  for  teaching  purposes  they  have  often 
required  modification." 

In  preparing  the  translation  considerable  difficulty  has  been 
experienced  in  finding  a  satisfactory  rendering  of  several  terms 
which  have  no  exact  equivalent  in  English.  I  may  here  especially 
mention  the  term  Vorblatt  (Porblad)  which  I  have  translated  by 
the  term  bracteole,  when  it  clearly  applied  to  the  first  leaf  (or 
leaves)  on  a  pedicel;  but  in  discussing  questions  of  general 
morphology  a  term  was  much  needed  to  include  both  vegetative 
and  floral  shoots,  and  for  this  I  have  employed  the  term  "  Fore- 
leaf."  Also,  the  term  "  Floral-leaf "  has  been  adopted  as  an 
equivalent  of  "Hochblatt,"  and  the  term  "bract"  has  been  limited 
to  a  leaf  subtending  a  flower. 

At  the  end  of  the  book  will  be  found  a  short  appendix  giving 
an  outline  of  some  of  the  earlier  systems  of  Classification,  with  a 
more  complete  account  of  that  of  Hooker  and  Bentham. 

In  a  book  of  this  character  it  is  almost  impossible  to  avoid 
some  errors,  but  it  is  hoped  that  these  will  be  comparatively  few. 
In  correcting  the  proof-sheets  I  have  received  invaluable  assist- 
ance from  Dr.  Warming  and  Dr.  Knoblauch,  who  have  kindly 
read  through  every  sheet,  and  to  whom  I  am  greatly  indebted  for 


Vlll  PREFACE. 

many  criticisms  and  suggestions.  1  have  also  to  thank  Mr.  I.  H. 
Burkill  for  his  kind  assistance  in  looking  over  the  proofs  of  the 
Monocotyledons  and  Dicotyledons,  and  Mr.  Harold  Wager  for 
kindly  reading  through  the  proofs  of  the  Algse  and  Fungi.  My 
thanks  are  also  especially  due  to  Mr.  B.  L.  Danielsen,  and  I  wish 
to  take  this  opportunity  of  acknowledging  the  very  considerable 
help  which  I  have  received  from  him  in  translating  from  the 
Original  Danish. 

M.  C.  POTTER. 
January,  1895. 


TABLE    OF   CONTENTS. 

BEING  THE  SYSTEM  OF  CLASSIFICATION  ADOPTED  IN 
THE  PRESENT  VOLUME. 

(The  Algce  and  Fungi  re-arranged  in  co-operation  with  Dr.  E.  Knoblauch,  the 
other  Divisions  as  in  the  3rcZ  Danish  Edition.} 


PAGE 

DIVISION  I.     THALLOPHYTA        ...         4 

A.  Sub-Division.     Myxomycetes,  Slime-Fungi 5 

B.  Sub-Division.     Algae. 8 

Class  1.     SYNGENETICJE     .         .  .         .         .         .         .         .14 

,,      2.      DlNOFLAGELLATA 16 

„      3.      DlATOME^E  . 18 

,,      4.       SCHIZOPHYTA         .........          22 

Family  1.     Schizophycese 22 

„      2.    Bacteria 26 

,   Class  5.     CONJUGATE 41 

,,     6.     CHLOKOPHYCEJE 46 

Family  1.     Protococcoideaj 47 

,,      2.     Confervoideee          .......  53 

„      3.     Siphoneae 59 

Class  7.     CHAKACE^; 64 

,,     8.     PH;EOPHYCE.E  (OuvE-BuoAVN  SEAWEEDS)     ....  68 

Family  1.     Phseosporeae 68 

,,       2.     Cyclosporero 73 

Class  9.     DICTYOTALES 76 

,,     10.     RHODOPHYCE^E  (RED  SEAWEEDS) 77 

Family  1.     Bangioideae 77 

2.    Floridese         .    '     .         .         .     '   .         .         .         .78 

C.  Sub-Division.     Fungi          .    " .84 

Class  1.    PHYCOMYCETES  .•  ' 96 

Sub-Class  1.     Zygomycetes "    96 

„         2.     Oomycetes        . 100 

Family  1.     Entomophthorales      .....     102 

„       2.     Chytridiales        .         .         .         .         .         .102 

„       3.     Mycosiphonales .  i         .  104 


TABLE    OF   CONTENTS. 

PAGE 

Class  2.    MESOMYCETES ...  108 

Sub- Class  1.     Hemiasci 108 

2.    Hemibasidii 109 

Class  3.     MYCOMYCETES  (HiGHEB  FUNGI) 114 

Sub- Class  1.     Ascomycetes     ........  114 

Series  1.    Exoasci •        •         •  116A 

„      2.    Carpoasci 118 

Family  1.     Gymnoascales 118 

„         2.     Perisporiales 119 

,,         3.     Pyrenomycetes 125 

4.     Hysteriales 132 

„        5.     Discomycetes 132 

6,    Helvellales  . 136 

Ascolichenes 136 

Sub-Class  2.     Basidiomycetes 144 

Series  1.    Protobasidomycetes 145 

„      2.    Autobasidiomycetes 157 

Family  1.     Dacryomycetes 159 

„       2.    Hymenomycetes 159 

3.     Phalloideje 172 

,,       4.     Gasteromycetes 173 

Basidiolichenes 176 

Fungi  Imperfect! 176 

DIVISION    II.     MUSCINE^E   (MOSSES)       .        .  181 

Class  1.     HEPATIC^ 188 

Family  1.     Marchantiese 190 

,,       2.     Antboceroteee 191 

,,       3.     Jungermanniesa 191 

Class  2.    Musci  FRONDOSI 192 

Family  1.     Spbagnese 193 

,,      2.     Schizocarpese ,t  •  195 

„      3.     Cleistocarpeae 195 

,,      4.     Stegocarpese 195 

DIVISION  III.     PTERIDOPHYTA     .        .        .198 

Class  1.    FILICIN.E 205 

Sub- Class  1.     Filices 205 

Family  1.     Eusporangiatee 210 

,,      2.     Leptosporangiatsa 212 

Sub-Class  2.    Hydropteridece 215 

Class  2.    EQUISEIIN.E  (HORSETAILS)  . 221 

Sub  Class  1.     Isosporous  Equisetince 221 

,,         2.     Heterosporous  Equisetincc         ...  .  225 

Class  3.     LYCOPODINJE  (CLUB  MOSSES) 226 

Sub- Class  1.     Lycnpodlea 22G 

„         2.    Selagmdlece 228 


TABLE    OF   CONTENTS.  XI 

PAGR 

TRANSITION  FROM  THE  CRYPTOGAMS  TO  THE  PHANEROGAMS      .        .        .  234 

Asexual  Generation  of  the  Cormophytes 234 

Sexual  Generation  ;  Fertilisation 243 

DIVISION    IV.     GYMNOSPERM^E     .         .         .251 

Class  1.    CYCADE^E  (CYCADS)     .........  252 

„   2.     CONIFERS  (PINE-TREES) 255 

Family  1.     Taxoideaj 259 

„       2.     Pinoidea 262 

Class  3.    GNETE^I 270 

Fossil  Gymnosperms 271 

DIVISION    V.     ANGIOSPERM^E         .        .        .273 

Class  1.    MONOCOTYLEDONES 274 

Family  1.     Helobiera .278 

„       2.     Glumiflorffl .283 

„      3.     Spadiciflorse 297 

„      4.    Enantioblast89 308 

„      5.    Liliifloraa 309 

„       6.     Scitaminea 323 

„       7.     Gynandrse 328 

Class  2.     DICOTYLEDONES 334 

Sub-Class  1.     Choripetalce 337 

Family  1.     Saliciflorffi 337 

„      2.    Casuariniflone 339 

3.    Querciflorae 340 

,,      4.    Juglandiflorsa 349 

5.     Urticiflorffi 351 

,,,      6.    Poly  goni  floras 358 

,,      7.     Curvembryaa 363 

^,,,      8.'  Cactifloree 375 

„      9.    Polycarpicea 377 

„     10.     Ehoeadinse 393 

„     11.    Cistiflor® 406 

,,     12.     Gruinales .        .416 

,,     13.     Columniferra 421 

„     14.     Tricoccse 430 

„     15.    Terebinthina? 435 

„     16.    Aesculinas 439 

„     17.    Frangulin89 443 

„     18.     ThymelaeineD 448 

„     19.    Saxifraginee 451 

„    20.    EosiflorssK 456 

„     21.    Leguminosse/ 466 

„     22.    Passiflorin® 475 

„    23.     MyrtifloraB 482 

^  „    24.    UmbelliflorsB/ 490 

„    25.    Hysterophyta 498 


xu 


TABLE    OF    CONTENTS. 


Sub-Class  2.     Sympetalce     . 

A.     PentacycliccK    . 
Family  26.     Bicornes    . 
„      27.     DiospyrmsB 
„      28.     Primulinas 

P.     Tetracyclicce    . 
Family  29.     Tubiflorae 
„      30.     Personates 
„      31.     Nuculiferse 
„      32.     Contort®  . 
„      33.     Kubiales    . 
,,      34.     Dipsacales 
„      35.     Campanulinse    . 
„      36.     Aggregates 


APPENDIX 
INDEX 


PAGE 

504 
506 
506 
510 
511 
514 
514, 
517 
531 
541 
548 
556 
560 
564 
574 
593 


COEEIGENDA. 

Page  9,  line  12  from  top,  for  Hydrodicton  read  Hydrodictyon. 
,,    14,  lines  1  and  2  from  top,  for  as  in  the- preceding  case  read  in  this  case. 
,,    14,     ,,     2  and  15  from  top,  for  zygote  read  oospore. 
For  a,  6  and  ii  read  ee,  03  and  ue  throughout. 

The  following  are  not  officinal  in  the  British  Pharmacopoeia : — page  316, 
Draccena  (Dragon's-blood),  Smilax  glabra  ;  p.  321,  "  Orris-root  "  ;  p.  326,  species 
of  Curcuma,  Alpinia  officinarum ;  p.  333,  Orchis-species  ("  Salep  ").  On  page 
206,  par.  4,  only  Pearl  Barley  is  offic.  in  the  Brit.  Phar. 


CLASSIFICATION   OP    THE   VEGETABLE   KINGDOM. 


The  Vegetable  Kingdom  is  arranged  in  5  Divisions. 

Division  I. — Thallophyta,  Stemless  Plants,  or  those  which 
are  composed  of  a  "  thallus,"  i.e.  organs  of  nourishment  which 
are  not  differentiated  into  root  (in  the  sense  in  which  this  term  is 
used  among  the  higher  plants),  stem,  or  leaf.  Vascular  bundles 
are  wanting.  Conjugation  and  fertilisation  in  various  ways; 
among  most  of  the  Fungi  only  vegetative  multiplication. 

In  contradistinction  to  the  Thallophytes  all  other  plants  are  called  "  Stem- 
plants '.'  ("  Cormophyta  "),  because  their  shoots  are  leaf-bearing  stems.  The 
name  Thallophyta  (Stemless-plants)  is  to  some  extent  unsuitable,  since  many 
of  the  higher  Algae  are  differentiated  into  stem  and  leaf. 

The  Thallophytes  are  again  separated  into  3  sub-divisions, 
namely : 

Sub-Division  A.— Myxomycetes,  Slime-Fungi,  with  only  1 

class. 

Sub-Division  B. — Algae,  with  10  classes: 
Class  1.     Syngeneticae. 

„     2.     Dinoflagellata,  Peridinea. 
„     3.     Diatomeae,  Diatoms. 
,,     4.     Schizophyta,  Fission  Algae. 
„     5.     Conjugatae. 
„     6.     Chlorophyceae,  Green  Algae. 
„     7.     Characeae,  Stone-worts. 
,,     8.     Phaeophyceae,  Brown  Algae. 
„     9.     Dictyotales. 
„  10.     Rhodophyceae,  -Red  Algse. 
Sub-Division  C. — Fungi,  with  3  classes: 
Class  1.     Phycomycetes. 
,,     2.     Mesomycetes. 
,.,     3.     Mycomycetes,  Higher  Fungi. 

Division  II. — Bryophyta   or   Muscineae,   Mosses.     These 
have    leaf -bearing    shoots,    but   neither    true   roots    nor    vascular 
w.  B  B 


2  CLASSIFICATION   OF   THE    VEGETABLE    KINGDOM. 

bundles.  The  lowest  Mosses  have,  however,  a  thallus.  Fertili- 
sation is  accomplished  by  means  of  self-motile,  spirally  coiled 
spermatozoids,  through  the  agency  of  water.  From  the  fertilised 
oosphere  a  "  fruit-body  "  (capsule)  with  unicellular  organs  of 
reproduction  (spores)  is  produced.  The  spore  on  germination 
gives  rise  to  the  vegetative  system,  which  bears  the  organs  of 
sexual  reproduction  ;  and  this  system  is  divided  into  two  stages 
— the  protonema,  and  the  leaf-bearing  plant  produced  on  it. 
Alternation  of  generations : 

I.  The    protonema  and    the    entire   nutritive  system    which 

bears  the  organs  of  sexual  reproduction. 
II.  The  capsule-like  sporangium,  with  spores. 

2  Classes  :   1.  Hepaticse,  Liverworts. 

2.  Musci,  Leafy  Mosses. 

Division  III— Pteridophyta  or  Vascular  Cryptogams, 
Fern-like  Plants  having  leaf-bearing  shoots,  true  roots,  and 
vascu,lar  bundles  with  tracheides  and  sieve-tubes.  Fertilisation 
as  in  the  Mosses.  From  the  fertilised  oosphere  the  leaf-bearing 
shoot  arises,  which  bears  on  its  leaves  the  reproductive  organs, 
the  spores,  in  capsule-like  sporangia.  From  the  germination  of 
the  spore  a  small  prothallium  is  formed,  which  bears  the  sexual 
reproductive  organs. 

Alternation  of  generations  : 

I.  Prothallium  with  organs  of  sexual  reproduction. 
II.  Leaf-bearing  shoot  with  capsule-like  sporangia. 

3  Classes:  1.  Filicinas,  True  Ferns. 

2.  Equisetinae,  Horsetails. 

3.  Lycopodinae,  Club-mosses. 

Division  IV. — Gymnospermse.  The  vegetative  organs  are  in 
the  main  similar  to  those  in  the  3rd  Division ;  special  shoots  are 
modified  into  flowers  for  the  service  of  reproduction.  From  the 
oosphere,  which  is  fertilised  by  means  of  the  pollen-tube,  the 
leaf -bearing  plant  is  derived ;  this  passes  the  first  period  of  its 
life  as  an  embryo  in  the  seed,  and  continues  its  development  when 
the  germination  of  the  seed  takes  place.  The  organs  correspond- 
ing to  the  spores  of  the  two  preceding  Divisions,  are  called  respec- 
tively the  pollen-grain  and  embryo-sac.  The  pollen-grains  are 
multicellular  ;  i.e.  they  contain  an  indistinct  prothallium.  In  the 
embryo-sac  a  prothallium,  rich  in  reserve  material  (endosperm),. 


CLASSIFICATION    OF    THE    VEGETABLE    KINGDOM.  3 

with,   female  organs  of  reproduction,  is  developed   BEFORE    FER- 
TILISATION.     The  pollen- grains  are  carried  by  means  of  the  wind 
to  the  ovules;  these  enclose  the  embryo-sac,  and  are  situated  on 
the  open  fruit-leaf  (carpel),  which  has  no  stigma. 
Alternation  of  generations  : 

I.  Pro  thallium  =  Endosperm  in  ovule. 

II.  Leaf-bearing  plant,  with  flowers  which  produce  the  pollen- 
sac  and  ovule. 
3  Classes :  1.  Cycadeaa. 

2.  Conifene. 

3.  Gnetaceae. 

Division  Y. — Angiospermse.  The  members  of  this  group  are 
very  similar  to  those  of  Division  IV.  The  ovules  are,  however, 
encased  in  closed  fruit-leaves  (ovary),  which  have  a  special  portion 
(stigma)  adapted  for  the  reception  and  germination  of  the  pollen- 
grains.  The  pollen-grains  are  bicellular,  but  with  only  a  mem- 
brane separating  the  two  nuclei ;  they  are  carried  to  the  stigma 
by  animals  (chiefly  insects),  by  the  wind,  or  by  some  other  means. 
Endosperm  is  not  formed  till  AFTER  FERTILISATION.  Alternation  of 
generations  in  the  main  as  in  the  Gymnosperms,  but  less  distinct ; 
while  the  sexual  generation,  the  prothallium,  with  the  organs  of 
fertilisation,  is  also  strongly  reduced. 

2  Classes:1  1.  Monocotyledones.     Embryo  with  one  seed-leaf. 
2.  Dicotyledones.     Embryo  with  two  seed-leaves. 

For  a  long  time  the  vegetable  kingdom  has  been  divided  into  CRYPTOGAMS  (so- 
called  because  their  organs  of  reproduction  remained  for  some  time  undis- 
covered), and  PHANEROGAMS  or  Flowering-plants  which  have  evident  sexual 
organs. 

The  first  three  divisions  belong  to  the  Cryptogams,  and  the  third  and  fourth 
divisions  to  the  Phanerogams.  This  arrangement  has  no  systematic  value,  but 
is  very  convenient  in  many  ways. 

The  Cryptogams  are  also  known  as  Spore -plants,  since  they  multiply  by- 
unicellular  organs  (spores),  and  the  Phanerogams  in  contradistinction  are 
called  Seed-plants  (Spermaphyta),  since  they  multiply  by  seeds,  multicellular 
bodies,  the  most  important  part  of  which  is  the  embryo  (a  plant  in  its 
infancy).  Mosses,  Ferns,  and  Gymnosperms  are  together  known  as  Arche.- 
goniatse,  since  they  possess  in  common  a  female  organ  of  distinct  structure, 
the  Archegonium. 

1  See  Angiospermae. 


4  THALLOPHYTA. 

DIVISION  I. 

THALLOPHYTA. 

The  thallus  in  the  simplest  forms  is  unicellular;  in  the  majority, 
however,  it  is  built  up  of  many  cells,  which  in  a  few  instances 
are  exactly  similar ;  but  generally  there  is  a  division  of  labour,  so 
that  certain  cells  undertake  certain  functions  and  are  constructed 
accordingly,  while  others  have  different  work  and  corresponding 
structure.  Vessels  or  similar  high  anatomical  structures  are 
seldom  formed,  and  the  markings  on  the  cell-wall  are  with  few 
exceptions  very  simple.  The  Myxomycetes  occupy  quite  an 
isolated  position  ;  their  organs  of  nourishment  are  naked  masses 
of  protoplasm  (plasmodia). 

As  regards  the  external  form,  the  thallus  may  be  entirely 
without  special  prominences  (such  as  branches,  members),  but 
when  such  are  present  they  are  all  essentially  alike  in  their 
origin  and  growth,  that  is,  disregarding  the  hair- structures 
which  may  be  developed.  A  shoot  of  a  Seaweed  or  of  a  Lichen, 
etc.,  is  essentially  the  same  as  any  other  part  of  the  plant ;  only 
among  the  highest  Algae  (Characeae,  certain  Siphonese,  Sar- 
gassum,  and  certain  Ked  Seaweeds)  do  we  find  the  same  differ- 
ences between  the  various  external  organs  of  the  plant  body  as 
between  stem  and  leaf,  so  that  they  must  be  distinguished  by 
these  names. 

Roots  of  the  same  structure  and  development  as  in  the  Seed- 
plants  are  not  found,  but  organs  of  attachment  (rhizoids  and 
haptera)  serve  partly  the  biological  functions  of  the  root. 

SYSTEMATIC  DIVISION  OF  THE  THALLOPHYTES.  To  the  Thallophytes 
belong  three  sub-divisions — Slime-Fungi,  Algae,  and  Fungi. 
Formerly  the  Thallophytes  were  divided  into  Algae,  Fungi,  and 
Lichens.  But  this  last  group  must  be  placed  among  the  Fungi, 
since  they  are  really  Fungi,  which  live  symbiotically  with  Algae. 
The  Slime-Fungi  must  be  separated  from  the  true  Fungi  as  a 
distinct  sub-division.  The  Algce  possess  a  colouring  substance, 
which  is  generally  green,  brown,  or  red,  and  by  means  of  which, 
they  are  able  to  build  up  organic  compounds  from  carbonic  acid 
and  water.  The  Bacteria,  especially,  form  an  exception  to  the 
Algas  in  this  respect ;  like  the  Fungi  and  Slime-Fungi  they  have 
as  a  rule  no  such  colouring  material,  but  must  have  organic  car- 
bonaceous food  ;  these  plants  form  no  starch,  and  need  no  light 


MTXOMYCETES.  5 

for  their  vegetation  (most  Fungi  require  light  for  fructification). 
The  Myxomycetes,  Bacteria,  and  Fungi  derive  their  nourish- 
ment either  as  saprophytes  from  dead  animal  or  vegetable  matter, 
or  as  parasites  from  living  animals  or  plants  (hosts),  in  which  they 
very  often  cause  disease. 

A  remark,  however,  must  be  made  with  regard  to  this  division.  Among  the 
higher  plants  so  much  stress  is  not  laid  upon  the  biological  relations  as  to  divide 
them  into  "  green  "  and  "  non-green";  Cuscuta  (Dodder),  a  parasite,  is  placed 
among  the  Convolvulaceae,  Neottia  and  Corallorhiza,  saprophytes,  belong  to  the 
Orchidaceas,  although  they  live  like  Fungi,  yet  their  relations  live  as  Algae.  In 
the  same  manner  there  are  some  colourless  parasitic  or  saprophytic  forms  among 
the  Algae,  and  stress  must  be  laid  upon  the  fact  that  not  only  the  Blue-green  Algae, 
but  also  the  Bacteria,  which  cannot  assimilate  carbonic-acid,  belong  to  the  Algae 
group,  Sehizophyceae.  The  reason  for  this  is  that  systematic  classifications  must 
be  based  upon  the  relationship  of  form,  development,  and  reproduction,  and  from 
this  point  of  view  we  must  regard  the  Bacteria  as  being  the  nearer  relatives  of 
the  Blue-green  Algae.  All  the  Thallophytes,  which  are  designated  Fungi  (when 
the  entire  group  of  Slime-Fungi  is  left  out),  form  in  some  measure  a  connected 
series  of  development  which  only  in  the  lower  forms  (Phycomycetes)  is  related 
to  the  Alga?,  and  probably  through  them  has  taken  its  origin  from  the  Alg83  ;  the 
higher  Fungi  have  then  developed  independently  from  this  beginning.  The 
distinction  of  colour  referred  to  is  therefore  not  the  only  one  which  separates  the 
Alga?  from  the  Fungi,  but  it  is  almost  the  only  characteristic  mark  by  which 
we  can  at  once  distinguish  the  two  great  sub-divisions  of  the  Thallophytes. 

The  first  fo>ms  of  life  on  earth  were  probably  "  Protistae,"  which  had  assimi- 
lating colour  material,  or  in  other  words,  they  were  Algae  because  they  could 
assimilate  purely  inorganic  food  substances,  and  there  are  some  among  these 
which  belong  to  the  simplest  forms  of  all  plants.  Fungi  and  Slime-Fungi  must 
have  appeared  later,  because  they  are  dependent  on  other  plants  which  assimi- 
late carbon.1 

Sri-Division  I.— MYXOMYCETES,  SLIME-FUNGI. 

The  Slime-Fungi  occupy  quite  an  isolated  position  in  the 
Vegetable  Kingdom,  and  are  perhaps  the  most  nearly  related  to 
the  group  of  Rhizopods  in  the  Animal  Kingdom.  They  live  in  and 
on  organic  remains,  especially  rotten  wood  or  leaves,  etc.,  on  the 
surface  of  which  their  sporangia  may  be  found. 

They  are  organisms  without  chlorophyll,  and  in  their  vegetative 
condition  are  masses  of  protoplasm  without  cell- wall  (plasmodia) . 
They  multiply  by  means  of  spores,  which  in  the  true  Slime-Fungi 2 

1  According  to  the  recent  investigations  of  Winogradsky  some  micro-organisms 
(Nitrifying-bacteria)  can  build  organic  from  inorganic  matter.    Sachs'  hypothesis 
that  the  first  organisms  must  necessarily  have  contained  chlorophyll  is  there- 
fore untenable. 

2  Myxogasteres,  Engler's  Syllabus,  p.  1. 


6 


MYXOMYCETES. 


are  produced  in  sporangia,  but  in  some  others  l  free.  The  spores 
are  round  cells  (Fig.  1  a)  which  in  all  the  true  Slime-Fungi  are 
surrounded  by  a  cell- wall.  The  wall  bursts  on  germination,  arid 
the  contents  float  out  in  the  water  which  is  necessary  for 
germination.  They  move  about  with  swimming  and  hopping 
motions  like  swarmspores  (e, /),  having  a  cilia  at  the  front  end  and 
provided  with  a  cell-nucleus  and  a  pulsating  vacuole.  Later  on 


FiG.l.— a-l  Development  of  "Fuligo"  from  spore  to  Myxamceba ;  a-m  are  magnified 
390  times  ;  m  is  a  Myxamoeba.  of  Lycogala  epidendron  ;  I'  three  Myxamosbse  of  Physarum 
album  about  to  unite;  o,  a  small  portion  of  plasmodium,  magnified  90  times. 

they  become  a  little  less  active,  and  creep  about  more  slowly,  while 
they  continue  to  alter  their  form,  shooting  out  arms  in  various 
places  and  drawing  them  in  again  (g,  h,  i,  k,  I,  tn)  ;  in  this  stage 
they  are  called  Myzamoebce. 

The  Myxamceba  grows  whilst  taking  up  nourishment  from  the 
material  in  which  it  lives,  and  multiplies  by  division.     At  a  later 
stage  a  larger  or  smaller  number  of  A1yxamceba3  may  be  seen  to 
1  Acrasieae  and  Plasmodiophorales,  ibid. 


MYXOMYCETES. 


coalesce  and  form  large  masses  of  protoplasm,  plasmodia,  which 
in  the  "  Flowers  of  Tan "  may  attain  the  size  of  the  palm  of  a 
hand,  or  even  larger,  but  in  most  others  are  smaller.  The  plas- 
modia are  independent,  cream-like  masses  of  protoplasm,  often 
containing  grains  of  carbonate  of  lime  and  colouring  matter  (the 
latter  yellow  in  the  Flowers  of  Tan). 
They  creep  about  in  the  decaying  matter 
in  which  they  live,  by  means  of  amoeboid 
movements,  internal  streamings  of  the  pro- 


FIG.  3.— Four  sporangia  of 
Stemonitis  J'usca,  fixed  on  a 
branch,  a  The  plasmodium. 


FIG.  2.— The  plasmodinm  (a)  of  Stemoni'is  fusca,  com- 
mencing to  form  into  sporangia  (b)  ;  drawn  on  July  9. 
The  dark-brown  sporangia  were  completely  formed  by 
the  next  morning ;  c-e  shows  the  development  of  their 
external  form. 

toplasm  continually  taking  place ;  finally 
they  creep  out  to  the  surface,  and  very 
often  attach  themselves  to  other  objects, 
such  as  Mosses,  and  form  sporangia  (Fig. 
2).  These  are  stalked  or  sessile  and  are 
generally  cylindrical  (Fig.  3),  spherical 
or  pear-shaped  (Fig.  4) ;  they  rarely  attain 
a  larger  size  than  that  of  a  pin's  head,  and 
are  red,  brown,  white,  blue,  yellow,  etc.,  with  a  very  delicate  wall. 
In  some  genera  maybe  found  a  "  Capillitium  "  (Fig.  4  cp),  or  net- 
work of  branched  fine  strands  between  the  spores.  Flowers  of 


FIG.  4. — Sporangium  of  Av- 
cyria  incarnata.  J5  closed;  C 
open ;  j>  wall  of  sporangium ; 
cp  capilitium. 


8  ALG2E. 

Tan  (Fuligo  septica)  has  a  fruit-body  composed  of  many  sporangia 
(an  JEthalium),  which  has  the  appearance  of  flat,  irregular,  brown 
cakes,  inside  the  fragile  external  layer  of  which  a  loose  powder, 
the  spores,  is  found.  It  generally  occurs  on  heaps  of  tanners'  bark, 
and  appears  sometimes  in  hot-beds  in  which  that  material  is  used, 
and  is  destructive  by  spreading  itself  over  the  young  plants  and 
choking  them. 

All  the  motile  stages  may  pass  into  resting  stages,  the  small  forms 
only  surrounding  themselves  with  a  wall,  but  the  large  ones  at  the 
same  time  divide  in  addition  into  polyhedral  cells.  When  favour- 
able conditions  arise,  the  walls  dissolve  and  the  whole  appears 
again  as  a  naked  (free-moving)  mass  of  protoplasm. 

To  the  genuine  Slime-Fungi  belong :  Arcyria,  Trichia,  Didymium, 
Physarum,  Stemonitis,  Lycogala,  Fidigo,  Spumaria,  Eeticularia. 

Some  genera  wanting  a  sporangium-wall  belong  to  the  Slime- 
Fungi  :  Ceratiomyxa,  whose  fruit-body  consists  of  polygonal 
plates,  each  bearing  stalked  spores;  Uictyostelium,  in  which  the 
swarm-stage  is  wanting  and  which  has  stalked  spores.  Plas- 
modiophora  brassicce  preys  upon  the  roots  of  cabbages  and  other 
cruciferous  plants,  causing  large  swellings.  PI.  alni  causes 
coral-shaped  outgrowths  on  the  roots  of  the  Alder  (Alnus). 
Phytomyxa  leguminosarum  may  be  found  in  small  knobs  (tubercles) 
on  the  roots  of  leguminous  plants.  It  is  still  uncertain  whether 
it  is  this  Fungus  or  Bacteria  which  is  the  cause  of  the  formation 
of  these  tubercles. 


Sub-Division  II. — ALGXE. 

Mode  of  Life.  The  Algae  (except  most  of  the  Bacteria)  are 
themselves  able  to  form  their  organic  material  by  the  splitting 
up  of  the  carbonic  acid  contained  in  the  water,  or  air  in  some 
cases,  and  for  this  purpose  need  light.  The  majority  live  in 
water,  fresh  or  salt,  but  many  are  present  on  damp  soil,  stones, 
bark  of  trees,  etc. 

With  the  exception  of  the  Bacteria,  no  saprophytes  have  actually 
been  determined  to  belong  to  this  group,  and  only  very  few 
true  parasites  (for  instance,  Phyllosiphon  arisari,  Mycoidea,  etc.), 
but  a  good  many  are  found  epiphytic  or  endophytic  on  other  Alga?, 
or  water  plants,  and  on  animals  (for  instance,  certain  tichizopliycecv 
and  Protococcoidece ;  Trichophilus  welckeri  in  the  hairs  of  Bradypus? 
the  Sloth),  and  several  species  in  symbiotic  relation  to  various 


ALG.E.  9 

Fungi  (species  of  Lichen),  to  Sponges  (e.g.  Trentepohlia  spongio- 
phila,  Struvea  deiicatula),  and  to  sundry  Infusoria  and  other 
lower  animals  as  Radiolarias,  Hydra,  etc.  (the  so-called  Zoochlorella 
and  Zooxantella,  which  are  perhaps  partly  stages  in  development 
of  various  Green  and  Brown  Algae). 

Vegetative  Organs.  The  cells  in  all  the  Algae  (excepting 
certain  reproductive  cells)  are  surrounded  by  a  membrane  which 
(with  the  exception  of  the  Bacteria)  consists  of  pure  or  altered 
cellulose,  sometimes  forming  a  gelatinous  covering,  at  other  times 
a  harder  one,  with  deposits  of  chalk  or  silica  formed  in  it.  The 
cell-nucleus,  which  in  the  Schizophyta  is  less  differentiated,  may 
be  one  or  more  (e.g.  Hydrodicton,  Siphonece)  in  each  cell.  Except- 
ing in  the  majority  of  the  Bacteria,  colour  materials  (of  which 
chlorophyll,  or  modifications  of  it,  always  seems  to  be  found)  occur, 
which  either  permeate  the  whole  cytoplasm  surrounding  the  cell- 
nucleus,  as  in  most  of  the  coloured  Schizophyta,  or  are  con- 
tained in  certain  specially  formed  small  portions  of  protoplasm 
(chromatophores). 

The  individual  at  a  certain  stage  of  development  consists  nearly 
always  of  only  one  cell ;  by  its  division  multicellular  individuals 
may  arise,  or,  if  the  daughter-cells  separate  immediately  after  the 
division,  as  in  many  of  the  simplest  forms,  the  individual  will, 
during  the  whole  course  of  its  existence,  consist  of  only  a  single 
cell  (unicellular  Algae).  In  multicellular  individuals  the  cells 
may  be  more  or  less  firmly  connected,  and  all  the  cells  of  the 
individual  may  be  exactly  alike,  or  a  division  of  labour  may  take 
place,  so  that  certain  cells  undertake  certain  functions,  and  are 
constructed  accordingly  ;  this  may  also  occur  in  parts  of  the  cell 
in  the  large  unicellular  and  multinuclear  Algae  (Siphoneae,  p.  62). 

The  cells  in  most  of  the  Algae  belong  to  the  parencliymatous  form  ; 
these,  however,  in  the  course  of  their  growth,  may  very  often  become 
somewhat  oblong;  in  many  Algae  (particularly  Fucoideae  and 
Florideae)  occur,  moreover,  hyphce-like  threads,  which  are  very  long, 
often  branched,  and  are  either  formed  of  a  single  cell,  or,  more 
frequentty,  of  a  row  of  cells,  having  a  well-pronounced  apical 
growth.  The  parenchymatous  as  well  as  the  hyphae-like  cells 
may,  in  the  higher  Algae  (especially  in  certain  Fucoideae  and 
Florideae),  be  further  differentiated,  so  that  they  form  well- 
defined  anatomico-physiological  systems  of  tissue,  i.e.  assimilating, 
conducting,  storing,  and  mechanical. 

With  regard  to  the  external  form,  the  thallus  may  present  no 


10  ALGJ). 

differentiation,  as  in  many  unicellular  Algae,  or  in  multicellular 
Algae  of  the  lower  order,  which  are  then  either  equally  developed 
in  all  directions  (e.g.  Pleurococcus,  Fig.  47),  or  form  flat  cell-plates 
(Merismopediutn)  or  threads  (Oscillaria,  Fig.  21).  The  first  step  in 
the  way  of  differentiation  appears  as  a  difference  between  apex 
and  base  (Rivularia,  Porphyra)  ;  but  the  division  of  labour  may 
proceed  so  that  differences  may  arise  between  vegetative  and 
reproductive  cells  (CEdogonium,  Fig.  54) ;  hairs  and  organs  of 
attachment  (rhizoids  and  haptera),  which  biologically  serve  as 
roots,  are  developed,  and  even  leaves  in  certain  forms  cf  high 
order,  belonging  to  different  classes  (e.g.  Caulerpa,  Fig.  59  ; 
Characece,  Fig.  61  ;  Sargassum,  Fig.  72  ;  and  many  Florideae). 

The  nonsexual  reproduction  takes  place  vegetatively,  in 
many  instances,  simply  by  division  into  two,  and  more  or  less  com- 
plete separation  of  the  divisional  products  (Diatomaceae,  Desmi- 
diaceae  (Fig.  36),  many  Fission-plants,  etc.),  or  by  detached  portions 
of  the  thallus  (e.g.  Caulerpa,  Ulva  lactuca,  etc. ;  among  many  Schizo- 
phycese,  small  filaments  known  as  Jiortnogonia  are  set  free),  or 
asexnally  by  special  reproductive  cells  (spores}  set  free  from  the 
thallus  ;  these  may  be  either  stationary  or  motile.  The  stationary 
reproductive  cells  (spores)  may  either  be  devoid  of  cell- wall  (te- 
traspores  of  the  Florideae),  or  may  possess  a  cell-wall ;  in  the  latter 
case  they  may  be  formed  directly  from  the  vegetative  cells,  gene- 
rally by  the  thickening  of  the  walls  (akinetes),  or  only  after  a 
process  of  re-juvenescence  (aplanospores) .  Aplanospores,  as  well 
as  akinetes,  may  either  germinate  immediately  or  may  become 
resting-cells,  which  germinate  only  after  a  period  of  rest. 

THE  MOTILE  ASKXUAL  REPRODUCTIVE  CELLS  are  spherical,  egg-  or 
pear-shaped,  naked,  swarmspores  (zoospores),  which  have  arisen  in 
other  cells  (zoosporangia),  and  propel  themselves  through  the 
water  by  means  of  cilia ;  or  they  are  Phyto-Amoebce,  which  have  no 
cilia  and  creep  on  a  substratum  by  means  of  pseudopodia.  The 
cilia,  which  are  formed  from  the  protoplasm  (in  the  Bacteria, 
however,  from  the  membrane),  are  mostly  situated  at  the  pointed 
and  colourless  end,  which  is  directed  forwards  when  in  motion, 
and  are  1,  2  (Fig.  5  B),  4  or  more.  Both  the  cilia  in  the  Brown 
AlgJB  are  attached  to  one  side  (Fig.  65);  they  are  occasionally 
situated  in  a  circle  round  the  front  end  (CEdogonium,  Fig.  6  a, 
and  Derbesia),  or  are  very  numerous  and  situated  in  pairs  dis- 
tributed over  a  large  part  or  nearly  ihe  whole  of  the  zoospore 
( Vaucheria) .  Besides  being  provided  with  one  or  more  nuclei 


ALGJ:. 


11 


(Vaucheria),  they  may  also  have  a  red  "eye  spot"  and  vacu- 
oles,  which  are  sometimes  pulsating,  i.e.  they  appear  and  re- 
appear at  certain  intervals.  The  swarmspores  move  about  in 
the  water  in  irregular  paths,  and  apparently  quite  voluntarily, 
revolving  round  their  longer  axes  ;  but  they  come  to  the  sur- 
face of  the  water  in  great  numbers  either  because  of  their 
dependence  on  light,  or  driven  by  warm  currents  in  the 
water,  or  attracted  by  some  passing  mass  of  food  material. 
The  swarmspores  germinate,  each  forming  a  new  plant,  as  their 
movement  ceases  they  surround  themselves  with  a  cell-wall, 
grow,  and  then  divide  ;  in  Fig.  6  6,  two  may  be  seen  in  the  con- 
dition of  germination,  and  about  to  attach  themselves  by  means 
of  the  front  end,  which  has  been  developed  into  haptera  (see  also 
Fig.  5  I?,  lowest  figure). 


FIG.  6.— Cladophora  glomcrata.    A  The  lower  cells  are  FIG.   6.  —  (Edogonium  -.    a 
full  of  swarmspores,  whilst  from  the   upper  oue  the  (free),  b  germinating  swarm- 
greater  part  have  escaped   through  the   aperture  m.  spores. 
B  Free  and  germinating  swarmspores. 

The  sexual  reproduction  here,  probably  in  all  cases,  con- 
sists in  the  coalescence  of  two  masses  of  protoplasm,  that  is,  in 
the  fusion  of  their  nuclei. 

The  simplest  and  lowest  form  is  termed  conjugation,  or  isoga- 
mous  fertilisation,  and  is  characterized  by  the  fact  that  the  two 
coalescing  cells  (termed  gametes)  are  equal,  or  almost  equal,  in  shape 
and  size  (the  female  gamete  in  the  Cutleriacece,  e.g.  Zanardinia 


12 


collaris,  Fig.  7,  is  considerably  larger  than  the  male  gamete).  The 
cell  in  which  the  gametes  are  developed  is  called  a  gametangium, 
and  the  reproductive  cell  formed  by  their  union — which  generally 

has  a  thick  wall  and  only 
germinates  after  a  short 
period  of  rest — is  termed  a 
zygote  or  zygnspore.  The  con- 
jugation takes  place  in  two 
ways  : — 

(a)  In  the  one  way  the 
gametes  are  motile  cells 
( planogametes,  zoogametes, 
Fig.  8),  which  unite  in 
pairs  during  their  swarming 
hither  and  thither  in  the 
water ;  during  this  process 
they  lie  side  by  side  (Fig.  8d), 
generally  at  first  touching  at 
the  clear  anterior  end,  and 
after  a  time  they  coalesce 
and  become  a  motionless  zy- 
gote, which  surrounds  itself 
with  a  cell-wall  (Fig.  8  e). 
This  form  of  conjugation  is  found  in  Ulothrix  (Fig.  8  d),  Acetdbu- 
laria,  and  other  Algae  (Figs.  45,  56,  66). 

(6)  Among  other  Algae  (e.g.  Diatomacece  and  Conjugatce),  the 
conjugating  cells  continue  to  be  surrounded  by  the  cell -wall  of 
the  mother-cell  (aplanogametes  in  an  aplanogametangiuin) ;  the 


FIG.  7. — Zanardinia  collaris.  A  Male  game- 
tangia  (the  small-celled)  and  female  gametangia 
(large-celled).  C  Female  gamete.  D Male  gamete. 
B  E  Fertilisation.  F  Zygote.  G  Germinating 
zygote. 


FIG.  8. — Ulothrix  zonata :  a  portion  of  a  thread  with  zoospores,  of  which  two  are  formed 
in  each  cell  (zoosporangium),  the  dark  spots  upon  them  are  the  "red  eye-spots  "  ;  1,  2,  3, 
4  depict  successive  stages  in  the  development  of  the  zoospores ;  b  a  single  zoospore,  at  v 
the  pulsating  vacuole ;  c  portion  of  a  thread  with  gametes,  of  which  sixteen  are  formed  in 
each  gametangium ;  d  gametes  free  and  in  conjugation;  e  conjugation  has  been  effected, 
and  the  formed  zygotes  are  in  the  resting  condition. 


ALG.E. 


13 


aplanogametangia  generally  grow  out  into  short  branches,  which 
lie  close  together  and  touch  one  another,  the  wall  at  the  point 
of  contact  is  then  dissolved  (Fig.  39).  Through  the  aperture  thus 
formed,  the  aplanogametes  unite,  as  in  the  first  instance,  and 
form  a  rounded  zygote,  which  immediately  surrounds  itself  with 
a  cell- wall.  Various  modifications  occur ;  compare  Figs.  37,  39, 
41,  43. 

The  highest  form  of  the  sexual  reproduction  is  the  Egg-  or 
Oogamous  fertilisation.  The  two  coalescing  cells  are  in  the  main 
unlike  each  other  in  form  as  well  as  size.  The  one  which  is  con- 
sidered as  the  male,  and  is  known  as  the  spermatozoid  (anther  ozoid), 
developes  as  a  rule  in  large  numbers  in  each  mother-cell  (antheri- 


if       ) 

FIG.  9.— Fertilisation  in  the  Bladder- 
wrack  (Fitcus  vesiculosus). 

dium);  they  are  often  self- 
motile  (except  in  the  Flori- 
deae,  where  they  are  named 
spermatia) ,  and  are  many 
times  smaller  than  the  other 
kind,  the  female,  which  is 
known  as  the  egg-cell 
(oosphere).  The  egg-cell  is 
always  a  motionless,  spheri-  FlG' 10— SP''««>PI«*  a«n«iin«. 

cal,  primordial  cell  which  can  either  float  about  freely  in  the 
water,  as  in  the  Fucaceee  (Fig.  9),  or  is  surrounded  by  a  cell-wall 
(oogonium)',  generally  only  one  oosphere  is  to  be  found  in  each 
oogonium,  but  several  occur  in  Sphceroplea  (Fig.  10).  The  re- 


14 

suit  of  the  spermatozoid  coalescing  with  the  egg-cell  is,  as  in 
the  preceding  case,  the  formation  of  a  zygote,  which  generally 
undergoes  a  period  of  rest  before  germination  (the  Florideae  are 
an  exception,  a  fruit-body,  cystocarp,  being  produced  as  the  result 
of  coalescence). 

An  example  of  fertilisation  is  afforded  by  the  Alga,  Spharoplea  annulina  (Fig. 
10).  The  filamentous  thallus  is  formed  of  cylindrical  cells  with  many  vacuoles  (r 
in  A) ;  some  cells  develope  egg-cells  (B),  others  spermatozoids  ((7),  the  latter  in 
a  particularly  large  number.  The  egg-cells  are  spherical,  the  spermatozoids  of 
a  club-  or  elongated  pear-shape  with  two  cilia  at  the  front  end  (G  ;  E  is  however 
a  swarmspore).  The  spermatozoids  escape  from  their  cells  through  apertures  in 
the  wall  (o  in  C)  and  enter  through  similar  apertures  (o  in  B)  to  the  egg-cells. 
The  colourless  front  end  of  the  spermatozoid  is  united  at  first  with  the  "re- 
ceptive spot"  of  the  egg-cell  (see  F),  and  afterwards  completely  coalesces  with 
it.  The  result  is  the  formation  of  a  zygote  with  wart-like  excrescences  (D). 

The  female  (parthenogenesis)  or  male  (androgenesis)  sexual  cell 
may,  sometimes  without  any  preceding  fertilisation,  form  a  new 
individual  (e.g.  Ulothrix  zonata,  Cylin  drocapsa,  etc.). 

Systematic  division  of  the  Algae.  The  Algae  are  divided 
into  the  following  ten  classes  : 

1.  SYNGENETICLE  ;  2.  DINOFLAGELLATA,  or  PERIDINEA  ;  3.  DIATO- 
MACE^l  ;  4.  SCHIZOPHYTA,  FlSSION-ALGJ; ;  5.  CONJUGATE;  6.  CHLORO- 
PHYCE^,  GREEN- ALGJS;  7.  CHARACEJ;,  STONEWORTS;  8.  PILEOPHYCEJ:  ; 
9.  DlCTYOTALES;  10.  RHODOPHYCE^. 

Among  the  lowest  forms  of  the  Algae,  the  Syngeneticae,  the 
Dinoflagellata,  and  the  unicellular  Volvocaceae  (Chlamydomoneae), 
distinct  transitional  forms  are  found  approaching  the  animal 
kingdom,  which  can  be  grouped  as  animals  or  plants  according  to 
their  method  of  taking  food  or  other  characteristics.  Only  an 
artificial  boundary  can  therefore  be  drawn  between  the  animal 
and  vegetable  kingdoms.  In  the  following  pages  only  those  forms 
which  possess  chromatophores,  and  have  no  mouth,  will  be  con- 
sidered as  Algae. 


Class  l.    Syngeneticae. 

The  individuals  are  uni-  or  multi-cellular,  free-swimming  or 
motionless.  The  cells  (which  in  the  multicellular  forms  are  loosely 
connected  together,  often  only  by  mucilaginous  envelopes)  are  naked 
or  surrounded  by  a  mucilaginous  cell- wall,  in  which  silica  is  never 
embedded.  They  contain  one  cell-nucleus,  one  or  more  pulsating 


SYNGENETIOE. 


15 


vacuoles,  and  one  to  two  band-  or  plate-like  chromatophores  with 
a  brown  or  yellow  colour,  and  sometimes  a  pyrenoid. 

Reproduction  takes  place  by  vegetative  division,  or  asexually  by 
zoospores,  akinetes  (or  aplanospores  ?).  Sexual  reproduction  is  un- 
known. They  are  all  fresh  water  forms. 

To   this  class  may  perhaps   be   assigned  the  recently   arranged   and  very 
little    known     orders    of     Calcocytaceec,     Murracytacece,    Xanthellacece,    and 
Dictyochacea,  which  partly  occur  in  the 
free  condition  in  the  sea,  in  the  so-called 
"  Plankton,"    and     partly    symbiotic    in 
various  lower  marine  animals. 

The  Syngeneticce  are  closely  re- 
lated to  certain  forms  in  the  animal 
kingdom,  as  the  Flagellatse. 

Order  1.  Chrysomonadinaceae.  In- 
dividuals, uni-  or  multi-cellular,  swim- 
ming in  free  condition,  naked  or  sur- 
rounded by  a  mucilaginous  covering.  The 
cells  are  generally  oval  or  elongated,  with 
2  (rarely  only  1)  cilia,  almost  of  the  same 
length,  and  generally  with  a  red  "  eye-spot 
only)  band-shaped  chromatophores.  Keproduction  by  the  longitudinal  divi- 
sion of  the  individual  cells  either  during  the  swarming,  or  during  a  resting 
stage ;  in  the  multicellular  forms  also  by  the  liberation  of  one  or  more  cells, 
which  in  the  latter  case  are  connected  together. 

A.  Unicellular :  Chromulina,  Cryptoglena,  Microglena,  Nephroselmis. 

B.  Multicellular  :  Uroglena,  Syncrypta  (Fig.  11),  Synura. 

Among  the  unicellular  Chrysomonadinaceae  are  probably  classed  some  forms 
which  are  only  stages  in  the  development  of  the  multicellular,  or  of  other 
Syiif/eneticce. 

Order  2.  Chrysopyxaceae  are  unicellular,  and  differ  mainly  from  the  pre- 
ceding in  being  attached  either  on  a  slime-thread  (Stylo- 
chrysalis),  or  enclosed  in  an  envelope  (Chrysopyxis,  Fig. 
12).  They  have  two  cilia,  and  multiply  by  longitudinal 
(Chrysopyxis)  or  transverse  division,  and  the  swarming 
of  one  of  the  daughter-individuals  (zoospore).  Division 
may  also  take  place  in  a  motionless  stage  (palmella-stage). 

Order  3.  Dinobryinaceae.  The  individuals  are  ori- 
ginally attached,  uni-  or  multi-cellular  ;  each  individual 
cell  is  distinctly  contractile,  and  fixed  at  the  bottom  of  a 
cup-shaped,  open  envelope.  Cilia  2,  but  of  unequal 
length.  Asexual  reproduction  by  zoospores,  which  are 
formed  by  straight  or  oblique  longitudinal  division  of 
the  mother-cell,  during  a  palmella-stage  which  is  pro- 
duced in  the  winter  aplanospores.  Epipyxis,  Dinobryon. 


FIG.  11. — Syncrypta  volvox :  the  multi- 
cellular  individual  is  surrounded  by  a 
mucilaginous  granular  envelope. 

'  at  their  base,  and  with  2  (rarely  1 


Fio.  12.— Chrysopyxis 
bipes  :  m  envelope,  EC 
chromatophure,  cv 
contractile  vacuole. 


16  DINOFLAGELLATA. 

Order  4.  Hydruraceae.  The  individuals  are  attached,  without  cilia,  multi- 
cellular,  branched,  and  with  apical  growth.  The  cells  are  spherical,  but  in 
the  final  stage  almost  spindle-shaped,  and  embedded  in  large  masses  of  mucilage. 
Asexual  reproduction  by  zoospores  which  are  tetrahedric,  with  1  cilia,  and  by 
resting  akinetes.  Hydrurus  is  most  common  in  mountain  brooks. 


Class  2.    Dinoflagellata. 

The  individuals  are  of  a  very  variable  form,  but  always  uni- 
cellular, and  floating  about  in  free  condition.  The  cell  is  dorsi- 
ventral,  bilateral,  asymmetric  and  generally  surrounded  by  a  colour- 
less membrane,  which  has  no  silica  embedded  in  it,  but  is  formed 
of  a  substance  allied  to  cellulose.  The  membrane,  which  exter- 
nally is  provided  with  pores  and  raised  borders,  easily  breaks  up  into 
irregularly-shaped  pieces.  In  the  forms  which  have  longitudinal  and 
cross  furrows,  two  cilia  are  fixed  where  these  cross  each  other,  and 
project  through  a  cleft  in  the  membrane  ;  one  of  these  cilia  projects 
freely  and  is  directed  longitudinally  to  the  front  or  to  the  rear,  the 
other  one  stretches  crosswise  and  lies  close  to  the  cell,  often  in  a 
furrow  (cross  furrow).  The  chromatophores  are  coloured  brown 
or  green  and  may  either  be  two  parallel  (Exuviella\  or  several 
radially  placed,  discs,  which  sometimes  may  coalesce  and  become  a 
star-shaped  chromatophore.  The  colouring  material  (pyrrophyl) 
consists,  in  addition  to  a  modification  of  chlorophyl,  also  of  phyco- 
pyrrin  and  peridinin  ;  this  colour  is  sometimes  more  or  less  masked 
by  the  products  of  assimilation  which  consist  of  yellow,  red  or 
colourless  oil  (?)  and  starch.  Cell-nucleus  one :  in  Polydinida  several 
nuclei  are  found ;  contractile  vacuoles  many,  which  partly  open  in 
the  cilia-cleft  (Fig.  13  grs).  In  some  an  eye-spot,  coloured  red  by 
haematochrome,  is  found.  Pyrenoids  occur  perhaps  in  Exuviella 
and  Amphidinium. 

THE  REPRODUCTION  takes  place  as  far  as  is  known  at  present, 
only  by  division.  This,  in  many  salt  water  forms,  may  take 
place  in  the  swarming  condition,  and,  in  that  case,  is  always 
parallel  to  the  longitudinal  axis.  The  daughter-individuals,  each 
of  which  retains  half  of  the  original  shell,  sometimes  do  not  separ- 
ate at  once  from  each  other,  and  thus  chains  (e.g.  in  Ceratium)  of 
several  connected  individuals  may  be  formed.  In  others,  the 
division  occurs  after  the  cilia  have  been  thrown  off  and  the  cell-con- 
tents rounded.  The  daughter-cells  then  adopt  entirely  new  cell-walls. 
A  palmella-stage  (motionless  division-stage)  sometimes  appears  to 


DINOFLAGELLATA. 


17 


take  place,  and  also  aplanospores  (?)  with  one  or  two  horn-like 
elongations  (e.g.  in  Peridiniwm  cinctum  and  P.  tabulatum) ;  at 
germination  one,  or  after  division,  two  or  more,  new  individuals 
may  be  formed. 

Sexual  reproduction  has  not  been  observed  with  certainty. 

The  Dinoflagellata  move  forward  or  backward,  turning  round 
their  longitudinal  axes ;  in  their  motion  they  are  influenced  by 
the  action  of  light.  The  motion  possibly  may  be  produced 
only  by  the  transverse  cilium,  which  vibrates  rapidly ;  whilst  the 
longitudinal  cilium  moves  slowly,  and  is  supposed  to  serve  mainly 
as  a  steering  apparatus.  They  live  principally  in  salt  water,  but 
also  in  fresh. 

Besides  the  coloured  forms,  which 
are  able  to  make  their  own  organic 
compounds  by  the  splitting  up  of  the 
carbonic  acid  contained  in  the  water, 
there  are  a  few  colourless  forms  (e.g. 
Gymnodinium  spirale),  or  such  as  do  not 
possess  chromatophores  (Polykrikos) ; 
these  appear  to  live  saprophytically, 
and  may  be  able  to  absorb  solid  bodies 
with  which  they  come  in  contact. 

Dinoflagellata  occur  in  the  "Plank- 
ton "  of  the  open  sea,  where  they  form 
together  with  Diatomacese  the  basis 
for  the  animal  life.  It  is  known  with 
certainty  that  some  salt  water  forms 
(like  the  Noctiluca,  which  belongs  to 
the  animal  kingdom  arid  to  which 
they  are  perhaps  related)  produce 
light,  known  as  phosphorescence. 


FIG.  13.— A  and  B  Glenodinium 
cinctum.  A  Seen  from  the  ventral 
side,  B  from  behind ;  fg  transverse 
cilium;  g  longitudinal  cilium;  ch 
chromatophores ;  a  starch  ;  n  cell- 
nucleus  ;  v  vacuole ;  oc  eye-spot ;  C 
Ceratium  tetraceros  from  the  ventral 
side;  r  the  right,  b  the  posterior 
horn ;  If  longitudinal  furrow  ;  gs 
cilium-cleft ;  v  vacuole  ;  g  longitudi- 
nal cilium.  (A  and  B  mag.  450 
times,  C  337  times.) 

Dinoflage llata  (Peridinea,  CilioJJagellata)  are  allied  through  their  lowest  form 
(Exuviella)  to  the  Syngeneticee  and  especially  to  the  order  Chrysomonadinaceas. 
They  may  be  divided  into  three  orders. 

Order  1.  Adinida.  Without  transverse  or  longitudinal  furrows,  but  en- 
closed in  two  shells,  and  with  two  parallel  chromatophores  in  each  cell. 
Exuviella,  Prorocentrum. 

Order  2.  Dinifera.  With  tranverse  and  generally  longitudinal  furrow.  Many 
radially-placed,  disc-formed  chromatophores.  The  most  common  genera  are — 
Ceratium  (Fig.  13),Peridinium,  Glenodinium  (Fig.  13),  Gymnodinium,  Dinophysis. 

Order  3.  Polydinida.  With  several  transverse  furrows,  no  chromatophores, 
and  several  cell-nuclei.  Only  one  genus — Polykrikos. 

\V.B.  C 


18  DIATOM  EJE. 

The  order  Polydinida  deviates  in  a  high  degree  from  the  other  Dinoflagellata, 
not  only  by  its  many  tranverse  furrows,  each  with  its  own  transverse  cilium,  and 
by  the  absence  of  chromatophores,  but  also  in  having  several  cell-nuclei  and  a 
Mnd  of  stinging  capsule,  which  otherwise  does  not  occur  within  the  whole 
class.  It  may  therefore  be  questionable  whether  this  order  should  really  be 
placed  in  the  vegetable  kingdom. 

Class  3.    Diatomeae. 

The  individuals — each  known  as  a  frustule — assume  very  various 
forms  and  may  be  unicellular  or  multicellular,  but  present  no 
differentiation;  many  similar  cells  may  be  connected  in  chains, 
embedded  in  mucilaginous  masses,  or  attached  to  mucilaginous 
stalks.  Tjbe  cells  are  bi-lateral  or  centric,  often  asymmetrical, 
slightly  dorsiventral  and  have  no  cilia ;  those  living  in  the  free 
condition  have  the  power  of  sliding  upon  a  firm  substratum.  The 
cell  contains  1  cell-nucleus  and  1-2  plate-shaped  or  several  disc- 
shaped  chromatophores.  The  colouring  material  "  Melinophyl " 
contains,  in  addition  to  a  modification  of  chlorophyl,  a  brown 
colouring  matter,  diatomin.  1  or  2  pyrenoids  sometimes  occur. 
Starch  is  wanting  and  the  first  product  of  assimilation  appears  to 
be  a  kind  of  oil  (?). 

The  cell-walls  are  impregnated  with  silica  to  such  a  degree  that 
they  are  imperishable  and  are  therefore  able  to  contribute  in  a 
great  measure  to  the  formation  of  the  earth's  crust.  The  structure 
of  their  cell- wall  is  most  peculiar  and  differs  from  all  other  plants 
(except  certain  Desmidiaceoe)  ;  it  does  not  consist  of  a  single  piece 
but  is  made  up  of  two — the  "  shells  " — (compare  Exuviella  and 
Prorocentrum  among  the  Dinoflagellata)  which  are  fitted  into  each 
other,  one  being  a  little  larger  than  the  other  and  embracing  its 
edge,  like  a  box  with  its  lid  (Fig.  14  B).  The  two  parts  which  cor- 
respond to  the  bottom  and  lid  of  the  box  are  known  as  valves. 
Along  the  central  line  of  the  valves  a  longitudinal  rib  may  often 
be  found,  interrupted  at  its  centre  by  a  small  cleft  (perhaps 
homologous  with  the  cilia-cleft  of  the  Dinoflagellata),  through 
which  the  protoplasm  is  enabled  to  communicate  with  the  exterior 
(Fig.  14  A).  It  is  principally  by  reason  of  the  valves,  which  bear 
numerous  fine,  transverse  ribs,  striae  or  warts,  etc.  (Figs.  14,  15,  17), 
that  the  Diatomege  have  become  so  well  known  and  employed  as 
test  objects  in  microscopical  science.  When  the  division  takes 
place,  the  two  shells  are  separated  a  little  from  each  other,  and 
after  the  cell-contents  have  divided  into  two  masses,  two  new 
shells  are  formed,  one  fitting  into  the  larger  valve,  the  other  one 


DTATOMEJ]. 


10 


into  the  smaller  valve  of  the  original  frustule.  The  latter  cell 
(frustule)  is  thus,  upon  the  whole, 
smaller  than  the  mother-cell,  and  as 
the  cells  do  not  increase  in  size,  some 
frustules  are  smaller  than  the  ones 
from  which  they  are  derived,  and  thus, 
by  repeated  divisions,  it  follows  that 
smaller  and  smaller  frustules  are  pro- 
duced. This  continued  diminution  in 
size  is,  however,  compensated  for  by 
the  formation,  when  the  cells  have 
been  reduced  to  a  certain  minimum, 
of  auxospores,  2-3  times  larger.  These 
may  either  be  formed  asexually  by  the 
protoplasm  of  a  cell  increasing,  round- 
ing off  and  surrounding  itself  with  a 
new  wall  (e.g.  Melosira)  or  after  con- 
jugation, which  may  take  place  with 
various  modifications :  1.  Two  indi- 
viduals unite  after  the  secretion  of  a 
quantity  of  mucilage,  and  the  valves 
then  commence  to  separate  from  each 
other,  on  the  side  which  the  two  indi-  FlG-  ".-Pinnuiarta .-  B,  from 

the  edge,  shows  the  valves  fitting 

viduals  turn  towards  each  other.      The     together ;  A,  a  valve. 


FIG.  15.— Various  Diatomacese.  A  Diatoma  vulgare.  B  Talellaria  flocculosa.  C  Navicul* 
tumida  (lateral  views).  D  Gomphonema  constrictum  (lateral  views).  E  Navicula  westll 
(lateral  views). 

protoplasmic  bodies  now  release  themselves  from  their  cell-wall, 
and  each  rounds  off  to  form  an  ellipsoidal  mass  ;  these  two  pro- 


20 


DIATOME.E. 


toplasmic  masses  (gametes)  coalesce  to  form  a  zygote,  the  cell- 
nuclei  and  chromatophores  also  fusing  together.  The  zygote  in- 
creases in  size,  and  surrounds  itself  with  a  firm,  smooth,  siliceous 
wall — the  perizonium.  The  auxospores,  whichever  way  they  arise, 
are  not  resting  stages.  The  germination  of  the  zygote  com- 
mences by  the  protoplasm  withdrawing  itself  slightly  from  the 
cell-wall  and  constructing  first  the  larger  valve,  and  later  on 
the  smaller  one ;  finally  the  membrane  of  the  zygote  bursts  (e.g. 
Himantidium).  2.  The  conjugation  occurs  in  a  similar  manner, 
but  the  protoplasm  of  the  cells  divides  transversely  before 
conjugation  into  two  daughter-cells.  Those  lying  opposite  one 
another  conjugate  (Fig.  16)  and  form  two  zygotes.  The  for- 
mation of  the  perizonium,  and  germination  take  place  as  in  the 
preceding  instance  (e.g.  Epithemia).  3.  Two  cells  place  them- 
selves parallel  to  each  other,  and  each  of  the  two  cell-contents, 


FIG.  16. — Conjugation  of  Cymbella  variabilis.  A,  The  protoplasm  in  the  two  cells  has 
divided  into  two  masses  ;  B  these  masses  coalesce  in  pairs;  the  cells  (B  C)  enclosed  in  a 
mucilaginous  matrix.  C  D  Auxospores  and  their  formation. 

without  coalescing,  becomes  an  auxospore.  The  formation  of 
the  wall  takes  place  as  in  the  preceding  case.  This  is  found  in 
the  Naviculea3,  Cymbelleae,  the  Gomphonemeae  (e.g.  Frustulia, 
Cocconema). 

The  Diatomacece  may  be  found  in  salt  as  well  as  in  fresh  water 
(often  in  such  masses  that  the  colour  of  the  water  or  mud  becomes 
yellow  or  brown ;  in  the  same  manner  the  genera  Chcetoceros, 
Rhizosolenia,  Coscinodiscus,  and  several  others,  form  large  slime- 
masses,  "  Plankton  "  on  the  surface  of  the  sea),  on  damp  soil  and 
in  dust  blown  by  the  wind.  They  occur  as  fossils  in  the  recent 
formations,  often  in  large  deposits  (siliceous  earth,  mountain 
meal),  as  in  the  cement  lime  in  Jutland,  the  alluvial  deposits 
beneath  Berlin,  in  clay  strata  beneath  peat  bogs,  in  guano,  etc. 


DIATOME^E.  21 

These  accumulations  of  fossilized  diatoms  are  used  in  the  manu- 
facture of  dynamite  and  in  various  manufactures. 

The  Diatomaceae  appear  nearest  to,  and  must  be  placed  as  a 
group  co-ordinate  with  the  Dinoflagellata,  as  they  doubtless  may 
be  supposed  to  derive  their  origin  from  forms  resembling  Exuviella, 
and  to  have  lost  the  cilia.  The  resemblances  to  the  Desmidiaceae 
which  are  striking  in  many  respects,  can  only  be  conceived  as 
analogies,  and  cannot  be  founded  upon  homologies,  and  it  is 
therefore  impossible  to  regard  them  as  proof  of  genetic  relation- 
ship. The  family  contains  only  one  order. 


FIG.  17.— Various  Diatomeae.  A  Synedra  radians.  B  EpitJiemia  turgida,  (from  the  two 
different  sides).  C  Cymbella  cuspidata.  D  Cocconeis  pediculus  (on  the  right  several  situated 
on  a  portion  of  a  plant,  on  the  left  a  single  one  more  highly  magnified). 

Order  1.  Diatomaceae.  This  order  may  be  divided  into  two 
sub-orders,  viz. — 

Sub-Order  1.  Placochromaticae.  The  chromatophores  are 
discoid,  large,  1  or  2  in  each  cell ;  the  structure  of  the  valves 
is  bilateral  and  always  without  reticulate  markings.  The  follow- 
ing groups  belong  to  this  sub-order :  Gomphonemece,  Cymbellecc, 
Amphorece,  Acknanthece,  Cocconeidece,  Naviculece,  Amphipleurece,  Pla- 
giotropidecr,  Amphitropidece,  Nitzchiece,  Surirayece,  and  Eunotiece. 

Sub-Order  2.  Coccochromaticae.  The  chromatophores  are 
granular,  small  and  many  in  each  cell.  The  structure  of  the  cells 
is  zygomorphic  or  centric,  often  with  reticulate  markings.  The 
following  groups  belong  to  this  sub-order  :  Fragilariece,  Meridiece, 
Tabellariece,  Licmopkorece,  BiddulpMece,  Anguliferce,  JEupodiscece, 
Coscinodiscece  and  Melosirece. 


22  SCH1ZOPHYTA. 

Class  4.    Schizophyta,  Fission-Algae. 

The  individuals  are  1 — many  celled  ;  the  thallus  consists  in 
many  of  a  single  cell,  in  others  of  chains  of  cells,  the  cells  dividing 
in  only  one  definite  direction  (Figs.  18,  21).  In  certain  Fission-Algae 
the  cell-chain  branches  (Fig.  30)  and  a  difference  between  the  an- 
terior and  the  posterior  ends  of  the  chain  is  marked ;  in  some,  the 
cells  may  be  united  into  the  form  of  flat  plates  by  the  cell-division 
taking  place  in  two  directions  ;  and  in  others  into  somewhat  cubi- 
cal masses,  or  rounded  lumps  of  a  less  decided  form,  by  the  divisions 
taking  place  in  three  directions  ;  or  less  defined  masses  may  be 
formed  by  the  divisions  taking  place  in  all  possible  directions. 

The  cell-walls  rarely  contain  cellulose,  they  often  swell  con- 
siderably (Figs.  20,  22),  and  show  distinct  stratifications,  or  they 
are  almost  completely  changed  into  a  mucilaginous  mass  in  which 
the  protoplasts  are  embedded,  e.g.  in  Nostoc  (Fig.  22),  and  in  the 
"ZooglcEa"  stage  of  the  Bacteria  (Fig.  27).  Sexual  reproduction 
is  wanting.  Vegetative  reproduction  by  division  and  the  separa- 
tion of  the  divisional  products  by  the  splitting  of  the  cell-wall  or 
its  becoming  mucilaginous  ;  among  the  Nostocaceaa,  Lyngbyaceao, 
Scytonemaceae,  etc.,  "  Hormogonia "  are  found ;  in  ChamcesipJion 
and  others  single  reproductive  akinetes  are  formed.  Many  Fission- 
AlgJB  conclude  the  growing  period  by  the  formation  of  resting 
akinetes  or  aplanospores. 

The  Schizophyta  may  be  divided  into  2  families : 

1.  SCHIZOPHYCEE. 

2.  BACTERIA. 

Family  1.     Schizophyceae,*  Blue-Green  Algae. 

All  the  Blue-green  Algas  are  able  to  assimilate  carbon  by  means 
of  a  colouring  material  containing  chlorophyll  (cyanophyll) ;  but 
the  chlorophyll  in  this  substance  is  masked  by  a  blue  (phycocyan), 
or  red  (phycoerythrin,  e.g.  in  Trichodesmium,  erythrceum  in  the  Red 
Sea)  colouring  matter  which  may  be  extracted  from  them  in  cold 
water  after  death.  The  colouring  matter,  in  most  of  them,  per- 
meates the  whole  of  the  protoplasm  (excepting  the  cell-nucleus), 
but  in  a  few  (e.g.  Qlaucocystis,  Phragmonema),  slightly  developed 
chromatophores  are  to  be  found.  Where  the  cells  are  united  into 
filaments  (cell-chains)  a  differentiation  into  apex  and  base  (Eivu- 
lariacecd)  may  take  place,  and  also  between  ordinary  vegetative 
cells  and  heterocysts;  these  latter  cannot  divide,  and  are  dis- 
*  Mykophyceae,  Cyauophyceae. 


SCHIZOPHYTA. 


23 


tinguished  from  the  ordinary  vegetative  cells  (Fig.  22  Ji)  by  their 
larger  size,  yellow  colour,  and  poverty  of  contents.  Branching 
sometimes  occurs  and  is  either  true  or  spurious. 


FIG.  16.— Microcoleus  lyngbyanus :  a  portion  of  a  filament,  the  thick  sheath  encloses  only 
one  cell-chain ;  in  one  place  a  cell  is  drawn  out  by  the  movement  of  the  cell-chain;  I  the 
cell-chain  has  divided  into  two  parts  ("  hormongonia ")  which  commence  to  separate 
from  each  other. 

The  cell-chain  in  the  spurious  branching  divides  into  two  parts, 
of  which  either  one  or  both  grow  beyond  the  place  of  division 
(Fig.  18)  and  often  out  to  both  sides  (e.g.  Scytonema\  the  divisions 
however,  always  take  place  transversely  to  the  longitudinal 
direction  of  the  cell-chain.  In  the  true  branching  a  cell  elon- 
gates in  the  direction  transverse  to  the  cell-chain,  and  the  division 
then  takes  place  nearly  at  right  angles  to  the  former  direction 
(Sirosiphoniacece). 


FIG.  19. — Cylindrospevmum  mojits :  a  resting  akinete  with  heterocyst ;  l-d  germinating 
stages  of  a  resting  akinete  ;  e  filament  with  two  heterocysts  and  the  formation  of  new 
akinetes ;  /  part  of  a  filament  with  a  heterocyst,  and  mature  resting  akinete. 

Cilia  are  wanting,  but  the  filaments  are  sometimes  self-motile 
(e.g.  hormogonia  in  Nostoc}  and  many  partly  turn  round  their  axes, 
partly  slide  forward  or  backward  (Oscillaria) . 

Reproduction  takes  place  by  spores  and  hormogonia  in  addition 


SCinZOPHYTA. 


to  simple  cell-division.  Hormogonia  are  peculiar  fragments  of  a 
cell-chain  capable  of  motion,  and  often  exhibit  a  vigorous  motion 
in  the  sheath,  until  at  last  they  escape  and  grow  into  a  new 
individual  (Fig.  18).  The  spores  are  reproductive  akinetes 
(Chamcesiphon,  etc.)  or  resting  akinetes ;  these  latter  arise  by  the 
vegetative  cells  enlarging  and  constructing  a  thick  cell- wall  (Fig. 
19  e/).  On  germination,  this  cell- wall  bursts  and  the  new  cell-chain 
elongates  in  the  same  longitudinal  direction  as  before  (Fig.  19  b  c). 
Many  {e.g.  Oscillaria)  may  however  winter  in  their  ordinary  vege- 
tative stage.  Aplanospores  are  wanting. 

The  Fission-  Algae  are  very  prevalent  in  fresh  water  and  on 
damp  soil,  less  so  in  salt  water ;  they  also  often  occur  in  water 
which  abounds  in  decaying  matter.  Some  are  found  in  warm 
springs  with  a  temperature  as  high  as  50°  C. 

The  Family  may  be  divided  into  2  sub-families : 

1.  HOMOCYSTE^;      (heterocysts     are     wanting) :      Chroococcacece, 
Lyngbyacece  and  Chamcesiphonacece. 

2.  HETEROCYSTE^     (heterocysts    present):     Nostocacece,    Rivul- 
ariacece,  Scytonemacece  and  Sirosiphoniacece. 

Order  1.  Chroococcaceae.  The  individuals  are  1 — many- 
celled,  but  all  the  cells  are  uniform, 
united  to  form  plates  or  irregular 
masses,  often  surrounded  by  a  mu- 
cilaginous cell-wall,  but  never  forming 
cell-chains.  Multiplication  by  division 
and  sometimes  by  resting  akinetes,  but 
reproductive  akinetes  are  wanting. 
Chroococcus,  Aphanocapsa,  Glceocapsa 
(Fig.  20) ,  Ccelosphcerium,  Merismopeditim, 
Glaucocystis,  Oncobyrsa,  Polycystis,  Gom- 
phosphceria. 

Order  2.        Lyngbyaceae    (Oscil- 
lariaceae).     The  cells  are  discoid  (Fig.  21),  united  to  straight  or 

spirally  twisted,  free  filaments, 
which  are  unbranched,  or  with 
spurious  branching.  The  ends 
of  the  cell-chains  are  similar. 
Heterocysts  absent.  Reproduc- 
tion by  synakinetes,  resting 
akinetes  are  wanting.  Oscillaria  (Fig.  21),  Spirulina,  Lyvgbya, 
Microcoleus,  Symploca,  Plectonema. 


FIG.  20.— Glceocapsa  atrata:  A, 
B,  C,  D,  E  various  stages  of 
development. 


FIQ.  21.— Oscillaria;  a  terminal,  b  central 
portion  of  a  filament. 


SCHIZOPHYTA.  25 

Order  3.  Chamsesiphonaceae.  The  individuals  are  1— 
many-celled,  attached,  unbranched  filaments  with  differentiation 
into  apex  and  base,  without  heterocysts.  Multiplication  by  re- 
productive akinetes;  resting  akinetes  are  wanting.  Dermocarpa, 
Clastidium,  Chamcesiphon,  Godlewskia,  Phragmonema. 

Order  4.  Nostocacese.  The  individuals  are  formed  of  mul- 
ticullular,  unbranched  filaments,  without  differentiation  into  apex 
and  base ;  heterocysts  present.  Reproduction  by  synakinetes  and 
resting  akinetes. 

Some  genera  are  not  mucilaginous,  e.g.  Cylindrospermum  (Fig. 
19).  The  cell-chains  in  others,  e.g.  Nostoc,  wind  in  between  one 


FIG.  22. — Nostoc  verrucosum.  A  The  plant  in  its  natural  size  ;  an  irregularly  folded 
jelly-like  mass.  B  One  of  the  cell-chains  enlarged,  with  its  heterocysts  (h),  embedded  in 
its  mucilaginous  sheath. 

another  and  are  embedded  in  large  structureless  jelly-like  masses, 
which  may  attain  the  size  of  a  plum  or  even  larger  (Fig.  22) ; 
sometimes  they  are  found  floating  in  the  water,  sometimes 
attached  to  other  bodies.  Other  genera  as  follows :  Aphanizo- 
menon  and  AnabcBna  (in  lakes  and  smaller  pieces  of  water) ; 
Nodularia  is  partly  pelagic.  Some  occur  in  the  intercellular 
spaces  of  higher  plants,  thus  Nostoc-forms  are  found  in  Anthoceros, 
Blasia,  Sphagnum,  Lemna,  and  in  the  roots  of  Cycas  and  Gunnera  ; 
Anabcena  in  Azolla. 

Order  5.  Rivulariaceae.  The  individuals  are  multicellular 
filaments,  with  differentiation  into  apex  and  base ;  spurious 
branching,  and  a  heterocyst  at  the  base  of  each  filament.  Re- 
production by  synakinetes  and  resting  akinetes,  rarely  by  simple 
reproductive  akinetes.  Eivularia,  Glceotrichia,  Isactis,  Calothrix. 

Order  6.  Scytonemaceae.  The  individuals  are  formed  of 
multicellular  filaments  with  no  longitudinal  division ;  differen* 
tiation  into  apex  and  base  very  slight  or  altogether  absent; 


26 


BACTERIA. 


branching  spurious  ;  heterocysts  present.  Reproduction  bj  syna- 
kinetes,  rarely  by  resting  akinetes  and  ordinary  reproductive 
akinetes.  Tolypothrix,  Scytonema,  Hassalia,  Microchcete. 

Order  7.  Sirosiphoniacese.  The  individuals  are  formed  of 
multicellular  threads  with  longitudinal  divisions  ;  true  branching 
and  heterocysts,  and  often  distinct  differentiation  into  apex  and 
base.  Reproduction  by  synakinetes,  rarely  by  resting  akinetes 
and  ordinary  reproductive  akinetes.  Hapalosiphon,  Stigonema, 
Capsosira,  Nostocopsis,  Mastigocoleus. 


Family  2.      Bacteria.* 

The  Bacteria  (also  known  as  Schizomycetes,  and  Fission-Fungi) 
are  the  smallest  known  organisms,  and  form  a  parallel  group  to 
the  Blue-green  Algae,  but  separated  from  these  Algae  by  the 
absence  of  their  colouring  material  ;  chlorophyll  is  perhaps  found 
in  a  few  Bacteria. 

The  various  forms  under  which  the  vegetative  condition  of  the 
Bacteria  appear,  are  termed  as  follows  : 

1.  GLOBULAR  FORMS,  cocci  (Figs.  27,  30  c) :   spherical  or  ellip- 
soidal, single  cells,  which,  however,  are  usually  loosely  massed 
together  and  generally  termed  " Micrococci." 

2.  ROD-LIKE  FORMS  :  more  or  less  elongated  bodies  ;  the  shorter 
forms  have  been  styled  "Bacterium  "  (in  the  narrower  sense  of  the 
word),  and  the  term  "Bacillus  "  has  been  applied  to  longer  forms 

which  are  straight 
and  cylindrical  (Figs. 
28,  29,  30  E). 

3.  THREAD-LIKE 
FORMS  :  unbranched, 
long,  round  filaments, 
resembling  those  of 
Oscillaria,  are  pos- 
sessed by  Leptothrix 
(very  thin,  non-granu  - 
lar  filaments  ;  Fig.  30 
J.,the  small  filaments) 
and  Beggiatoa  (thicker 
filaments,  with  strong,  refractile  grains  or  drops  of  sulphur  (Fig. 

*  The  Bacteria  are  more  usually  included  under  Fungi.  It  seems  better,  how- 
ever, to  place  them  under  the  Algae  in  a  separate  class  with  the  Schizophyceee. 


FIG.  23. — Spirillum  sanguineum.  Four  specimens.  One 
has  two  cilia  at  the  same  end,  the  sulphur  grains  are  seen 
internally. 


BACTERIA. 


27 


31);  often  self-motile).     Branched  filaments,  with  false  branching 
like  many  Scytonemacece,  are  found  in  Cladothrix  (Fig.  30  B,  G). 

4.  SPIRAL  FORMS  :  Rod-like  or  filamentous  bodies,  which  more 
or  less  strongly  resemble  a  corkscrew  with  a  spiral  rising  to  the 
left.     In  general  these  are  termed  Spirilla  (Fig.  23) ;  very  attenu- 
ated spirals,  Vibriones  (standing  next  to  Fig.  30  M)  ;  if  the  filaments 
are  slender  and  flexible  with  a  closely  wound  spiral,  Spirochcetce 
(Fig.  24). 

5.  The  MERISMOPEDITJM- 
FORM,    consisting    of    rounded 
cells   arranged   in    one   plane, 
generally  in  groups  of  four,  and 
produced  by  divisions  perpen- 
dicular to  each  other. 

6.  The  SARCINA-FORM,  con- 
sisting of  roundish  cells  which 
are  produced  by  cellular  divi- 
sion in  all  the  three  directions 
of  space,  united  into  globular 
or  ovoid   masses   ("parcels") 
e.g.    Sarcina   ventriculi    (Figs. 


FIG.  24. —  Spirochcete  obermeieri,  in  active 
motion  (b)and  shortly  before  the  termination 
of  the  fever  (c) ;  a  blood  corpuscles. 


25,  26). 

All  Bacteria  are  unicellular.  In  the  case  of  the  micrococci  this 
is  self-evident,  but  in  the  "  rod,"  "  thread,"  and  "  spiral  "  Bacteria, 
very  often  numerous  cells  remain  united  together  and  their  indi- 
vidual elements  can  only  be  recognised  by  the  use  of  special  re- 
agents. 


FIG.  25. — Sarcina  ventriculi.  One  sur- 
face only  is  generally  seen.  Those  cells 
which  are  drawn  with  double  contour  are 
seen  with  the  correct  focus,  and  more 
distinctly  than  those  cells  lying  deeper 
drawn  with  single  contour. 


FIG.  26. — Sarcina  minuta  :  a-d  succes- 
sive stages  of  one  individual  (from  4-10 
p.m.) ;  /  an  individual  of  32  cells. 


The  condition  termed  "  Zoogloea,"  which  reminds  us  of  Nostoc,  is 
produced  by  the  cells  becoming  strongly  mucilaginous.  A  number 
of  individuals  in  active  division  are  found  embedded  in  a  mass  of 
mucilage,  which  either  contains  only  one,  or  sometimes  more,  of 


28  BACTERIA. 

the  above-named  forms.  The  individuals  may  eventually  swarm 
out  and  continue  their  development  in  an  isolated  condition. 
Such  mucilaginous  masses  occur  especially  upon  moist  vegetables 
(potatoes,  etc.),  on  the  surface  of  fluids  with  decaying  raw  or 
cooked  materials,  etc.  The  mucilaginous  envelope  is  thrown  into 
folds  when  the  Bacteria,  with  their  mucilaginous  cell-walls, 
multiply  so  rapidly  that  there  is  no  more  room  on  the  surface  of 
the  fluid. 

The  cells  of  the  Bacteria  are  constructed  like  other  plant-cells 
in  so  far  as  their  diminutive  size  has  allowed  us  to  observe  them. 
The  cell-wall  only  exceptionally  shows  the  reactions  of  cellulose 
(in  Sarcina,  Leuconostoc ;  also  in  a  Vinegar-bacterium,  Bacterium 
xylinum)  ;  a  mucilaginous  external  layer  is  always  present.  The 
body  of  the  cell  mostly  appears  to  be  an  uniform  or  finely  granu- 
lated protoplasm.  Very  few  species  (e.g.  Bacillus  virens)  contain 
chlorophyll ;  others  are  coloured  red  (purple  sulphur  Bacteria) ; 
the  majority  are  colourless.  Bacillus  amylobacter  shows  a  reaction 
of  a  starch-like  material  when  treated  with  iodine  before  the 
spore-formation.  Some  Bacteria  contain  sulphur  (see  p.  37). 
The  body,  which  has  been  described  as  a  cell-nucleus,  is  still  of  a 
doubtful  nature. 

Artificial  colourings  with  aniline  dyes  (especially  methyl- violet, 
gentian-violet,  methylene-blue,  fuchsin,  Bismarck-brown  and 
Vesuvin)  play  an  important  part  in  the  investigations  of 
Bacteria. 

MOVEMENT.  Many  Bacteria  are  self-motile;  the  long  filaments  of 
Beggiatoa  exhibit  movements  resembling  those  of  Oscillaria.  In 
many  motile  forms  the  presence  of  cilia  or  flagella  has  been  proved 
by  the  use  of  stains  ;  many  forms  have  one,  others  several  cilia 
attached  at  one  or  both  ends  (Fig.  23)  or  distributed  irregularly 
over  the  whole  body ;  the  cilia  are  apparently  elongations  of  the 
mucilaginous  covering  and  not,  as  in  the  other  Alga3  of  the  proto- 
plasm. In  Spirochcete  the  movement  is  produced  by  the  flexibility 
of  the  cell  itself.  Generally  speaking,  the  motion  resembles  that 
of  swarm- cells  (i.e.  rotation  round  the  long  axis  and  movement  in 
irregular  paths) ;  but  either  end  has  an  equal  power  of  proceeding 
forwards. 

The  swarming  motion  must  not  be  confounded  with  the  hopping  motion  of  the 
very  minute  particles  under  the  microscope  (Brownian  movement). 

VEGETATIVE   REPRODUCTION  takes  place  by  continued  transverse 


BACTERIA.  29 

division ;    hence  the  name   "  Fission-Fungi  "   or  "  Fission- Algae," 
has  been  applied  to  the  Bacteria. 

SPORES.  The  spores  are  probably  developed  in  two  ways.  In 
the  ENDOSPOROUS  species  (Figs.  28,  29),  the  spore  arises  as  a  new 
cell  inside  the  mother-cell.  The  spores  are  strongly  refractile, 
smaller  than  the  mother-cell,  and  may  be  compared  to  the 
aplanospores  of  other  Algae.  In  addition  to  these  there  are  the 
ARTHROSPOROUS  species  in  which  the  cells,  just  as  in  Nostoc  and 
other  Blue-green  Algae,  assume  the  properties  of  spores  without 
previously  undergoing  an  endogenous  new  construction,  and  are 
able  to  germinate  and  form  new  vegetative  generations  (Fig.  27). 
The  formation  of  spores  very  often  commences  when  the  vegetative 
development  begins  to  be  restricted. 


FIG.  27. — Leuconostoc  mesenterioides :  a  a  zooglcea,  natural  size ;  b  cross  section  of 
zoogloea;  c  filaments  with  spores;  d  mature  spores;  e-i  successive  stages  of  germina- 
tion ;  in  e  portions  of  the  ruptured  spore-wall  are  seen  on  the  external  side  of  the  muci- 
laginous covering,  (b-i  magnified  520.) 

The  spores  germinate  as  in  JVbsfoc  by  the  bursting  of  the  external 
layer  of  the  cell-wall,  either  by  a  transverse  or  longitudinal  cleft, 
but  always  in  the  same  way,  in  the  same  species  (Fig.  28,  example 
of  transverse  cleft). 

DISTRIBUTION.  Bacteria  and  their  germs  capable  of  development, 
are  found  everywhere,  in  the  air  (dust),  in  surface  water,  and 
in  the  superficial  layers  of  the  soil.  The  number  varies  very 
much  in  accordance  with  the  nature  of  the  place,  season,  etc. 
They  enter,  together  with  air  and  food,  into  healthy  animals  and 
occur  always  in  their  alimentary  tract. 


30 


BACTEE1A. 


GROWTH  AND  REPKODUCTION  depend  upon  the  conditions  of 
temperature.  There  is  a  certain  minimum,  optimum  and  maxi- 
mum for  each  species  ;  for  instance  (in  degrees  Centigrade) — 


d 


FIG.  28. — Bacillus  megaterium :  a  outline  of  a 
living,  vegetative  cell-rod  ;  6  a  living,  motile,  pair  of 
rods ;  p  a  similar  4-celled  rod  after  the  effects  of 
iodine  alcohol ;  c  a  5-celled  rod  in  the  first  stages  of 
spore-formation ;  d-/  successive  stages  of  spore- 
formation  in  one  and  the  same  pair  of  rods  (in  the 
course  of  an  afternoon) ;  r  a  rod  with  mature  spores ; 
g^-g3  three  stages  of  a  5-celled  rod,  with  spores  sown 
in  nutritive  solution ;  Ti'-Ti2,  t,  fc,  I  stages  of  germina- 
tion ;  m.  a  rod  in  the  act  of  transverse  division, 
grown  out  from  a  spore  which  had  been  sown  eight 
hours  previously.  (After  de  Bary ;  a  mag.  250,  the 
other  figures  600  times). 


Bacillus  subtilis 

B.  anthracis 

Spirillum  cholerce  asiaticce 


Bacterium,  tuberculosis 


Minim. 
+    6 
15 
8 


Opt.' 

c.  30 

20-25 

37 


28     37-38 


FIG.  29.— Bacillus  amylobacter. 
Motile  rods,  partly  cylindrical 
and  without  spores,  partly 
swollen  into  various  special 
shapes  and  with  spore-forma- 
tion in  the  swelling,  s  Mature 
spore,  with  thick  mucilaginous 
envelope.  (After  de  Bary ; 
mag.  600  times,  with  the  excep- 
tion of  s,  which  is  more  highly 
magnified.) 


Maxim. 
+   50 
43 

40  (but  grows  only 
feebly  if  under 
16°). 
42 


The  functions  of  life  cease  on  a  slight  excess  of  the  maximum 
or  minimum  temperature,  numbness  setting  in  when  either  of 
these  limits  is  passed.  Crenothrix-thres^ds  provided  with  muci- 
laginous envelopes  may,  according  to  Zopf,  sustain  a  tempera- 
ture of — 10°.  Some  Bacteria  are  said  to  be  able  to  resist  the 
exposure  to  as  low  a  temperature  as — 110°  for  a  short  time.  It  is 
not  known  at  what  degree  of  cold  the  death  of  the  Bacteria  oc- 
curs :  the  greatest  degree  of  heat  which  the  vegetative  cells  can 


BACTERIA.  31 


withstand  is  about  the  same  as  that  for  other  vegetative  plant- 
cells,  namely,  about  50-60°  C.  Certain  Bacteria,  e.g.  B.  thermo- 
philus.  grow  and  thrive  vigorously  at  70°  C.  Many  spores,  on  the 
contrary,  are  able  to  bear  far  higher  temperatures  (in  several 
species  a  temperature  for  some  duration  of  above  100°,  those  of 
Bacillus  subtilis,  for  instance,  can  withstand  for  hours  a  tempera- 
ture of  100°  in  nutrient  solutions  ;  the  spores  remain  capable  of 
development  after  exposure  to  a  dry  heat  of  123°  C.). 

The  Desiccation  of  the  air,  if  prolonged,  kills  many  forms  when 
in  the  vegetative  condition.  The  spores  however  can  bear  a  much 
longer  period  of  dryness,  some  even  several  years. 

OXYGEN.  Some  species  cannot  live  without  a  supply  of  free 
oxygen  (Aerobic),  e.g.  the  Vinegar-bacteria,  the  Hay-bacilli,  the 
Anthrax-bacilli,  the  Cholera- Microspira.  Other  species  again  thrive 
vigorously  without  supply  of  free  oxygen,  and  are  even  checked  in 
their  development  by  the  admission  of  air  (Anaerobic),  e.g.  the 
butyric  acid  Bacterium  (Clostridium  butyricum  =  Bacillus  amy- 
lobacter).  '  A  distinction  may  bo  drawn  between  obligate  and 
facultative  aerobics  and  obligate  and  facultative  anaerobics. 
Several  Bacteria,  producing  fermentation,  may  grow  without  the 
aid  of  oxygen  when  they  are  living  in  a  solution  in  which  they 
can  produce  fermentation ;  but,  if  this  is  not  the  case,  they  can. 
only  grow  when  a  supply  of  oxygen  is  available.  A  great 
number  of  the  pathogenic  Bacteria  belong  to  the  facultative 
anaerobics. 

A  luminous  Bacterium  (Bacillus  plwsphorescens)  which  in  tho 
presence  of  a  supply  of  oxygen  gives  a  bluish-white  light,  has 
been  found  in  sea- water.  Phosphorescent  Bacteria  have  fre- 
quently been  observed  upon  decaying  sea-fish,  as  well  as  on  the 
flesh  of  other  animals  ;  by  transferring  the  Bacteria  from  cod  fish 
to  beef,  etc.,  the  latter  may  be  made  luminous. 

Organic  carbon  compounds  are  indispensable  for  all  Bacteria, 
(except,  as  it  appears,  for  the  nitrifying  organisms),  as  they  can 
only  obtain  the  necessary  supplies  of  carbon  from  this  source.  The 
supplies  of  nitrogen,  which  also  they  cannot  do  without,  can  be  ob- 
tained equally  as  well  from  organic  compounds  as  from  inorganic 
salts,  such  as  saltpetre  or  ammonia-compounds.  The  various  "ash- 
constituents  "  are  also  essential  for  their  nourishment. 

While  Moulds  and  Yeast-Fungi  grow  best  in  an  acid  substratum, 
the  Bacteria,  on  the  other  hand,  generally  thrive  best  in  a  neutral 
or  slightly  alkaline,  one. 


32  BACTERIA. 

In  sterilization,  disinfection,  and  antisepsis,  means  are  employed 
by  which  the  Bacteria  are  killed,  or  checked  in  their  development, 
for  instance,  by  heat  (ignition,  cooking,  hot  vapours,  hot  air,  etc.), 
or  poisons  (acids,  corrosive  sublimate).  The  process  of  preserv- 
ing articles  of  food,  in  which  they  are  boiled  and  then  hermeti- 
cally sealed,  aims  at  destroying  the  Bacteria,  or  the  spores  of 
those  which  already  may  be  present  in  them,  and  excluding 
all  others. 

As  the  Bacteria  are  unable  to  assimilate  carbon  from  the  car- 
bonic acid  of  the  air,  but  must  obtain  it  from  the  carbon- com- 
pounds already  in  existence  in  the  organic  world,  they  are  either 
saprophytes  or  parasites.  Some  are  exclusively  either  the  one  or 
the  other,  obligate  saprophytes  or  parasites.  But  there  are 
transitional  forms  among  them,  some  of  which  are  at  ordinary 
times  saprophytes,  but  may,  when  occasion  offers,  complete  their 
development  wholly  or  partly  as  parasites — facultative  parasites  ; 
others  are  generally  parasitic,  but  may  also  pass  certain  stages  of 
development  as  saprophytes — facultative  saprophytes. 

All  chlorophyll-free  organisms  act  in  a  transforming  and  dis- 
turbing manner  on  the  organic  compounds  from  which  they  obtain 
their  nourishment,  and  while  they  themselves  grow  and  multiply, 
they  produce,  each  after  its  kind,  compounds  of  a  less  degree  of 
complexity,  i.e.  they  produce  fermentation,  putrefaction,  sometimes 
the  formation  of  poisons,  and  in  living  beings  often  disease. 

Those  organisms  which  produce  fermentation  are  called  ferments  • 
this  word,  however,  is  also  employed  for  similar  transformations 
in  purely  chemical  materials  (inorganic  ferments  or  enzymes). 
Many  organic  ("  living ")  ferments,  among  which  are  Yeast- 
cells  and  Bacteria,  give  off  during  their  development  certain 
inorganic  and  soluble  ferments  (enzymes)  which  may  produce 
other  transformations  without  themselves  being  changed.  Dif- 
ferent organisms  may  produce  in  the  same  substratum  different 
kinds  of  transformation ;  alcoholic  fermentation  may  for  instance 
be  produced  by  different  species  of  Fungi,  but  in  different  pro- 
portions, and  the  same  species  produces  in  different  substrata, 
different  transformations  (e.g.  the  Vinegar-bacteria  oxydize  diluted 
alcohol  to  vinegar,  and  eventually  to  carbonic  acid  and  water). 

In  the  study  of  Bacteria  it  is  absolutely  necessary  to  sterilize  the  vessels 
employed  in  cultivation,  the  apparatus,  and  nutrient  solutions,  i.e.  to  free  them 
from  Bacteria  germs  and  also  to  preserve  the  cultures  from  the  intrusion  of  any 
foreign  germs  ("pure-cultures").  A  firm,  transparent,  nutritive  medium  is 


BACTERIA.  33 

frequently  employed.  -This  maybe  prepared  by  adding  to  , the  nutrient  solu- 
tions (broth)  either  gelatine,  or — when  the  Bacteria  are  to  be  cultivated  at 
blood-heat — serum  of  sheep's  or  calf's  blood,  agar-agar  or  carragen  ;  serum 
alone  may  in  itself  serve  as  a  nutrient  medium.  The  so-called  "plate-cul- 
tures "  are  frequently  employed,  i.e.  the  germs  are  isolated  by  shaking  them 
with  the  melted  liquid  nutrient  gelatine,  which  is  then  spread  on  a  glass  plate 
and  allowed  to  coagulate  ;  when  later  on  the  individual  germs  grow  into 
colonies,  these  remain  separate  in  the  solid  substratum  and  it  is  easy  to  pursue 
'heir  further  development.  Similar  plate-cultures  may  also  be  cultivated  in 
test-tubes  and  on  microscopic  slides.  The  slides  and  glass  plates  must  be 
placed  in  "  moist  chambers"  free  from  Bacteria,  By  sowing  a  few  cells  (if  pos- 
sible one)  by  means  of  a  fine  platinum  wire,  pure  cultures  for  further  investi- 
gation may  be  obtained. 

In  order  to  prove  the  relationship  between  pathogenic  Bacteria  and  certain 
diseases,  the  experimental  production  of  pathogenic  Bacteria  by  the  inoculation 
of  Bacteria  from  pure  cultures  into  healthy  animals,  is  very  important. 

It  has  not  so  far  been  possible  to  establish  a  classification  of  the 
Bacteria,  as  the  life-history  of  many  species  has  not  yet  been 
sufficiently  investigated.1  The  opinions  of  botanists  are  at 
variance,  in  many  cases,  with  regard  to  the  forms  of  growth  of  a 
particular  kind.  Some  species  are  pleomorphic  (many-formed) 
while  others  possess  only  one  form. 

The  following  Bacteria  are  Saprophytes  : — 

Cladothrix  dichotoma  is  common  in  stagnant  and  running  water 
which  is  impregnated  with  organic  matter ;  the  cell-chains  have 
false  branching.  According  to  Zopf,  Leptothrix  ochracea  is  one  of 
the  forms  of  this  species  which,  in  water  containing  ferrous  iron 
(e.g.  as  FeC03),  regularly  embeds  ferric-oxide  in  its  sheath  by  means 
of  the  activity  of  the  protoplasm.  Leptothrix  odiracea  and  other 
Iron-bacteria,  according  to  Winogradsky  (1888),  do  not  continue 
their  growth  in  water  free  from  protoxide  of  iron ;  while  they 
multiply  enormously  in  water  which  contains  this  salt  of  iron. 
The  large  masses  of  ochre-coloured  slime,  found  in  meadows,  bogs, 
and  lakes,  are  probably  due  to  the  activity  of  the  Iron-bacteria. 

Those  forms  which,  according  to  Zopf's  views,  represent  the- 
forms  of  development  of  Cladothrix  dichotoma  are  placed  together 
in  Fig.  30.  A  represents  a  group  of  plants,  seventy  times 
magnified,  attached  to  a  Vancheria.  The  largest  one  is  branched 
like  a  tree,  with  branches  of  ordinary  form;  a  specimen  with, 
spirally  twisted  branches  is  seen  to  the  right  of  the  figure,  at 
the  lower  part  some  small  Leptothrix-like  forms.  .B  shows  the- 

1  On  the  classification  of  Bacteria  see  "  System  der  Bakterien,"  by  Dr. 
W.  Migula.  Jena,  1897. 

W.  B.  D 


BACTK1IIA. 


manner    of   branching   and   an    incipient  Coccws- formation.     G   a 
Coccus-m&ss  whose  exit  from  the  sheath  has  been  observed.    D  the 


FIG.  30.— Cladothrix  dichotoma. 


same  mass  as  C  after  the  course  of  a  day,  the  Cocci  having 
turned  into  rods.  E  a  group  of  Cocci  in  which  some  have  deve- 
loped into  shorter  or  longer  rods.  F  one  of  these  rods  before 


BACTERIA.  35 


and  after  treatment  with  picric  acid,  which  causes  the  chain-like 
structure  to  become  apparent.  Q  a  portion  of  a  plant  with  con 
spicuous  sheath,  two  lateral  branches  are  being  formed.  H  part 
of  a  plant,  whose  cells  have  divided  and  form  Cocci.  The  original 
form  of  the  cells  in  which  the  Cocci  are  embedded  may  still  be 
recognised.  I.  Leptothrix-filsim.en.ts  with  conspicuous  mucilaginous 
sheath,  from  which  a  series  of  rods  is  about  to  emerge  ;  the  rod 
near  the  bottom  is  dead,  and  has  remained  lying  in  the  sheath. 
K  part  of  a  plant  which  is  forming  Cocci,  those  at  the  top  are  in 
the  zoogloea-stage,  at  the  base  they  are  elongating  to  form  rods 
and  Leptothrix-nl&ments.  L  a  portion  of  a  branched  Cladothrix, 
which  divides  into  motile  Bacillus-iorms ;  the  rays  at  the  free  i 
•ends  indicate  the  currents  which  the  cilia  produce  in  the  water.  •„ 
M  a  spirally-twisted,  swarming  filament,  before  and  after  division 
into  halves.  N  part  of  a  tree-like  zooglcea  with  Cocci  and  short 
rods. — All  of  these  spirilla,  zooglcea,  etc.,  which  Zopf  has  con- 
nected with  Glad,  dichotoma,  are  according  to  Winogradsky,  inde- 
pendent organisms. 

Micrococcus  urece  produces  urinal  fermentation  (transformation  of 
urinal  matter  into  ammonium  carbonate)  ;  aerobic  ;  round  cells 
generally  united  to  form  bent  chains  or  a  zoogloea. — Several  other 
kinds  of  Bacteria  have  the  same  action  as  this  one  :  in  damp 
soil  containing  ammonia-compounds,  saltpetre-formations  are  pro- 
duced by  M.  nitrificans  and  several  different  kinds  of  Bacteria. 

Micrococcus  prodigiosus  is  found  on  articles  of  food  containing 
starch;  "bleeding  bread"  is  caused  by  this  Bacterium,  which  has 
the  power  of  forming  a  red  pigment ;  it  also  occurs  in  milk,  and 
produces  lactic  acid. 

Leuconostoc  mesenterioides  is  the  frog-spawn  Bacterium  (Fig.  27) 
,  which  is  found  in  sugar  manufactories,  and  has  the  power  of 
producing  a  viscous  fermentation  in  saccharine  solutions  which 
have  been  derived  from  plants,  e.g.  in  beetroot-sugar  manu- 
factories, where  large  accumulations  of  mucilage  are  formed  at 
the  expense  of  the  sugar,  with  an  evolution  of  carbonic  acid.  The 
•cell-rows,  resembling  somewhat  a  pearl  necklace,  have  thick 
mucilaginous  cell-walls,  and  form  white  "  Nostoc  "-lumps.  The 
mucilage  eventually  deliquesces  and  the  cells  separate  from  each 
other ;  arthrospores  ? — Similar  viscous  deteriorations  occur  in 
beer  and  wine,  which  may  then  be  drawn  out  into  long,  string 
like  filaments — "  ropiness." 

Bacterium  aceti,  the  Vinegar-bacterium,   oxidizes   alcohol  into 


36  BACTERIA. 

acetic  acid  (acetous-fermentation)  and  forms  a  greyish  covering 
of  Bacteria  ("  Vinegar-mother  ")  on  the  surface  of  the  liquid  ;  the 
acetic  acid  formed,  becomes  by  continued  oxidization  by  B.  aceti, 
again  transformed  into  carbonic  acid  and  water.  Aerobic; 
short  cylindrical  cells,  often  united  into  chains,  or  to  form  a 
zoogloea  ;  sometimes  also  rod-  and  spindle-shaped.  The  Vinegar- 
bacteria  and  other  kinds  with  ball-  or  rod-forms  sometimes  become 
swollen,  spindle-shaped,  or  oval  links;  they  are  supposed  to  be 
diseased  forms1  ("Involution-forms"). 

Bacillus  laciicus  (Bacterium,  acidi  lactici,  Zopf)  is  always  found 
in  milk  which  has  stood  for  some  time,  and  in  sour  foods  (cabbage, 
cucumbers,  etc.)  ;  it  turns  the  milk  sour  by  producing  lactic  acid 
fermentation  in  the  sugar  contained  in  the  milk  ;  the  lactic  acid 
formed,  eventually  causes  the  coagulation  of  the  casein.  It  re- 
sembles the  Vinegar-bacteria,  occurring  as  small  cylindrical  cells, 
rarely  in  short  rows;  not  self-motile. — Several  other  Bacteria  appear 
to  .act  in  the  same  way,  some  occurring  in  the  mouth  of  human! 
beings  ;  some  of  these  Bacteria  give  to  butter  its  taste  and  flavour. 

The  'kefir-grains  which  are  added  to  milk  for  the  preparation  of 
kefir,  contain  in  large  numbers  a  Bacterium  (Dispora  caucasica) 
in  the  zooglcea-form,  a  Yeast-fungus,  and  Bacillus  lacticus.  Kefir 
is  a  somewhat  alcoholic  sour  milk,  rich  in  carbonic  acid ;  it  is  a 
beverage  manufactured  by  the  inhabitants  of  the  Caucasus,  from 
the  milk  of  cows,  goats,  or  sheep,  and  is  sometimes  used  as  a 
medicine.  In  the  production  of  kefir,  lactic  acid  fermentation 
takes  place  in  one  part  of  the  sugar  contained  in  the  milk,  and 
alcoholic  fermentation  in  another  part,  and  the  casein  which  had 
become  curdled  is  partially  liquefied  (peptonised)  by  an  enzyme  of 
a  Zooglcea-bacterium. 

Bacillus  amylobacter  (Bacillus  butyricusj,  the  Butyric- acid-bac- 
terium (Fig.  29),  is  a  very  common  anaerobic  which  produces 
fermentation  in  sugar  and  lactic-acid  salts,  and  whose  principal 
product  is  butyric  acid.  It  destroys  articles  of  food  and  (together 
with  other  species)  plays  a  part  in  the  butyric  acid  fermentation 
which  is  necessary  in  the  making  of  cheese  ;  it  is  very  active 
wherever  portions  of  plants  are  decaying,  in  destroying  the  cellu- 
lose in  the  cell-walls  of  herbaceous  plants,  and  is  thus  useful  in 
the  preparation  of  flax  and  hemp.  The  cells  are  self-motile, 
generally  cylindrical,  sometimes  united  into  short  rows  ;  endo- 

1  According  to  Hansen  these  are  not  disease  forms,  but  occur  regularly  under 
certain  conditions,  e.g.  temperature. 


BACTERIA. 


37 


sporous;  the  spore-forming  cells  swell,  assume  very  different  forms, 
and  show  granulose  reaction.  The  germ-tube  grows  out  in  the 
direction  of  the  long  axis  of  the  spore. 

Bacillus  subtilis,  the  Hay-bacillus,  is  developed  in  all  decoctions 
of  hay ;  a  slender,  aerobic,  self-motile  Bacillus ;  endosporous  (aplano- 
spores) ;  the  spore-wall  ruptures  transversely  on  germination. 

Grenothrix  kilhniana  occurs  in  the  springs  of  many  baths,  in 
wells,  in  water  or  drain-pipes. 

Beggiatoa  (parallel  with  the  Blue-green  Alga  Oscillaria).  Long 
filaments  formed  of  cylindrical  cells  which  are  attached  by  one  of 
the  ends,  but  which  are  nearly  always  free  when  observed.  The 
filaments,  like  those  of  OsciUaria,  describe  conical  figures  in  their 
revolutions,  the  free  filaments  slide  upwards  and  parallel  with  one 
another;  sheaths  are  wanting;  strongly 
refractive  sulphur  drops  are  found  in  the 
interior.  The  Beggiatoas  are  the  most 
prevalent  Sulphur-bacteria.  They  occur, 
very  commonly  in  large  numbers,  wher- 
ever plant  or  animal  remains  are  de- 
caying in  water  in  which  sulphuretted 
hydrogen  is  being  formed ;  thus,  for  ex- 
ample, B.  alba  (Fig.  31)  occurs  frequently 
as  a  white  covering  or  slimy  film  on  mud 
containing  organic  remains.  B.  miralrilis  is 
remarkable  for  its  size  and  its  strong  peristaltic 
movements.  The  Sulphur-bacteria  oxidize 
the  sulphuretted  hydrogen,  and  accu- 
mulate sulphur  in  the  shape  of  small 
granules  of  soft  amorphic  sulphur,  which 
in  the  living  cell  never  passes  over  into 
the  crystalline  state.  They  next  oxidize 
this  sulphur  into  sulphuric  acid,  which 
is  immediately  rendered  neutral  by  ab- 
sorbed salts  of  calcium,  and  is  given 
off  in  the  form  of  a  sulphate,  thus  CaC03  is  principally  changed 
into  Ca  S04.  In  the  absence  of  sulphur  the  nutritive  processes 
are  suspended,  and  consequently  death  occurs  either  sooner  or 
later.  The  Sulphur- bacteria  may  exist  and  multiply  in  a  fluid 
which  only  contains  traces  of  organic  matter,  in  which  organisms 
devoid  of  chlorophyll  are  not  able  to  exist.  The  Beggiatoas 
very  frequently  form  white,  bulky  masses  in  sulphur  wells  and 


PIG.  31.  — Beggiatoa  alba:  a 
from  a  fluid  containing  abund- 
ance of  sulphuretted  hydrogen ; 
b  after  lying  24  hours  in  a  solu- 
tion devoid  of  sulphuretted 
hydrogen ;  e  after  lying  an 
additional  48  hours  in  a  solution 
devoid  of  sulphuretted  hydro- 
gen, by  this  means  the  trans- 
verse walls  and  vacuoles  have 
become  visible. 


38  BACTERIA. 

in  salt  water,  the  traces  of  organic  material  which  the  sulphur 
water  contains  proving  sufficient  for  them.  The  cellulose-fermenta- 
tion, to  which  the  sulphur  wells  in  all  probability  owe  their  origin,  mainly 
procures  them  suitable  conditions  for  existence.  The  CaC03  and  H2S, 
formed  during  the  cellulose  fermentation  by  the  reduction  of  CaS04  is 
again  changed  into  CaS04  and  C02  by  the  Sulphur-bacteria  (Winogradsky, 
1887).— Other  Sulphur-bacteria,  the  so-called  purple  Sulphur-bacteria,  e.g. 
B.  roseo-persicina,  Spirillum  sanguineum  (Fig.  23),  Bacterium  sulfuratum,  etc., 
have  their  protoplasm  mixed  with  a  red  colouring  matter  (bacterio-purpurin) 
which,  like  chlorophyll,  has  the  power,  in  the  presence  of  light,  of  giving  off 
oxygen  (as  proved  by  T.  W.  Englemann,  1888,  in  oxygen-sensitive  Bacteria). 
The  three  purple  Sulphur-bacteria  mentioned,  are,  according  to  Winogradsky, 
not  pleomorphic  kinds  but  embrace  numerous  species. 

Many  Spirilli  (Spirillum  tenue,  S.  undula,  S.plicatile,  and  others) 
are  found  prevalent  in  decaying  liquids. 

Bacteria  (especially  Bacilli)  are  the  cause  of  many  substances 
emitting  a  foul  odour,  and  of  various  changes  in  milk. 

Parasitic  Bacteria  live  in  other  living  organisms  ;  but  the 
relation  between  "  host "  and  parasite  may  vary  in  considerable 
degree.  Some  parasites  do  no  injury  to  their  host,  others  produce 
dangerous  contagious  diseases  ;  some  choose  only  a  special  kind 
as  host,  others  again  live  equally  well  in  many  different  ones. 
There  are  further  specific  and  individual  differences  with  regard 
to  the  predisposition  of  the  host,  and  every  individual  has  not  the 
same  receptivity  at  all  times. 

THE  HARMLESS  PARASITES  OF  HUMAN  BEINGS.  Several  of  the 
above  mentioned  saprophytes  may  also  occur  in  the  alimentary 
canal  of  human  beings;  e.g.,  the  'Hay-bacillus,  the  Butyric-acid- 
bacillus,  etc. ;  but  the  gastric  juice  prevents  the  development  of 
others,  at  all  events  in  their  vegetative  condition.  Sarcina  ven- 
tricidi,  "packet-bacterium,"  is  only  known  to  occur  in  the  stomach 
and  intestines  of  human  beings,  and  makes  its  appearance  in  cer- 
tain diseases  of  the  stomach  (dilation  of  the  stomach,  etc.)  in  great 
numbers,  without,  however,  being  the  cause  of  the  disease.  It 
occurs  in  somewhat  cubical  masses  of  roundish  cells  (l^ig.  25). 

LESS  DANGEROUS  PARASITES.  In  the  mouth,  especially  between 
and  on  the  teeth,  a  great  many  Bacteria  are  to  be  found  (more 
than  fifty  species  are  known),  e.g.  Leptothrix  buccalis  (long,  brittle, 
very  thin  filaments  which  are  united  into  bundles),  Micrococci  in 
large  lumps,  Spirochcete  cohnii,  etc.  Some  of  them  are  known  to 
be  injurious,  as  they  contribute  in  various  ways  to  the  decay  of  the 
teeth  (caries  dentiuni) ;  a  Micrococcus,  for  instance,  forms  lactic  acid 


BACTERIA.  39 

in  materials  containing  sugar  and  starch,  and  the  acid  dissolves 
the  lime  salts  in  the  external  layers  of  the  teeth  :  those  parts  of 
the  teeth  thus  deprived  of  lime  are  attacked  by  other  Bacteria,  and 
become  dissolved.  Inflammation  in  the  tissues  at  the  root  of  a 
tooth,  is  probably  produced  by  septic  materials  which  have  been 
formed  by  Bacteria  in  the  root-canal. 

DANGEROUS  PARASITES.  In  a  large  number  of  the  infectious  dis- 
eases of  human  beings  and  animals,  it  has  been  possible  to  prove 
that  parasitic  Bacteria  have  been  the  cause  of  the  disease.  Various 
pathogenic  Bacteria  of  this  nature,  belonging  to  the  coccus,  rod, 
and  spiral  Bacteria  groups,  are  mentioned  in  the  following :  — 

Pathogenic  Micrococci.  Staphylococcus  pyogenes  aureus  pro- 
duces abscesses  of  various  natures  (boils,  suppurative  processes  in 
internal  organs).  The  same  effects  are  produced  by — 

Streptococcus  pyogenes,  which  is  the  most  frequent  cause  of  malig- 
nant puerperal  fever;  it  is  perhaps  identical  with — 

Streptococcus  erysipelatis,  which  is  the  cause  of  erysipelas  in 
human  beings. 

Diplococcus  pneumonice  (A.  Frankel)  is  the  cause  of  pneumonia, 
and  of  the  epidemic  cerebro-spinal  meningitis. 

Gonococcus  (Neisser)  is  the  cause  of  gonorrhea  and  inflammation 
of  the  eyes. 

Pathogenic  Rod- Bacteria.  Bacterium  cholerm  gallinarum, 
an  aerobic,  facultative  parasite  which  produces  fowl- cholera  among 
poultry  ;  it  is  easily  cultivated  on  various  substrata  as  a  sapro- 
phyte. The  disease  may  be  conveyed  both  through  wounds  and 
by  food,  and  may  also  be  communicated  to  mammals. 

Bacillus  anthracis,  the  Anthrax  bacillus  (Fig.  32),  chiefly  attacks 
mammals,  especially  herbivorous  animals  (house  mice,  guinea-pigs, 
rabbits,  sheep,  cattle),  in  a  less  degree  omnivorous  animals  (includ- 
ing human  beings),  and  in  a  still  less  degree  the  Carnivores. 
Aerobic.  Cylindrical  cells,  3-4  times  as  long  as  broad,  united  into 
long  rod-like  bodies,  which  may  elongate  into  long,  bent,  and 
twisted  filaments.  Not  self-motile.  Endosporous.  Germination 
takes  place  without  the  throwing  off  of  any  spore-membrane  (com- 
pare Hay-bacillus  p.  37  which  resembles  it).  Contagion  may  take 
place  both  by  introduction  into  wounds,  and  from  the  mucous 
membrane  of  the  intestines  or  lungs,  both  by  vegetative  cells  and 
by  spores  ;  in  intestinal  anthrax,  however,  only  by  spores.  The 
Bacillus  multiplies  as  soon  as  it  has  entered  the  blood,  and  the 
anthrax  disease  commences.  The  Bacilli  not  only  give  off  poison, 


40 


BACTERIA. 


but  also  deprive  the  blood  of  its  oxygen.  Vegetative  cells  only 
occur  in  living  animals.  This  species  is  a  facultative  parasite  which 
in  the  first  stage  is  a  saprophyte,  and  only  in  this  condition  forms 
spores. 

Bacillus  tuberculosis  produces  tuberculosis  in  human  beings,  also 
in  domestic  animals  (perlsudit}.  It  is  a  distinct  parasite,  but  may 
also  live  saprophytically.  It  is  rod-formed,  often  slightly  bent, 
and  is  recognised  principally  by  its  action  with  stains  (when 
stained  with  an  alkaline  solution  of  methyl-blue  or  carbolic  f  uch- 


FIG.  32.— Anthrax  bacillus  (Bactl-  FIG.  33.— Anthrax  bacillus.     The  formation  of 

lus  anthracis)  with  red  i,b)  and  white  the  spores  ;  magnified  450  times, 

(a)  blood- corpuscles. 

sin,  it  retains  the  colour  for  a  long  time  even  in  solutions  of  mineral 
acids,  in  contrast  with  the  majority  of  well-known  Bacteria):  it 
probably  forms  spores  which  are  able  to  resist  heat,  cfryness,  etc. 

Bacillus  leprce  produces  leprosy  ;  Bacillus  mallei  produces  glanders ;  Bacillus 
tetani,  tetanus  (the  tetanus  bacillus  is  very  common  in  soil ;  anaerobic) ;  Bacillus 
diphtheria,  diphtheria;  Bacillus  typhosus,  typhoid  fever,  etc. 

Pathogenic  Spiral  Bacteria.  Spirochocte  obermeieri  (Fig 
24)  produces  intermittent  fever  (febris  recurrens) ;  it  makes  its 
appearance  in  the  blood  during  the  attacks  of  fever,  but  it  is  not 
to  be  found  during  intervals  when  there  is  no  fever.  Obligate 
parasite. 

Spirillum  cholerce  asiaticce  (Microspira  comma)  without  doubt 
produces  Asiatic  cholera  ;  an  exceedingly  motile  spirillum,  which 
is  also  found  in  short,  bent  rods  (known  as  the  "Comma-bacillus"), 


BACTERIA.  40A 

it  lives  in  the  intestines  of  those  attacked  by  the  disease,  and 
gives  off  a  strong  poison  which  enters  the  body.  It  is  easily 
cultivated  as  a  saprophyte. 

A  great  many  circumstances  seem  to  show  that  a  number  of 
other  infectious  diseases  (syphilis,  small-pox,  scarlet-fever,  measles, 
yellow-fever,  etc.)  owe  their  origen  to  parasitic  Bacteria,  but  this 
has  not  been  proved  with  certainty  in  all  cases. 

It  has  been  possible  by  means  of  special  cultivations  (ample 
supply  of  oxygen,  high  temperature,  antiseptic  materials)  to  pro- 
duce from  the  parasitic  Bacteria  described  above  (e.gr.  the  fowl-cholera 
and  the  anthrax  Bacteria)  physiological  varieties  which  are  distinct 
from  those  appearing  in  nature  and  possess  a  less  degree  of 
"virulence,"  i.e.  produce  fever  and  less  dangerous  symptoms  in 
those  animals  which  are  inoculated  with  them.  The  production 
of  such  physiological  varieties  has  come  to  be  of  great  practical 
importance  from  the  fact  that  they  are  used  as  vaccines,  i.e.  these 
harmless  species  produce  in  the  animals  inoculated  with  them 
immunity  from  the  malignant  infectious  Bacteria  from  which 
they  were  derived.  This  immunity  is  effected  by  the  change  of 
the  products  of  one  or  more  of  the  Bacteria,  but  we  do  not  yet 
know  anything  about  the  way  in  which  they  act  on  the  animal 


FIG.  34.— a  and  b  The  same  blood-cell 
of  a  Frog :  a  in  the  act  of  engulfing  an 
anthrax-bacillus ;  b  after  an  interval  of 
a  few  minutes  when  the  bacillus  has 
been  absorbed. 


organism.  The  white  blood  corpuscles,  according  to  the  Met- 
schnikoff,  play  the  part  of  "Phagocytes"  by  absorbing  and 
destroying  the  less  virulent  Bacteria  which  have  entered  the 


40B 


BACTERIA. 


blood,  and  by  so  doing  they  are   gradually  enabled  to  overcome 
those  of  a  more  virulent  nature. 

BACTERIA  PATHOGENIC  TO  PLANTS.  Recent  investigations  have 
thoroughly  proved  the  parasitic  action  of  Bacteria  upon  plant- 
tissues,  and  certain  diseases  of  plants  have  undoubtedly  been  traced 
to  these  organisms,  though  their  behaviour  as  vegetable  parasites 
in  other  cases  has  not  been  strictly  denned.  The  following 
examples  may  be  quoted: — 


FIG.  31x. — A  Cell  from  Turnip  attacked  by  Pseudomonas  destructans.  The  bacteria  are 
seen  in  the  cell-cavity  and  along  the  track  of  the  middle  lamella.  Tue  cell-wall  is  much 
swollen  ;  at  a  it  is  just  beginning  to  separate  along  the  middle  lamella;  at  b  the  dissociation 
is  more  strongly  marked. 

B  P.  destructans  with  single  polar  flagellum. 


Pseudomonas  destructans  causes  the  White  Rot  of  the  Turnip 
(Brassica  napus).  The  action  of  this  Bacterium  upon  the  cell> 
of  its  host  is  similar  to  that  of  certain  parasitic  Fungi ;  it  prodiices 
a  toxin  which  kills  the  protoplasm,  and  further  secretes  an  enzyme 
which  has  the  power  of  dissolving  the  middle  lamella,  and  causing 
the  softening  and  swelling  of  the  cell-wall.  By  this  means  the 
Bacteria  rapidly  invade  the  tissues  of  the  host,  burrowing  through 
the  cell- walls  and  extending  into  the  cell-cavities  and  intercellular 
spaces  (Fig.  34A.)  ;  the  cell- sap  escapes  with  the  death  of  the  cells, 


CONJUGATE.  41 

and  the  root  is  reduced  to  a  mere  watery  pulp.  This  organism 
generally  infects  the  plant  through  a  wound,  but  it  is  able  to 
enter  the  epidermis  of  undeveloped  leaves. 

Pseudomonas  campestris  produces  a  brown  discolouration  of 
the  vascular  tissue  in  the  roots  and  leaves  of  Cruciferous  plants. 
It  attacks  the  woody  portion  of  the  vascular  bundles,  and  is 
found  in  great  numbers  blocking  up  the  wood- vessels.  The  in- 
fection of  the  leaves  may  take  place  through  the  water-pores,  and 
spreads  thence  along  the  vascular  bundles  to  other  parts  of  the 
plant. 

Pseudomonas  Tiyacinthi  is  pathogenic  to  Hyacinths,  spreading 
especially  through  the  vascular  system.  Yellow  spots  are  found 
on  the  bulbs  and  foliage  which  are  due  to  the  vessels  becoming 
filled  with  a  yellow  mucilage  produced  by  the  Bacteria. 

Bacillus  solanacearum  attacks  the  Tomato,  Egg-plant,  and 
Potato,  causing  the  withering  and  final  death  of  the  young 
shoots. 

Bacillus  betce  gives  rise  to  the  "  Gummosis "  of  the  Sugar 
Beet, 

Micrococcus  tritici  is  an  organism  which  infects  the  grain  of 
Wheat,  and  preys  upon  the  proteids  and  starch,  forming  what  is 
known  as  "  Pink  Wheat." 

Bacillus  mali,  it  has   lately  been  shown,  is   the  cause   of  the 
cankers  so  common  on  the  Apple  and  Pear.     This  parasite  is  pro 
bably  carried  by  Aphides,  and  effects  an  entrance  through   the 

wouuds  caused  by  these  insects. 

Class  5.     Conjugatae. 

The  Alga3  belonging  to  this  class  have  chlorophyll,  and  pyre- 
noids  round  which  starch  is  formed.  The  cells  divide  only  in  one 
direction,  they  live  solitarily,  or  united  to  form  filaments  which 
generally  float  freely  (seldom  attached).  Swarm-cells  are  want- 
ing. TJie  fertilisation  is  isogamous  (conjugation}  and  takes  place 
by  means  of  aplano gametes.  The  zygote,  after  a  period  of  rest, 
produces,  immediately  on  germination,  one  or  more  new  vegetative 


42  CONJUGATE. 

individuals ;    sometimes  akinetes    or   aplanospores   are  formed   in 
addition.     They  only  occur  in  fresh  or  slightly  brackish  water. 

Order  1.  Desmidiaceae.  The  cells  generally  present  markings 
on  the  outer  wall,  and  are  mostly  divided  into  two  symmetrical 
halves  by  a  constriction  in  the  middle,  or  there  is  at  least  a 
symmetrical  division  of  the  protoplasmic  cell-contents.  The  cell- 
wall  consists  nearly  always  of  two  layers,  the  one  overlapping 
the  other  (Fig.  35  0).  The  cells  either  live  solitarily  or  are  united 
into  unbranched  filaments.  The  mass  of  protoplasm  formed  by 
the  fusion  of  the  two  conjugating  cells  becomes  the  zygote,  which 
on  germination  produces  one  (or  after  division  2,  4  or  8)  new 
vegetative  individual.  The  chromatophores  are  either  star-,  plate-, 
or  band-shaped,  and  regularly  arranged  round  the  long  axis  of  the 
cell. 


FIG.  35.— .4  Cell  of  Gymnozyga  Irelissonii,  external  view  showing  the  distribution  of  the 
pores.  B  A  portion  of  the  membrane  of  SLauvastrum  licorne  with  pores  containing  proto- 
plasmic projections.  C  Cell-wall  of  Hyalotheca  tnucosa  during  cell-division  ;  the  central 
part,  being  already  formed,  shows  the  connection  with  the  divisional  wall. 

The  Desmidiaceoe  are  not  able  to  swim  independently,  many 
species,  however,  show  movements  of  different  kinds  by  rising  and 
sliding  forward  on  the  substratum.  These  movements,  which  are 
partly  dependent  upon,  and  partly  independent  of  light  and  the 
force  of  gravitation,  are  connected  with  the  protrusion  of  a  mu- 
cilaginous stalk.  The  mucilage,  which  sometimes  surrounds  the 
whole  individual,  may  acquire  a  prismatic  structure,  it  is  secreted 
by  the  protoplasmic  threads  which  project  through  certain  pores 
definitely  situated  in  the  walls  (Fig.  35  A,  B). 

VEGETATIVE  MULTIPLICATION  takes  places  by  division.  A  good 
example  of  this  is  found  in  Cosmarium  botrytis  (Fig.  36  A-D}. 
The  nucleus  and  chromatophores  divide,  and  simultaneously  the 
central  indentation  becomes  deeper,  the  outer  wall  is  then  ruptured 
making  a  circular  aperture  through  which  the  inner  wall  protrudes 
forming  i  short,  cylindrical  canal  between  the  two  halves  to  which 
it  is  attached  (Fig.  36  (7).  After  elongation  the  canal  is  divided 
by  a  central  transverse  wall,  which  commences  as  a  ring  round  its 


CONJUGATE. 


inner  surface  and  gradually  forms  a  complete  septum.  The  divid- 
ing wall  gradually  splits,  and  the  tw  individuals  separate  from 
each  other,  each  one  having  an  old  and  a  new  half.  The  two 


FIG.  36.— Cosmarium  lotrytis.    A-D  Different  stages  of  cell-division. 

daughter-cells  bulge  out,  receive  a  supply  of  contents  from  the 
parent-cells,  and  gradually  attain  their  mature  size  and  develop- 
ment (Fig.  36  B-D}.  Exceptions  to  this  occur  in  some  forms. 

CONJUGATION  takes  place  in  the  simplest  way  in  Mesotcenium,  where 
the  two  conjugating  cells  unite  by  a  short  tube  (conjugation-canal), 
which  is  not  developed  at  any  particular  point.  The  aplanogametes 
merge  together  after  the  dissolution  of  the  dividing  wall,  like  two 
drops  of  water,  almost  without  any  trace  of  preceding  contraction, 
so  that  the  cell-wall  of  the  zygote  generally  lies  in  close  contact 
with  the  conjugating  cells.  The  conjugating  cells  in  the  others 
lie  either  transversely  (e.g.  Cosmarium,  Fig.  37  d  ;  Staurastrum, 
etc.),  or  parallel  to  one  another  (e.g.  Penium,  Closterium,  etc.),  and 


FIR.  37. — Cosmarium  meneghinii:  a-c  same  individual  seen  from  the  side,  from  the  end, 
and  from  the  edge ;  d-/  stages  of  conjugation  ;  g-i  germination  of  the  zygote. 

emit  a  short  conjugation-canal  (Fig.  37  d}  from  the  centre  of  that 
side  of  each  cell  which  is  turned  towards  the  other  one.  These 
canals  touch,  become  spherical,  and  on  the  absorption  of  the  divid- 
ing wall  the  aplanogametes  coalesce  in  the  swollen  conjugation- 
canal  (Fig.  37  e),  which  is  often  surrounded  by  a  mucilaginous 
envelope.  The  zygote,  which  is  often  spherical,  is  surrounded  by 
a  thick  cell-wall,  consisting  of  three  layers ;  the  outermost-  of  these 


44  CONJUGATE 

sometimes  bears  thorn-like  projections,  which  in  some  species 
are  simple  (Fig.  37  /),  in  others  branched  or  variously  marked  ; 
in  some,  however,  it  remains  always  smooth  (e.g.  Tetmemorus, 
Desmidium).  Deviation  from  this  mode  of  conjugation  may  occur 
within  certain  genera  (e.g.  Glosterium,  Penium).  Upon  germina- 
tion the  contents  of  the  zygote  emerge,  surrounded  by  the  inner- 
most layers  of  the  wall  (Fig.  37  g,  h),  and  generally  divide  into 
two  parts  which  develope  into  two  new  individuals,  placed  trans- 
versely to  each  other  (Fig.  37  i) ;  these  may  have  a  somewhat  more 
simple  marking  than  is  generally  possessed  by  the  species. 


FIG.  38. — Desmidiacese.  A  Closterium  moniliferum  ;  B  Penium  crassiusculnm ;  C 
Micr aster ias  Lruncata  (front  and  end  view)  ;  D  Euastrum  elegans ;  E  Staura>>trum  muticum, 
(end  view). 

The  most  frequent  genera  are  : — 

A.  Solitary  cells  :     MESOT^ENIUM,   PENIUM   (Fig.   38    B),   CYLINDBOCYSTIS, 
EUASTBUM    (Fig.   38  D),   MICEASTEBIAS  (Fig.  38  C),  COSMABIUM  (Fig.  36,  37), 
XANTHIDIUM,  STAUBASTBUM  (Fig.  38  E),  PLEUROT^NIUM,  DOCIDIUM,  TETMEMOBUS, 
CLOSTEBIUM  (Fig.  38  A),  SPIBOT^NIA. 

B.  Cells  united  into  filaments :    SPH^BOZOSMA,   DESMIDIUM,   HYALOTHECA, 
GYMNOZYGA,  ANCYLONEMA,  GONATOZYGON. 

Order  2.  Zygnemaceae.  Cell-wall  without  markings.  The 
cells  are  cylindrical,  not  constricted  in  the  centre,  and  (generally) 
united  into  simple,  unbranched  filaments.  The  whole  contents  of 
the  conjugating  cells  take  part  in  the  formation  of  the  zygote, 
which  on  germina.tion  grows  out  directly  into  a  new  filament. 

Spirogyra  is  easily  recognised  by  its  spiral  chlorophyll  band  ; 
Zyynema  has  two  star-like  chromatophores  in  each  cell  (Fig.  40)  ; 
both  these  genera  are  very  common  Algae  in  ponds  and  ditches. 


CONJUGATE. 


45 


The   conjugation   among   the   Zygnemacese   takes  place  in  the 
following  manner :  the  cells  of  two  filaments,  lying  side  by  side,  or 


A  B 

FIG.  39.— Spirogyra  longata.  A  At  the  commencement  of  conjugation,  the  conjugation- 
canals  begin  to  protrude  at  a  and  touch  one  another  at  b  ;  the  spiral  chlorophyll  band  and 
•cell-nuclei  (fc)  are  shown.  B  A  more  advanced  stage  of  conjugation;  a,  a'  the  rounded 
female  and  male  aplanogametes  :  in  b'  the  male  aplanogamete  is  going  over  to  and 
uniting  with  the  female  aplanogamete  (b). 

two  cells,  the  one  being  situated  above  the  other  in  the  same  fila- 
ment (Fig.  41),  push  out  small  protuberances  opposite  each  other 
(Fig.  39  A,  a,  6)  ;  these  finally  meet,  and  the  dividing  wall  is  ab- 
sorbed so  that  a  tube  is  formed  connecting  one  cell  with  the  other; 


FIG.  40.— A  cell  of  Zygnema.    S  Pyrenoid. 


FIG.  41.— Zygnema  insigne,  with  zygote. 


the  protoplasmic  contents  round  off,  and  the  whole  of  these  contents 
of  one  of  the  cells  glides  through  the  conjugation-tube  and  coalesces 

with  that  of  the 
other  (Fig.  39  B\ 
the  aggregate  mass 
then  rounds  off,  sur- 
rounds itself  with 

FIG.  42.— Germinating  zygote  of  Spirogyra  jugalis :  the       a  Cell-Wall,    and    be- 
.young  plant  is  still  unicellular ;  the  end  which  is  still  in 
the  wall  of  the  zygote  is  elongated  and  root-like;   tho 
•chromatophore  divides  and  forms  the  spiral  band. 


comes  a  zygote.     A 
distinct  •     difference 


46  CHLOROPHYCEJ;. 

may  be  found  between  the  cells  in  the  two  filaments,  those  in  the 
one  whose  protoplasmic  contents  pass  over  being  cylindrical,  while 
those  of  the  recipient  one  are  more  barrel-shaped,  and  of  a  larger 
diameter.  The  former  may  be  regarded  as  a  male,  the  latter  as 
a  female  plant.  The  zygote  germinates  after  a  period  of  rest,  and 
grows  out  into  a  new  filament  (Fig.  42). 

Order  3.  Mesocarpaceae.  The  cell-walls  are  glabrous,  uncon- 
stricted  in  the  centre,  and  united  into  simple  unbranched  filaments. 
The  chromatophore  consists  of  an  axial  chlorophyll-plate,  with 
several  pyrenoids.  The  zygote  is  formed  by  the  coalescence  of  two 
cells  (Fig.  43)  (sometimes  three  or  four),  but  the  whole  proto- 
plasmic contents  of  the  cells  do  not  take  part  in  this  process,  a 
portion  always  remaining  behind ;  the  aplanogametes  coalesce  in 
the  conjugation-canal.  The  zygote  thus  formed  appears  incapable 
of  germination  until  after  3-5  divisions.  Of  the  cells  so  formed, 


FIG.  48. — Mougeotia  calcarea.  Cells  showing  various  modes  of  conjugation  :  at  m  tripar- 
tition  ;  pg  quadripartition ;  s  quinquipartition  of  the  zygote. 

only  one  is  fertile,  the  sterile  cells,  according  to  Pringsheim,  con- 
stituting a  rudimentary  sporocarp.  The  germinating  cells  grow 
out  into  a  new  filament.  In  this  order,  conjugation  has  been  ob- 
served between  two  cells  of  the  same  filament.  The  Mesocarpacea? 
thrive  best  in  water  which  contains  lime. 

Class  6.    Chlorophyceae  (Green  Algae). 

These  Algse  are  coloured  green  by  chlorophyll,  seldom  in  com- 
bination with  other  colouring  matter,  and  then  especially  with  red. 
The  product  of  assimilation  is  frequently  starch,  which  generally 
accumulates  round  certain  specially  formed  portions  of  protoplasm 
termed  pyrenoids.  The  thallus  is  uni- or  multi-cellular;  in  the 
higher  forms  (certain  Siphonese)  the  organs  of  vegetation  attain 
differentiation  into  stem  and  leaf.  The  asexual  reproduction  takes 
place  in  various  ways ;  the  sexual  reproduction  is  effected  by  con- 
jugation of  motile  gametes,  or  by  oogamous  fertilisation.  The 


PEOTOCOCCOIDE^}.  47 

swarm-cells  (zoospores,  gametes,  and  spermatozoids)  are  con- 
structed symetrically,  and  have  true  protoplasmic  cilia,  these 
generally  being  attached  to  the  front  end  of  the  swarm-cells. 
Most  of  these  Algae  live  in  water  (fresh  or  salt);  some  are  found 
upon  damp  soil,  stones,  or  tree- stems,  and  some  live  enclosed  in 
other  plants. 

The  Class  is  divided  into  three  families  : — 

1.  PROTOCOCCOIDE^  :       Volvocaceae,      Tetrasporacese,      Chloro- 
sphaeracese,  Pleurococcaceae,  Protococcaceae,  Hydrodictyaceae. 

2.  CONFERVOIDE.ZE  :      Ulvaceae,    Ulothricaceae,     Chaetophoraceae, 
Mycoideaceae,    Cylindrocapsaceae,    GEdogoniaceae,     Coleochsetacese,. 
Cladophoraceae,  Gomontiaceae,  Sphaeropleaceae. 

3.  SIPHONED  :  Botrydiaceae,  Bryopsidaceaa,  Derbesiaceae,  Yau- 
cheriaceae,  Phyllosiphonaceae,  Caulerpaceae,  Codiaceae,  Valoniaceae,. 
Dasycladaceae. 

Family  1.     Protococcoidese. 

The  AlgaB  which  belong  to  this  group  are  nni-  or  multi-cellular 
with  the  cells  more  or  less  firmly  connected,  sometimes  in  a. 
definite,  sometimes  in  an  indefinite  form  (Fig.  47).  Colonies  are 
formed  either  by  division  or  by  small  unicellular  individuals  be- 
coming united  in  a  definite  manner  ;  the  colonies  formed  in  this 
latter  way  are  termed  Ccvnobia.  Apical  cells  and  branching  are 
absent.  Multiplication  by  division  ;  asexual  reproduction  by  zoo- 
spores,  rarely  by  akinetes.  Sexual  reproduction  may  be  wanting, 
or  it  takes  place  by  isogamous,  rarely  by  oogamous  fertilisation. 

Some  are  attached  by  means  of  a  stalk  to  other  objects  (Chara- 
cium,  Fig.  49),  others  occur  as  "  Endophytes  "  in  the  tissues  of 
certain  Mosses  or  Phanerogams,  e.g.  Chlorochytrium  lemnce,  in  Lemna 
trisulca  ;  Endosphcera.,  in  the  leaves  of  Potamogeton,  Mentha  aquatica, 
and  Peplis  portula ;  Phyllobium,  in  the  leaves  of  Lysimachia  num- 
mularia,  Ajuga,  Chlora,  and  species  of  Grasses ;  Scotinosphcera  in 
the  leaves  of  Hypnum  and  Lemna  trisulca ;  the  majority,  however, 
live  free  in  water  and  in  damp  places.  Many  species  which  were 
formerly  considered  to  belong  to  this  family  have  been  proved  to 
be  higher  Algae  in  stages  of  development. 

Order  1.  Volvocaceae.  The  individuals  in  this  order  are  either 
uni-  or  multi-cellular,  and  during  the  essential  part  of  their  life 
are  free-swimming  organisms.  They  are  generally  encased  in  a 

mucilaginous  envelope,  through  which  2-6  cilia  project  from  every 
\v.  u  E 


48  PROTOCOCCOIDE^;. 

coll.  The  vegetative  reproduction  takes  place  by  the  division  of 
all,  or  a  few,  of  the  cells  of  the  individual  ;  in  some  a  palmella-stage 
is  found  in  addition.  The  sexual  reproduction  takes  place  by 
isogamous  or  oogamous  fertilisation. 

The  Volvocaceae  may  be  considered  to  include  the  original  forms  of  the  Chloro- 
phyceae.  because,  among  other  reasons,  the  motile  stage  is  here  the  most  promi- 
nent; they  also  form  the  connecting  link  between  the  animal  Flagellata,  and 
forms  intermediate  to  the  Syngeneticte  may  perhaps  be  found  amongst  them. 
Three  series  of  green  Alga?  may  be  supposed  to  have  taken  their  origin  from  the 
Volvocaceae  :  CONJUGATED  (Desmidiacea)  which  have  lost  the  swarming  stage,  but 
whose  conjugation  is  the  nearest  to  the  fertilisation  in  Chlamydomonas  pulvis- 
ciilus  :  the  PROTOCOCCACE.E  in  which  the  vegetative  divisions  have  disappeared, 
while  the  swarming  stage  continues  to  be  present,  though  of  shorter  duration  ; 
and  TETRASPORACE.S:,  in  which  the  vegetative  divisions  are  more  prominent, 
whilst  the  swarming  stage  is  less  so. 


A.  UNICELLULAR  INDIVIDUALS.  The  principle  genera  are  : 
mydomonas,  Sphcerella,  Phacotus.  —  Sphcerella  nivalis  is  the  Alga 
which  produces  the  phenomenon  of  "  Red  Snow,"  well  known  en 
high  mountains  and  on  ice  and  snow  fields  in  the  polar  regions. 
The  red  colouring  matter  which  appears  in  this  and  other  green 
Algas,  especially  in  the  resting  cells,  is  produced  by  the  alteration 
of  the  chlorophyll. 

Phacotus  lenticularis  has  an  outer  covering  incrusted  with  lime, 
Avhich,  at  death,  or  after  division,  opens  out  into  two  halves. 
Species  may  be  found  among  Chlamydomonas,  in  which  conjuga- 
tion takes  place  between  gametes  of  similar  size  without  cell-wall, 

but  in  C.  pulvisculus  conjugation 
takes  place  between  male  and 
female  aplanogametes  which  are 
surrounded  by  a  mucilaginous 
envelope. 

B.    MULTICELLULAR  INDIVIDUALS, 

The  most  important  genera  are 
Gonium,  Stephanosphcera,  Pandor- 
ina,  Eudorina,  Volvox.  —  Gonium 
has  4  or  16  cells  arranged  in  a 

definite    pattern    in    a  flat    plate 
Fig.  41—  Gonium  pectorale.  /Ti1.         .  ..          ^       7      .         /-ri.        .  „, 

(Fig.  44).      Pandonna   (Fig.  45), 

has  16  cells  arranged  in  a  sphere  (Fig.  45  A).  The  vegetative 
reproduction  takes  place  in  this  way  :  each  cell,  after  having 
rounded  off,  and  after  the  withdrawal  of  the  cilia,  divides  itself 


PEOTOCOCCOIDE^. 


49 


into  16  new  ones  (Fig.  45  B),  each  forming  a  new  individual, 
which  soon  grows  to  the  size  of  the  mother-individual.  It  was 
in  this  Alga  that  the  conjugation  of  self-motile  gametes  was 
first  discovered  by  Pringsheim,  1869.  When  conjugation  is 
about  to  take  place,  each  cell  divides  into  sixteen,  as  in  vegeta- 
tive reproduction,  but  the  16  x  16  cells  all  separate  from  one 
another  (Fig.  45  (7,  female  gametes,  and  _D,  male  gametes),  and 


FIG.  45. — Pandorina  morum. 


swarm  solitarily  in  the  water.  The  male  are,  most  frequently, 
smaller  than  the  female,  but  otherwise  they  are  exactly  alike ; 
they  are  more  or  less  pear-shaped,  with  a  colourless  anterior  end, 
2  cilia,  a  red  "  eye-spot,"  etc.  After  swarming  for  some  time  they 
approach  each  other,  two  and  two,  generally  a  large  and  a  smaller 
one,  and  come  into  contact  at  their  colourless  end;  in  a  few 
moments  they  coalesce  and  become  one  cell  (Fig.  45  E,  .F),  this 


50  PROTOCOCCOIDEjE. 

has  at  first  a  large  colourless  anterior  end,  4  cilia,  and  2  "  eye- 
spots"  (Fig.  45  G),  but  these  soon  disappear  and  the  cell  becomes 
uniformly  dark-green  and  spherical,  and  surrounds  itself  with  a 
thick  cell-wall,  losing  at  the  same  time  its  power  of  motion:  the 
zygote  (Fig.  45  H)  is  formed,  and  becomes  later  on  a  deep  red 
colour.  On  the  germination  of  the  zygote,  the  protoplasmic  cell- 
contents  burst  open  the  wall  (Fig.  45  J),  and  emerge  as  a  large 
swarmspore  (Fig.  45  K]  which  divides  into  16  cells,  and  the  first 
small  individual  is  formed  (Fig.  45  L,  M). 

Eudorina  is  like  Pandorina  in  structure,  but  stands  somewhat 


FIG.  4G.—Volvox  globator,  sexual  inili vicinal :    a  antheridia  which  have  formed  spermato- 
zoids ;    6  oogonia. 

higher,  since  the  contrast  between  the  conjugating  sexual  cells  is- 
greater,  the  female  one  being  a  motionless  oosphere. 

The  highest  stage  of  development  is  found  in  Volvox  (Fig.  46). 
The  cells  are  here  arranged  on  the  circumference  of  a  sphere,  and 
enclose  a  cavity  filled  with  mucilage.  The  number  of  these  cells 
may  vary  from  200-22,000,  of  which  the  majority  are  vegetative 
and  not  reproductive,  but  some  become  large,  motionless  oospheres 
(Fig.  46  6) ;  others,  which  may  appear  as  solitary  individuals,, 
divide  and  form  disc- shaped  masses  of  from  8-256  small  spermato- 


FROTOCOCCOIDE.E. 


51 


zoids  (Fig.  46  a).  After  the  oosphere  has  been  fertilised  by  these, 
the  oospore  surrounds  itself  by  a  thick,  sometimes  thorny  cell- 
wall,  and  on  germination  becomes  a  new  individual  of  few  cells. 
A  few  cells  conspicuous  by  their  larger  size  may  be  found  (1-9,  but 
generally  8)  in  certain  individuals,  and  these  provide  the  vegeta- 
tive reproduction,  each  forming  by  division  a  new  individual. 

Order  2.  Tetrasporaceae  reproduce  both  by  vegetative  divisions  and 
swarmspores,  some  have  also  gamete-conjugation.  The  principal  genera  are : 
Tetraspora,  Apiocystis,  Dactylococcus,  Dictyosphcerium,  Chloranyium. 

Order  3.     Chlorosphaeracese.     Chlorosphcera. 

Order  4.  Pleurococcaceae.  In  this  order  the  swarm-stages 
and  sexual  reproduction  are  entirely  absent.  Vegetative  repro- 
duction by  division.  The  principal  genera  are  :  Pleurococcus  (Fig. 
47),  Scenedesmus  (Fig.  48),  Raphidium,  Oocystis,  Schizoclilamys, 
Crucigenia,  Selenastrum. — Pleurococcus  vulgaris  (Fig.  47)  is  one  of 
the  most  common  Algae  throughout  the  world,  occurring  as  green 
coverings  on  tree-stems,  and  damp  walls,  and  it  is  one  of  the  most 
common  lichen-gonidia. 


FIG.  47. — Pieurococcas  vulgaris. 


FIG.  48  —  S. 


uiiadricauda. 


Order  5.  Protococcacese.  The  cells  are  motionless,  free  or 
affixed  on  a  stalk  (e.g.  Characium,  Fig.  49),  either  separate  or 
loosely  bound  to  one  another ;  they  never  form  multicellular 
individuals.  Multiplication  by  division  is 
learly  always  wanting.  Reproduction  takes 
)lace  by  swarmspores,  which  have  1  or  2  cilia, 
ind  sexual  reproduction  in  some  by  gamete- 
mjugation.  The  principal  genera  are  :  Ghlo- 
coccum,  Chlorochytrium,  Ghlorocystis,  Scotino- 
sphcera,  Endosphcera,  Phyllobitim,  Characium, 
Ophiocytium,  Sciadium. 

Order  6.  Hydrodictyaceae.  The  indi- 
viduals are  unicellular  but  several  unite  after 
the  zoospore-stage  into  definitely  formed 
families  (coenobia).  Ordinary  vegetative  division  is  wanting,  but 


—  Characium 
strictum.  A  The  cell-con- 
tents have  divided  into 
many  swarmspores.  B 
Swarmspores  escaping. 


52 


PROTOCOCCOIDEJ;. 


asexual  reproduction  takes  place  by  zoospores  (or  by  motionless 
cells  without  cilia),  which  unite  and  form  a  family  similar  to  the 
mother-family,  inside  the  mother-cell,  or  in  a  mucilaginous  enve- 
lope. Where  sexual  reproduction  is  found  it  takes  place  by 
gamete-conjugation.  The  principal  genera  are  :  Pediastrum  (Fig. 
50),  Ccelastrum,  Hydrodictyon  (Fig.  51). 

The  coenobium  of  Hydrodictyon  reticulatum  (Water-net)  is 
formed  of  a  large  number  of  cells  which  are  cylindrical,  and 
attached  to  one  another  by  the  ends  (Fig.  51).  The  asexual 
reproduction  takes  place  by  zoospores,  which  are  formed  in  large 
numbers  (7,000-20,000)  in  each  mother-cell,  within  which  they 
move  about  for  a  time,  and  then  come  to  rest  and  arrange  them- 
selves into  a  new  net  (Fig.  51  A)  which  is  set  free  by  the  dissolu- 
tion of  the  wall  of  the  mother-cell,  grows,  and  becomes  a  new 


FIG.  50.— Pediastrum  a 


FIG.  51.  —  Hydrodictyon  retici.latum. 
A  A  cell  where  the  zoospores  are  on  the 
point  of  arranging  themselves  to  form  a 
net.  B  A  cell  with  gametes  swarming  out. 


cceriobium.  The  sexual  reproduction  takes  place  by  gamete-conju- 
gation. The  gametes  are  formed  in  the  same  manner  as  the  zoo- 
spores,  but  in  larger  numbers  (30,000-100,000),  and  swarm  out  of 
the  mother-cell  (Fig.  51  B}.  The  zygote  forms,  on  germination, 
2-5  large  zoospores,  each  with  one  or  two  cilia,  these  generally 
swarm  about  for  a  time,  and  after  a  period  of  rest  become  irregular 
thorny  bodies  (polyhedra)  ;  their  contents  again  divide  into  zoo- 
spores,  the  thorny  external  coating  of  the  polyhedra  is  cast  off, 
and  the  zoospores,  surrounded  by  the  dilated  internal  coating, 
unite  to  form  a  small  family,  which  produces  several  others  in  the 
manner  described. 


CONFERVOIDE^;. 


53 


Family  2.     Confervoideae. 

The  individuals  are  always  multicellular,  the  cells  firmly 
bound  together  and  united  into  unbranched  or  branched  filaments, 
expansions,  or  masses  of  cells  which  grow  by  intercallary  divisions 
or  have  apical  growth.  In  the  first  seven  orders  the  cells  are 
uninuclear,  but  the  cells  of  the  remaining  three  orders  contain 
several  nuclei.  Asexual  reproduction  by  zoospores,  akinetes  or 
aplanosporos.  Sexual  reproduction  by  isogamous  or  oogamous 
fertilisation. 

The  Confervoidese,  through  the  Ulvaceae,  are  connected  with  the  Tetra- 
sporacete,  and  from  the  Coleochcctacece,  which  is  the  most  highly  developed 
order,  there  are  the  best  reasons  for  supposing  that  the  Mosses  have  taken 
their  origin.  The  Cladophoracece  show  the  nearest  approach  to  the  Siplionece. 

Order  1.  Ulvaceae.  The  thallus  consists  of  one  or  two  layers 
of  parenchymatous  cells,  connected  together  to  form  either  a  flat 
membrane  (Monostroma,  Ulva)  or  a  hollow  tube  (Enteromorpha), 
and  may  be  either  simple,  lobed,  or  branched.  Reproduction 
takes  place  by  detached  portions  of  the  thallus ;  or  asexually  by 
zoospores  or  akinetes.  Gramete- conjugation  is  known  to  take 
place  in  some  members  of  this  order,  the  zygote  germinating 
without  any  resting-stage.  The  majority  are  found  in  salt  or 
brackish  water. 

Order  2.  Ulothricaceae.  The  thallus  consists  normally  of  a 
simple  unbranched  filament  (sometimes  a  small  expansion  con- 


PIG.  52.— UlotUrix  zonata:  a  portion  of  a  filament  with  zoospores,  which  are  formed 
two  in  each  cell  (zoosporangium);  the  dark  spots  are  the  red  "eye-spots";  1,  2,  3,  4, 
lenote  successive  stages  in  the  development  of  the  zoospores ;  b  a  single  zoospore,  v 
le  pulsating  vacuole;  c  portion  of  a  filament  with  gametes,  sixteen  are  produced  in  each 
Kametanginm  ;  d  free  gametes,  solitary  or  in  the  act  of  conjugation;  e  the  conjugation  is 
completed,  and  the  formed  zygote  has  assumed  the  resting-stage. 

listing  of  one  layer  of  cells  is  formed,  as  in  Schizomeris  and 
Prasiola  which  were  formerly  described  as  separate  genera). 
Asexual  reproduction  takes  place  by  means  of  zoospores  (with 


54  CONFERVOIDE5). 

1,  2,  or  4  cilia),  akinetes  or  aplanospores ;  tlie  last  named  may 
germinate  immediately,  or  only  after  a  period  of  rest.  Sexual  re- 
production takes  place  by  the  conjugation  of  gametes  of  about  the 
same  size,  each  having  two  cilia  (Fig.  52  d).  The  zygote  of 
Ulothrix,  on  germination,  produces  a  brood  of  zoospores  which 
swarm  for  a  time  and  then  elongate  to  become  t/7o£/m'#-filaments 
(alternation  of  generations).  The  gametes  may  also  germinate 
without  conjugation  in  the  same  manner  as  the  zoospores.  The 
principal  genera  are :  Ulothrix,  Hormidium,  Conferva,  Micro- 
spora. —  Ulothrix  zonata  is  very  common  in  running  fresh  water. 
Nearly  all  the  species  of  Hormidium  occur  on  damp  soil,  tree- 
stems  and  stones. 

Order  3.  Chsetophoraceae.  The  thallus  consists  of  a  single, 
branched,  erect  or  creeping  filament  of  cells,  often  surrounded  by 
mucilage.  The  cells  have  only  one  nucleus.  Asexual  reproduc- 
tion by  zoospores  with  2  or  4  cilia,  by  akinetes,  or  aplanospores., 
In  many,  conjugation  between  gametes  with  2  cilia  may  be  found. 
They  approach  on  one  side,  Ulothricaceae,  and  on  the  other,  My- 
coideaceae.  The  principal  genera  are  :  Stigeoclonium,  Draparnaldia, 
Chwtophora,  Entoderma,  Aphanochcete,  Herposteiron,  Phceothamnion, 
Ohlorotyliuin,  Trichophilus,  Gongrosira,  Trentepohlia.  Most  of  the 
species  of  Trentepohlia  are  coloured  red  by  the  presence  of  a  red 
colouring  material,  which  occurs  in  addition  to  the  chlorophyll. 
They  are  aerial  Algse  which  live  on  stones  (T.  jolithus,  "violet 
stone,"  so  named  on  account  of  its  violet-like  odour  in  rainy 
weather),  on  bark  and  old  wood  (T.  umbrwa),  or  on  damp  rocks 
(T.  aurea}.  Trichophilus  welckeri  lives  in  the  hair  of  Bradypus. 

Order  4.  Mycoideaceae.  The  thallus  is  discoid,  consisting  of  one  or  more 
cell-layers,  and  is  always  attached.  Asexual  reproduction  by  zoospores  with  2 
or  4  cilia.  Sexual  reproduction  in  some  species  by  the  conjugation  of  gametes 
with  2  cilia.  This  order  forms  the  connecting  link  between  CJicctophoraceae 
and  Coleochcetacea.  The  species  occur  in  fresh  water  (Chtetopeltis)  as  well  as 
in  salt  (Pringsheimia),  on  the  carapace  of  tortoises  (Dermatophyton=Epiclem- 
niydia),  or  endophytic  between  the  cuticle  and  the  epidermal  cells  of  the 
leaves  of  tropical  plants,  destroying  the  leaf-tissue  (Mycoidea). 

Order  5.  Cylindrocapsaceae.  The  thallus  consists  of  a 
simple  (rarely,  in  parts,  formed  of  many  rows)  unbranched 
filament,  attached  in  the  young  condition,  which  has  short  cells 
with  a.  single  nucleus,  and  is  enveloped  in  a  thick  envelope  with 
a  laminated  structure.  Asexual  reproduction  by  zoospores  with 
2  cilia,  which  are  formed  1,  2,  or  4  in  each  vegetative  cell.  The 


CONFEKVOIDE2E. 


55 


antheridia  are  produced  by  a  single  cell,  or  a  group  of  cells,  in  a 
filament,  dividing  several  times  without  increasing  in  size.  Two 
egg-shaped  spermatozoids,  each  with  2  cilia  (Fig.  53  D),  are 
formed  in  each  antheridium,  and  escape  through  an  'aperture  in 
the  side ;  in  the  first  stages  they  are  enclosed  in  a  bladder-like 
membrane  (Fig. 
53  B,  0).  Other 
cells  of  the  fila- 
ment swell  out 
and  form  oogonia 
(Fig.  53  A),  which 
resemble  those 
of  (Edogonium. 
After  fertilisa- 
tion, the  oospore 
surrounds  itself 
with  a  thick  wall, 
and  assumes  a 
reddish  colour. 
The  germination 
is  unknown. 
The  unfertilised 
oospheres  remain 
-  green, divide  often 
into  2-4  daughter- 
cells,  and  grow 
into  new  fila- 
ments. 

This    order, 
which     'only     in- 
cludes   one    genus,    Cylindrocapsa,  forms  the  connecting  link  be- 
tween   Ulothricacece  and  CEdogoniacece.     The  few  species  (4)  occur 
only  in  fresh  water. 

Order  6.  CEdogoniaceae.  The  thallus  consists  of  branched 
(Bulbochcete)  or  unbranched  ((Edogoni.um)  filaments,  attached  in 
the  early  stages.  The  cells  may  be  longer  or  shorter,  and  have 
one  nucleus.  Asexual  reproduction  by  zoospores,  which  have  a 
chaplet  of  cilia  round  the  base  of  the  colourless  end  (Fig.  6  a). 
Sexual  reproduction  takes  place  by  oogamous  fertilisation.  On 
the  germination  of  the  oospore,  4  zoospores  are  formed  (Fig.  54  F). 
They  occur  only  in  fresh  or  slightly  brackish  water.  The  division 


FIG.  53. — Cylindrocapsa  involuta.  A  Oo<*onium  with  oosphere 
(o)  surrounded  by  spermatozoids  (s).  S  Two  antheridia,  each 
with  two  spermatozoids.  C-  Spermatozoids  surrounded  by 
their  bladder-like  membrane.  D  Free  spermatozoid. 


56 


CONFERVOIDES. 


of  the  cells  takes  place  in  quite  a  peculiar  and  unusual  manner. 
At  the  upper  end  of  the  cell  which  is  about  to  divide,  a  ring-shaped 
thickening  of  soft  cellulose  is  formed  transversely  round  the  wall ; 
the  cell-nucleus  of  the  mother-cell  and  the  protoplasm  then  divide 
by  a  transverse  wall  into  two  portions  of  similar  size,  and  the 

cell-wall  bursts  trans- 
versely along  the  cen- 
tral line  of  the  thick- 
ened ring.  The  cell- 
wall  thus  divides  into 
two  parts — the  upper 
one  short,  the  "cap," 
and  the  lower  one 
much  longer,  the 
"sheath."  The  por- 
tions of  the  original 


J 


cell-wall  now  separate 
from  each  other,  the 
cellulose  ring  extend- 


§ 
K\  \/  /12IKI      an   additional    length 
[mP)  ^p\  I  JBH      of    cell-  wall    between 

them.  The  cap  and 
sheath  will  project  a 
little  in  front  of  the 
piece  thus  inserted. 
The  dividing-  wall  be- 
tween the  two  new 
cells  is  formed  near  to 
the  uppermost  edge 
of  the  sheath,  and 
gradually  becomes 
thicker  and  firmer. 
The  inserted  piece  of 
wall  forms  the  larger 
part  of  the  wall  of  the 
upper  cell  :  the  re- 
mainder is  formed  by 
the  cap.  This  mode 
of  division  is  repeated  exactly  in  the  same  way,  and  new  caps  are 
formed  close  below  the  first  one,  one  for  every  division. 


FIG.  5i.— A  (Edogonium  ciliatum.  A  Female  plant  with 
three  oogonia  (og)  and  dwarf-males  (m).  B  An  oogonium 
with  spermatozoid  (z)  seen  entering  the  oosphere  (o)  hav- 
ing passed  through  an  aperture  near  the  summit  of  the 
oogonium;  m  dwarf-male.  C  Ripe  oospore.  D  (Edogo- 
nium gemelHparnm.  F  Portion  of  a  male  fiiamm.t  from 
which  spertnatozoids  (z)  are  emerging.  E  Portion  of 
filament  of  Bulbochcete ;  tlie  upper  oogonium  still  en- 
closes the  oospore,  in  the  central  one  the  oospore  is  es- 
caping while  the  lower  one  is  empty.  FFour  zoospores 
developed  from  an  oospore.  G  Zcospore  germinating. 


CONFERVOIDEJ;. 


57 


Fertilisation  takes  place  in  the  following  way.  The  oogonium 
is  a  large  ellipsoidal,  swollen  cell  (og,  in  Fig.  54  A),  whose  contents 
are  rounded  off  into  an  oosphere  with  a  colourless  receptive-spot 
(see  B)  ;  an  aperture  is  formed  in  the  wall  of  the  oogonium,  through- 
which  the  spermatozoids  are  enabled  to  enter  (B).  The  sper- 
matozoids  are  produced  either  directly,  as  in  D  (in  pairs),  in  basal 
cells  of  the  filament,  or  indirectly.  In  the  latter  case  a  swarm- 
spore  (androspore)  is  formed  which  comes  to  rest,  attaches  itself 


FIG.  55  —  Coleochcete  pulvinata.  A  A  portion  of  a  thallus  with  organs  of  reproduction; 
a  oogonium  before,  b  after  fertilisation;  can  antheriditim,  closed;  ci  open,  with  emerg- 
ing spermatozoid.  B  Ripe  oogonium,  with  envelope.  G  Germination  of  the  oospore.  D 
Zoospore.  E  Spermatozoid. 

to  an  oogonium,  germinates,  and  gives  rise  to  a  filament  of  a  very 
fe\v  cells — dwarf-male  (A,  B,  m).  The  spermatozoids  are  formed 
in  the  upper  cell  of  the  dwarf-male  (m),  and  are  set  free  by  the 
summit  of  the  antheridium  lifting  off  like  a  lid.  On  the  germina- 
tion of  the  oospore  ((7),  which  takes  place  in  the  following  spring, 
4  zoospores  are  produced  (F)  (i.e.  the  sexual  generation) ;  these 
swarm  about  for  a  time,  and  ultimately  grow  into  new  filaments. 

Order  7.  Coleochaetaceae.  The  thallus  is  always  attached, 
and  of  a  disc-  or  cushion-shape,  formed  by  the  dichotomous 
branching  of  filaments  of  cells  united  in  a  pseudo-parenchy- 
matous  manner.  Each  cell  has  only  one  nucleus.  Asexual  re- 
production by  zoospores  with  2  cilia  (Fig.  55  D),  which  may 
arise  in  all  the  cells.  Sexual  reproduction  by  oogamous  fertili- 
sation. The  spermatozoids  resemble  the  swarmspores,  but  are 


v>8 

smaller  (E),  and  originate  singly  (in  the  species  figured)  in  small 
conical  cells  (c,  d  in  A}.  The  oogonia  are  developed  at  the 
extremities  of  certain  branches  :  they  are  bottle-shaped  cells  with 
very  long  arid  thin  necks  (trichogyne) ,  open  at  the  end  (a  in  A)  • 
at  the  base  of  each  oogonium  is  a  spherical  oosphere.  The  sper- 
matozoids  reach  the  oosphere  through  the  trichogyne,  or  through 
an  aperture  in  the  wall  when  the  trichogyne  is  absent,  and  fertili- 
sation having  taken  place,  the  oogonium  becomes  surrounded  by  a 
cell-layer  (envelope),  which  grows  out  from  the  cells  near  its  base 
(6  in  A),  and  in  this  way  a  kind  of  fruit  is  formed  (B)  (spermocarp, 
cystocarp) . 

The  oospore,  next  spring,  divides  and  forms  a  parenchymatous 
tissue  (homologous  with  the  Moss-sporophyte) ;  this  bursts  open 
the  envelope  (0),  and  a  zoospore  (homologous  with  the  spores  of 
the  Moss-capsule)  arises  in  each  of  the  cells,  and  produces  a  new 
Coleochcete.  We  have  then,  in  this  case,  a  still  more  distinct 
alternation  of  generations  than  in  (Edogonium.  Only  one  genus, 
Coleochcete,  is  known,  but  it  contains  several  species,  all  living  in 
fresh  water. 

Order  8.  Cladophoracese.  This  order  is  probably  derived 
from  the  Ulothricaceae.  The  thallus  consists  of  a  single,  un- 
branched  or  branched  filament,  generally  with  an  apical  cell. 
The  cells  have  each  2  "or  more  nuclei.  Asexual  reproduction  by 
zoospores  with  2  or  4  cilia,  and  by  akinetes.  Conjugation  of 
gametes  with  2  cilia  is  found  in  some  genera.  They  occur  in  salt 
as  well  as  in  fresk  water.  The  principal  genera  are :  Urospora, 
Chcetomo^ha,  llhizoclonium,  Cladophora ;  of  the  last  named  genus 
the  species  C.  lanosa  and  C.  rupestris  are  common  in  salt  water ; 
C.  fracta  and  C.  glomerata  in  fresh  water. 

Order  9.  Gomontiaceae.  Gomontia  polyrrhiza,  the  only  species  hitherto 
known,  is  found  on  old  calcareous  shells  of  certain  salt  water  Molluscs. 

Order  10.  Sphaeropleacese.  The  thallus  consists  of  free,  un- 
branched  filaments,  with  very  elongated  multin^clear  cells.  The 
vegetative  cells  form  no  zoospores.  Sexual  reproduction  by 
oogamous  fertilisation  (see  page  13,  Fig.  10  B).  The  oospore  has 
a  thick  wall  (Fig.  10  D)  studded  with  warts,  and  assumes 
colour  resembling  red  lead.  It  germinates  only  in  the  following 
spring,  and  produces  1-8  zoospores,  each  with  2  cilia  (Fig.  10  J57), 
which  grow  into  new  filaments.  Only  one  species,  Sphceroplea 
annulina,  is  known. 


SIPHONEJ1.  5&> 

Family  3.     Siphonese. 

The  thallus  has  apical  growth,  and  in  the  vegetative  condition, 
consists  generally  of  one  single  (in  the  Valoniaceae  most  frequently 
of  more)  multinuclear  cell,  which  may  be  much  branched,  and 
whose  separate  parts  in  the  higher  forms  (e.g.  Bryopavf,  Fig.  57 ;: 
Caulerpa,  Fig.  59,  etc.)  may  be  differentiated  to  perform  the 
various  physiological  functions  (as  root,  stem  and  leaf).  Vegeta- 
tive multiplication  by  detached  portions  of  the  thallus  (gemmae)  ; 
asexual  reproduction  by  zoospores,  akinetes,  or  aplanospores. 
Sexual  reproduction  by  gamete-conjugation,  rarely  by  oogamous 
fertilisation.  The  zygote  or  oospore  germinates  as  a  rule  without 
any  resting-stage. 

Most  of   the   Siphoneas   occur  in  salt  water  or  on  damp   soil. 


FIG.  56.—  Botrijdium  granulatum  •.  a  an  entire  plant  forming  swa-rmspores ;  b  swarm- 
spores  ;  c  an  individual  with  gametangia  ;  d,  gamete;  e,  f,  g  conjugation;  h  zygote 
seen  from  above  ;  i  the  same  in  a  lateral  view. 

Many  (e.g.  Dasycladacew)  are  very  much  incrusted  with  lime,  and 
occur,  in  the  fossilized  condition,  in  the  deposits  from  the  Cretaceous 
period  to  the  present  time.  The  Siphoneas  are  connected  by  their 
lowest  forms  (BotrydiaceoK  or  Valonid]  with  the  Protococcaceae,  but 
show  also,  through  the  Valoniaceae,  points  of  relationship  to  the 
Cladophoracece. 

Order  1.  Botrydiaceae.  The  thallus  in  the  vegetative  condi- 
tion is  unicellular,  club-shaped,  with  a  small  single  (Codiolum)  or 
repeatedly  dichotomously  branched  system  of  colourless  rhizoids 
(Botrydium,  Fig.  56  a),  by  which  it  is  attached  to  objects  immersed 
in  salt  water  (Codiolum)  or  to  damp  clay  soil  (Botrydium). 
Asexual  reproduction  by  zoospores  with  one  (Botrydium}  or  two. 


60 


SIPHONED. 


cilia,  and  by  apla.nospores.  The  sexual  reproduction  is  only 
known  in  Botrydium,  and  takes  place  in  the  following  manner : 
in  the  part  of  the  thallus  which  is  ahove  ground  arid  in  an  active 
vegetative  condition,  several  round  cells  (Fig.  56  c)  are  formed, 
which  may  be  green  or  red  according  as  they  grow  under  water, 
or  exposed  to  the  strong  light  of  the  sun..  These  cells  must  be 
considered  as  "  gametangia  "  as  they  produce  many  gametes  (d) 
provided  with  two  cilia.  The  zygote  (h,  i)  formed  by  the  conju- 
gation (e,  /,  </)  may  either  germinate  immediately,  or  become 
thick-walled  resting-cell  of  an  irregular  angular  form. 

Order  2.  Bryopsidaceae.  The  thallus  in  the  vegetative  condition  is  uni- 
cellular, and  consists  at  the 
lower  extremity  of  branched 
rhizoids.  while  the  upper 
portion  is  prolonged  into  a 
stem-like  structure  of  un- 
limited growth,  producing, 
acropetally,  benches  and 
leaf-like  structures.  The 
latter  have  limited  growth, 
and  are  separated  by  a  cross 
wall  from  the  stem,  and 
come  gametangia,  or  dro{ 
off.  The  gametes  have  twc 
FIG.  67.— Bnjnvais  plumosa.  A  the  plant,  natural  size.  cilia,  and  are  of  two  kinds  : 

B  A  portion  (enlarged)  which  shows  the  growing  point        the  femal      which  are 

•(t>),  ana  the  leaves  derived  f rom  it  in  acropetal  succes-  , 

8ion.  and    large    and    the    male, 

which     are'    of      brownish 

colour  and  smaller.     Zoospores  or  any  other  method  of  asexual  reproduction 
are  unknown.     Only  one  genus,  Eryopsis,  living  in  salt  water. 

OrderS.  Derbesiaceae.  Only  one  genus,  Derbesia,  living  in  saltwater.  The 
y.oospores,  which  are  formed  in  a  few  lateral,  swollen  zoosporangia,  possess  one 
nucleus  which  has  arisen  through  the  coalescence  of  several,  and  they  resemble 
the  zoospores  of  (Edogonium  by  having  a  circle  of  cilia  attached  at  the  base 
of  the  colourless  spot. 

Order  4.  Vaucheriaceae.  The  thallus  consists,  in  the  vegetative 
condition,  of  a  single  irregularly  or  dichotomously  branched  cell, 
without  differentiation  into  stem  or  leaf  ;  root-like  organs  of  attach- 
ment may  however  occur.  Asexual  reproduction  by  zoospores, 
which  are  formed  singly  in  the  extremity  of  a  branch  cut  off  by 
a  transverse  wall.  They  contain  many  nuclei,  and  bear  small 
cilia  situated  in  pairs,  which  give  the  appearance  of  a  fine  "  pile  " 
•covering  the  whole  or  a  great  part  of  the  surface.  Akinetes, 


SIPHONED. 


61 


aplanospores,  and  phytoamoebae  (naked  masses  of  protoplasm, 
without  cilia,  which  creep  like  an  amoeba  on  a  substratum)  may 
occur  under  certain  conditions. 

The  sexual  reproductive  organs  are  formed  on  short  lateral 
branches,  and  are  separated  from  the  vegetative  cell  (Fig.  58  A}  by 
cell-walls.  Numerous  spermatozoids,  each  with  two  cilia,  are 
developed  in  the  coiled  antheridium  (A,  ~b).  The  oogonium  is  a 
thick,  egg-shaped,  often  oblique  cell,  with  its  protoplasm  rounded 
into  an  oosphere,  which  has  a  hyaline  "  receptive-spot "  (A,  *)  im- 
mediately beneath  the  aperture  formed  in  the  wall  of  the  oogonium. 
A  slimy  mass,  which  serves  to  receive  the  spermatozoids,  is  formed 
in  some  species  in  this  aperture.  The  spermatozoids  when  liber- 
ated swim  towards  and  enter  the  oosphere,  which  then  immedia- 


A 

FIG.  53. — Vaucheria  sessilis. 
receptive  spot.    B  Oospore. 


A  Fertilisation ;   b  the  antheridia ;   a  the  oogonia  ;   a  the 


tely  surrounds  itself  with  a  thick  cell-wall.  The  mature  oospore 
(B)  contains  a  large  quantity  of  oil.  At  germination  the  outer 
cell-wall  bursts  and  a  new  plant  is  formed.  There  is  only  one 
genus,  Vaucheria,  with  species  living  in  salt  as  well  as  in  fresh 
water  and  on  damp  soil. 

Order  5.  Phyllosiphonacese  are  parasites  in  the  leaves  and  stalks  of 
Flowering-plants. 

Order  6.  Caulerpacese.  The  thallus  has  distinct  differentation 
into  root,  stem  and  leaf-like  members  (Fig.  59)  ;  it  is  unicellular. 
Within  the  cell,  strong,  branched  threads  of  cellulose  extend 
from  one  side  to  the  other  serving  as  stays  to  support  the 
thallus.  Reproduction  takes  place  by  detached  portions  of  the 
thallus ;  no  other  modes  of  reproduction  are  known.  This  order 
may  most  approximately  be  classed  with  the  Bryopsidacece.  The 
genus  Caulerpa  consists  of  more  than  seventy  species  which 
inhabit  the  tropical  seas. 

Order  7.    Codiaceae.    The  thallus  has  various  forms,  but  with- 


62 


SIPHONED. 


out  distinct  differentiation  in  stem-  or  leaf-structures,  sometimes 
(e.g.  Halimeda)  it  is  very  much  incrusted  with  lime.  In  the 
early  stages  it  is  unicellular  (later,  often  multicellular),  very 
much  branched,  with  the  branches,  at  any  rate  partly,  so  united 
or  grown  in  amongst  one  another  (Fig.  60)  that  an  apparently 
parenchymatous  cellular  body  is  formed.  Akinetes  oraplanosporos 
are  wanting;  zoospores  (or 
gametes  ?)  may  be  developed  in 
some  species,  however,  in  special 
swollen  sporangia.  Fertilisation 
similar  to  that  in  Bryopsis  occurs 
perhaps  in  G odium.  They  are 
all  salt  water  forms. 

Order  8.  Valoniaceae.  The  thal- 
lus  is  generally  multicellular,  without 
differentation  into  stem-  or  leaf-struc- 
tures, but  the  cells  are  sometimes 
united  together  and  form  a  leaf-like 
reticulate  expansion  (e.g.  Anadyomene). 
Zoospores  are  known  in  some,  and 
they  are  then  formed  directly  in  the 
vegetative  cells.  In  others  (e.g. 
Valdliia),  a  mass  of  protoplasm,  which 
7i)  ay  be  separated  through  the  damag- 
ing of  a  cell,  can  surround  itself  with 


Fia.  59.— Caulerpa  prolifera  (natural  size). 

a  cell-wall,  and  grow  into  a  new  plant.  No  other  modes  of  reproduction  are 
known.  The  most  important  genera  are  :  Valonia,  Siphonocladus,  Chamcedoris, 
Struvea,  Mtcrodictyon,  Anadyomene.  They  are  all  salt  water  forms. 

As  already  pointed  out,  the  Valoniacece  occupy  a  somewhat  central  position 
among  the  Siphoneae,  and  present  points  of  similarity  and  contrast  with  the 
Botrydiacece  and  the  Bryopsidacece  through  Valonia,  with  the  Dasycladacece 
through  Chamcedoris,  and  also  with  the  Cladophoraceee  through  Siphonocladus, 
and  Struvea. 


SIPHONED. 


.63 


Order  9.  Dasycladaceae.  The  thallus  consists  of  an  axile 
longitudinal  cell,  destitute  of  transverse  walls,  attached  at  the 
base  by  root-like  organs  of  attachment,  and  producing  acropetally 
whorls  of  united,  single  or  branched,  leaf-like  structures  with 
limited  growth.  Asexual  reproduction  is  wanting.  Sexual  re- 
production by  conjugation  of  gametes  which  arise  in  separate, 
fertile  leaves,  either  directly  or  from  aplanospores,  which  develope 
into  gametangia.  The  principal  genera  are  :  Acetabularia,  Dasy- 
cladus,  Neomeris,  Cymopolia.  All  marine. 

The  curiously  shaped 
Acetabularia  mediterra- 
nea  grows  gregariously 
on  limestone  rocks,  and 
shells  of  mussels  in  the 
Mediterranean  ;  it  re- 
sembles a  minute  um- 
brella with  a  small  stem, 
sometimes  as  much  as 
nine  centimetres  in 
height,  and  a  shade 
which  may  be  more  than 
one  centimetre  in  di- 
ameter. The  cell-mem- 
brane is  thick,  and 
incrusted  with  carbonate 
and  oxalate  of  lime. 
Only  the  lower,  root- 
like  part  of  the  thallus, 
which  penetrates  the 
calcareous  substratum- 
survives  the  winter,  and 
may  grow  up  into  a  new 

plant.  The  sterile  leaves,  which  drop  off  early,  are  dichoto- 
mously  branched  and  formed  of  cylindrical  cells  separated 
from  each  other  by  cross-walls,  but  they  are  not  grown  to- 
gether. The  shade  is  formed  by  a  circle  of  70-100  club-shaped 
rays  (fertile  leaves)  grown  together,  in  each  ray  40-80  aplano- 
spores are  formed,  which  become  liberated  at  the  breaking  of  the 
shade,  and  later  on  arc  changed  to  gametangia  (compare  Botry- 
dium)  which  open  by  a  lid  and  allow  a  large  number  of  egg- 
shaped  gametes  with  two  cilia  to  escape.  Gametes  from  various> 
w.  B.  F 


FIG.  60. — Halimeda  opuntia.    Plant  (natural  size), 
B  Part  of  a  longitudinal  section. 


6:1  CHAKACEJ3. 

gametangia  conjugate  with  one  another;  the  product  of  the  conju- 
gation swarms  about  for  some  time,  rounds  off,  and  then  surrounds 
itself  with  a  cell-wall.  The  zygote  germinates  after  a  period  of 
rest  and  then  produces  a  sexual  plant.  The  aplanospores  (game- 
tangia) thus  represent  the  sexual  generation. 

Class  7.    Characeae. 

The  thallus  has  a  stem  with  nodes  and  internodes ;  and  whorls 
of  leaves,  on  which  may  be  developed  the  antheridia  and  oogonia, 
are  borne  at  the  nodes.  Vegetative  reproduction  by  bulbils  and 
accessory  shoots.  Zoospores  are  wanting.  The  antheridia  are 
spherical,  and  contain  a  number  of  filaments  in  which  the  spirally 
coiled  spermatozoids,  each  with  two  cilia,  are  formed.  The  oogonium 
is  situated  terminally,  and  is  at  first  naked,  but  becomes  later 
on  surrounded  by  an  investment,  and  forms  after  fertilisation  the 
so-called  "  fruit."  The  oospore,  after  a  period  of  rest,  germinates 
by  producing  a  "  proembryo,"  from  which  the  young  sexual  plant 
arises  as  a  lateral  branch.  The  Characeae  are  distinguished  by  the 
structure  of  their  vegetative  system  as  well  as  by  the  spirally- 
coiled  spermatozoids,  and  stand  as  an  isolated  group  among  the 
Thallophytes,  of  which,  however,  the  Siphoneae  appear  to  be  their 
nearest  relations.  They  were  formerly,  but  wrongly,  placed  near 
the  Mosses.  The  class  contains  only  one  order,  the  Characeae. 

Order  1.  Characeae.  Algae  with  a  peculiar  odour,  often 
incrusted  with  lime,  and  of  a  brittle  nature.  They  generally  grow 
gregariously  in  large  masses  at  the  bottom  of  fresh  and  brackish 
water,  and  are  from  a  few  inches  to  more  than  a  foot  in  height. 
The  stem  has  long  internodes  which  in  Nitella  are  formed  of  one 
cylindrical  cell;  in  Chara  of  a  similar  cell,  but  closely  surrounded 
by  a  cortical  layer  of  smaller  ones.  The  protoplasm  in  contact 
with  the  cell- wall  exhibits  in  a  well-marked  degree  the  movement 
of  rotation  (cyclosis),  carrying  the  chlorophyll  corpuscles  along 
with  it.  The  internodes  are  separated  from  each  other  by  a  layer 
of  small  cells  (nodal  cells)  from  which  the  leaves  are  produced. 
The  leaves  are  borne  in  whorls  of  from  5-12  which  regularly 
alternate  with  one  another  as  in  the  higher  verticillate  plants  ;  a 
branch  is  borne  in  the  axil  of  the  first  formed  leaf  of  each  whorl 
(Fig.  61  A,  n). 

The  leaves  are  constructed  in  the  same  manner  as  the  stem ; 
they  are  divided  into  a  series  of  joints,  but  have  only  a  limited 


CHARACE^l.  65 

power  of  growth  ;  their  terminal  cell,  too,  is  not  enclosed  by  a 
cortex.  Leaflets  are  borne  at  their  nodes.  The  growth  of  the 
stem  is  unlimited,  and  proceeds  by  means  of  an  apical  cell  (Fig. 
62  .s).  The  apical  cell  divides  into  a  segment-cell  and  a  new 
apical  cell.  The  segment-cell  then  divides  by  a  tivmsverse  wall 
into  two  cells,  one  lying  above  the  other  ;  the  lower  one,  without 


FIG.  61.— Cltarafrajilis.  A  Portion  of  a  plant,  natural  size.  B  Portion  of  a  leaf  b,  with 
'leaflets  j3'-|3" ;  a  antherldiiim ;  c  oogonium.  C  A  shield.— Nitella  flexilis.  D  Filament  from 
antheridium  with  sperm  itozoids.  E  Free  spermatozoid!". 

Any  further  division,  becomes  one  of  the  long,  cylindrical,  inter- 
nodal  cells  (Fig.  62  m),  and  the  upper  one  (Fig.  62  n)  divides 
by  vertical  walls  to  form  the  nodal  cells.  The  cortical  cells 
(Fig.  62  r)  which  surround  the  long  internodal  cells  of  Chara,  are 
derived  from  the  divisions  of  the  nodal  cells ;  the  cells  covering 
'the  upper  portion  of  an  internodal  cell  being  derived  from  the 


66 


CHARACE.E. 


node  immediately  above  it,  and  those  in  the  lower  part  of  the 
internode  from  the  node  below  it. 

The  organs  of  reproduction  are  very  conspicuous  by  their  colour 
and  form.  They  are  always  situated  on  the  leaves,  the  plants 
being  very  frequently  monoecious.  The  antheridia  (Fig.  61  B,  a) 
are  modified  leaflets  or  the  terminal  cell  of  a  leaf;  they  are  spheri- 
cal and  become  red  when  mature.  Their  wall  consists  of  £ 
"shields,"  i.e.  of  plate-like  cells,  4  of  which  cover  the  upper  half, 
and  are  triangular ;  the  4  round  the  lower  half,  to  which  the  stalk 
of  the  antheridia  is  attached,  being  quadrilateral,  with  sides  of 
unequal  length.  The  shields  (Fig.  61  C)  have  dentated  edges, 


FIG.  G2. — Chara  fragilis :  s  apical  cell; 
n,n  nodal  cells;  in  internodal  cells;  U, 
Itl  leaves ;  r,  r  the  cortical  cells. 


FIG.  03.  —  Oogoiiiutn  of  Chara  :  K 
"crown";  u  receptive  spot;  s  sperma- 
tozoids. 


with  the  teeth  fitting  into  one  another,  and  their  faces  ornamented 
with  ridges.  From  the  centre  of  the  internal  face  of  each  shield 
(C)  a  cylindrical  cell,  the  manubrium,  projects  nearly  as  far  as  the 
centre  of  the  antheridium  ;  at  the  inner  end  of  each  of  the  manu- 
bria  a  spherical  cell,  the  capitulum,  is  situated.  Each  capitulum 
bears  six  secondary  capitula,  from  each  of  which  four  long  coiled 
filaments  (0,  D)  project  into  the  cavity  of  the  antheridium. 
These  filaments  are  divided  by  transverse  walls  into  from  100-200 
discoid  cells,  in  each  of  which  a  biciliated,  coiled  spermatozoid  is- 
developed  (D,  E)  from  the  nucleus.  The  spermatozoid  s  escape 
from  their  mother-cell  and  are  set  free  by  the  shields  separating 
from  one  other. 


CHARACE^E. 


67 


The  female  organ  of  reproduction  (Fig.  61  B,  63)  is  a  small 
modified  shoot,  whose  apical  cell  functions  as  an  oogonium,  its 
protoplasm  forming  the  oosphere,  which  has  a  colourless  receptive- 
spot  at  the  summit  (Fig.  63  u~).  The  oogonium  is  situated  on  a 
nodal  cell,  from  which  5  cells  grow  out  in  a  circle  and  coil  round 
the  oogonium,  covering  it  with  a  close  investment.  These  cells 
divide  once  or  twice  at  the  top,  so  that  5  or  10  small  cells  are 
cut  off,  which  project  above  the  oogo- 
nium and  form  the  so-called  "crown" 
(Fig.  63  &).  The  crown  either  drops  off 
at  fertilisation,  or  its  cells  separate  to 
form  a  central  canal  for  the  passage  of 
the  spermatozoids.  The  wall  of  the 
oosphere l  above  the  receptive  spot  be- 
comes mucilaginous,  and  allows  the 
spermatozoid  to  fuse  with  the  oosphere. 
The  oospore,  on  germination  (Fig.  64  sp), 
becomes  a  small  filamentous  plant,  of 
limited  growth  (Fig.  64  i,  d,  q,  pi) — the 
proembryo — and  from  this,  as  a  lateral 
outgrowth,  the  sexual  generation  is  pro- 
duced. 

The  order  is  divided  into  two  sub- 
orders : — 

A.  NITELLE^}.     The  crown  consists  of 
1Q  cells;    cortex  absent:    Nitella,   Toly- 
pella. 

B.  CHAREJ;.     The  crown  consists  of  5 
•cells  ;  cortex  present  :  Tolypellopsis,  Lam- 
prothamnus,  Lychnothamnus,  Chara. 

Chara  crinita  is  parthenogenetic ;  in 
large  districts  of  Europe  only  female 
plants  are  found,  yet  oospheres  are 
formed  capable  of  germination. 

About  40  species  of  fossilized  Chara, 
determined  by  their  carpogonia,  are 
known  in  the  geological  formations  from 
the  Trias  up  to  the  present  day. 


FiG.6t.— Chara  fragilis.  Ger- 
minating oospore  (sp);  i,d,g,pl, 
form  together  the  proembryo  j 
rhizoids  (ic")  are  formed  at  d; 
w'  the  so-called  tap-root;  at  g 
are  the  first  leaves  of  the  sexual 
plant.which  appears  as  a  lateral 
bud. 


1  Before  fertilisation  the  oosphere  divides  and  cuts  off  at  the  base  one  or 
more  cells  (polar  bodies?),  termed  "  wendungszellen." 


68 

Class  8.   Phaeophyceae  (Olive-Brown  Seaweeds). 

The  Phaeophyceee  are  Alga?,  with  chromatophores  in  which  the 
chlorophyll  is  masked  by  a  brown  colour  (phycophasin).  The  pro- 
duct of  assimilation  is  a  carbohydrate  (fucosan),  never  true  starch. 
In  the  highest  forms  (Fucacece),  the  thallus  presents  differentiation 
into  stem,  leaf,  and  root-like  structures.  The  asexual  reproduction 
takes  place  by  means  of  zoospores.  The  sexual  reproduction  is 
effected  by  the  coalescence  of  motile  gametes,  or  by  oogamous 
fertilisation.  The  swarm-cells  are  monosymmetric,  each  moved 
by  two  cilia  which  are  true  protoplasmic  structures,  and  generally 
attached  laterally  (Fig.  65).  The  Phasophyceee  are  almost  entirely 
salt-water  forms;  a  few  species  of  Lithoderma  live  in  fresh  water. 

The  class  is  divided  into  two  families  : — 

1.  PMIOSPORE^;  :    1    Sub-Family,   Zoogonicaa;    2   Sub-Family, 
Acinetae. 

2.  CYCLOSPORE^;  :  Fucaceoe. 

Family  1.     Phaeosporeae. 

The  family  consists  of  multicellular  plants,  whose  cells  are 
firmly  united  together  to  form  a  thallus;  this,  in  the  simplest 
cases,  may  be  a  branched  filament  of  cells  (Ectocarpus),  or,  in  the 
highest,  may  resemble  a  stem  with  leaves  (Laminariacece} ,  while 
all  transitional  forms  may  be  found  between  these  two.  The 
thallus  grows  by  intercalary  divisions  (e.g.  Ectocarpus],  or  by  an 
apical  cell  (e.g.  Sphacelaria)  ;  pseudo-parenchymatous  tissue  may 
sometimes  be  formed  by  cells,  which  were  originally  distinct, 
becoming  united  together.  The  size  of  the  thallus  varies  ;  in 
some  species  it  is  quite  small — almost 
microscopical, — while  in  the  largest  it 
is  many  metres  in  length. 

The  vegetative  cells  in  the  lower 
forms  are  nearly  uniform,  but  in  those 
which  are  more  highly  developed 
(Laminariacece  and  Fucaceic\  they  are- 
FIG.  65.— Swarmspore  of  CMeria  sometimes  so  highly  differentiated  that 
mechanical,  assimilating,  storing  and 

conducting  systems  may  be  found  ;  the  last  named  systems  are 
formed  of  long  cells  with  perforated,  transverse  walls,  which  bear 
a  strong  resemblance  to  the  sieve-tubes  in  the  higher  plants. 

The  colouring  niatter  in  the  living  cells  ("  phoeophyl  ")  contains 


PH.EOSPORE.E.  69 

chlorophyll;  but  this  is  concealed  by  a  brown  ("  phycophsein  "), 
and  a  yellow  ("  phycoxanthin  ")  colouring  material,  and  hence  all 
these  Alga3  are  a  lighter  or  darker  yellow-brown.  Starch  is  not 
formed.  Asexual  reproduction  takes  place,  (1)  by  zoospores 
which  arise  in  unilocular  zoosporangia,  and  are  monosymmetric, 
with  two  cilia  attached  laterally  at  the  base  of  the  colourless 
anterior  end  (Fig.  65),  the  longer  one  being  directed  forwards 
and  the  shorter  backwards  ;  or  (2)  by  aplanospores  (?). 

Sexual  reproduction  has  only  been  discovered  in  a  few  cases,  and 
takes  place  by  means  of  gametes  (oogamous  fertilisation  perhaps 


Fi«i.  66.  —  Ectocarpus  kiliculosus.  I  a-/ 
A  f  dmale  gamete  in  the  various  stages 
of  coming  to  rest.  II  A  motionless 
female  gamete  surrounded  by  male 
gametes.  Ill  a-e  Stages  in  the  coal- 
escence of  male  and  female  gametes. 


FIG.  67.  —  Zanardim'a  coUaris.  A  Male 
gametangia  (the  smaller  celled)  and  female 
gametangia  (the  larger  celled).  C  Female 
gamete.  D  Male  gamete.  B,  E  Fertilisation. 
F  Zygote.  6  Germinating  zygote. 


occurs  in  the  Tilopteridae).  The  gametes  have  the  same  structure 
as  the  zoospores,  an<i  arise  in  multilocular  gametangia ;  these,  like 
the  zoosporangia,  are  outgrowths  from  the  external  surface,  or 
arise  as  modifications  from  it.  The  conjugating  gametes  may  be 
similar  (e.g.  Ectocarpus  pusillus),  or  there  may  be  a  more  or  less 
pronounced  difference  of  sex,  an  indication  of  which  is  found  in 
Ectocarpus  siliculosus  (Fig.  66).  When  the  gametes  in  this 
species  have  swarmed  for  a  time,  some,  which  are  generally  larger, 


70 

are  seen  to  attach  themselves  by  one  of  the  cilia,  which  by  degrees 
is  shortened  to  form  a  kind  of  stalk  (compare  the  upper  gamete  in 
Fig.  66  II)  ;  these  are  the  female  gametes,  which  now  become  sur- 
rounded by  a  number  of  males  endeavouring  to  conjugate  with 
them,  but  only  one  succeeds  in  effecting  fertilisation.  The 
protoplasm  of  the  two  gametes  coalesces  (Fig.  66  III),  and  a 
zygote  (e)  is  formed.  The  male  gametes  which  do  not  conjugate 
may  germinate,  but  the  plants  derived  from  them  are  much  weaker 
than  those  produced  by  the  zygotes.  Strongly  pronounced  sexual 
differences  are  found  in  the  Cutleriaceae,  in  which  order  the  male 
and  female  gametes  arise  in  separate  gametangia  (Fig.  67.4).  The 
male  gametes  (Fig.  67  D)  are  much  smaller  than  the  female 
gamete  (Fig.  67  0) ;  the  latter,  after  swarming  for  a  short  time, 
withdraws  the  cilia,  and  is  then  ready  to  become  fertilised  (Fig. 
67  J5,  E\  thus  we  have  here  a  distinct  transition  to  the  oogamous 
fertilisation  which  is  found  in  the  Fucaceae.  Alteruation  of  gene- 
rations is  rarely  found. 

1.     Sub-Family.     Zoogonicae. 
Reproduction  by  means  of  gametes  and  zoospores. 
Order  1.     Ectocarpaceae.     The  thallus    consists  of   single    or 
branched  filaments  with  intercalary  growth,  extending  vertically 
from  a  horizontal,  branched  filament  or  a  disc,  but  sometimes  it  is 
reduced  to  this  basal  portion  only.     Zoosporangia  and  gametangia 
(for  fertilisation  see   Fig.  66)  are  either  outgrowths  or  arise  by 
the  transformation  of  one  or  several  of  the  ordinary  cells.     The 
most  common  genera  are  :  Ectocarpus  and  Pylaiella. 

Order  2.  Choristocarpacese.  Choristocarpus,  Discosporangium. 
Order  3.  Sphacelariaceae.  The  thallus  consists  of  small, 
parenchymatous,  more  or  less  ramified 
shoots,  presenting  a  feather-like  appear- 
ance. In  the  shoots,  which  grow  by 
means  of  an  apical  cell  (Fig.  68  £),  a 
cortical  layer,  surrounding  a  row  of 
central  cells,  is  present.  Sporangia  and 
gametangia  are  outgrowths  from  the 
main  stem  or  its  branches.  Sphacelaria, 
Choetopteris  are  common  forms. 

Order  4.     Encoeliaceae.     Punctaria,  Aspero- 

coccus,  Phyllitis  fascia. 
FIG.  68.— Apex  of  the  thallus 

otCha-wterisplumoBo.  S  Api-  Order  5-  Stnanaceae.  ptriaria,  Phlao- 
<•«!  cell.  spora. 


PH-EOSPOEE.E. 


71 


Order  6.     Dictyosiphonaceae.     Dictyosiphon. 

Order  7.     Desmarestiaceae.     Desmarestia  aculeata  is  common. 

Order  8.     Myriotrichiaceae.     Myriotrichia. 

Order  9.  Elachistaceae.  Elachista  fucicola  is  &  common  epiphyte  on 
species  of  Fitcus. 

Order  10.  Chordariaceae.  The  shoot-systems  are  often  surrounded  by 
mucilage.  Chordaria  ;  Leathesia  difformis  occurs  as  rounded,  brown-green 
masses  of  the  size  of  a  nut,  generally  attached  to  other  Seaweeds. 

Order  11.     Stilophoracese.     Stilophora  rhizodes  is  common. 

Order  12.     Spermatochnaceae.     Spermatochnus  paradoxus  is  common. 

Order  13.     Sporochnaceae.     Sporochnus. 

Order  14.  Ralfsiaceae.  Ralfsia  verrucosa  is  common  as  a  red-brown  incrus- 
tation on  stones  and  rocks  at  the  water's  edge. 

Order  15.  Lithodermataceae.  Some  species  of  the  genus  Lithodenna  occur 
in  fresh  water. 

Order  16.  Laminariaceae.  The  thallus  is  more  or  less  leathery, 
and  has  generally  a  root-like  lower  part  (Fig.  69)  which  serves  to 
attach  it,  and  a  stalk  or  stem-like  part,  terminated  by  a  large  leaf- 
like  expansion.  Meristematic  cells 
are  situated  at  the  base  of  the  leaf, 
and  from  these  the  new  leaves  are 
derived.  The  older  leaf  hust 
pushed  away  by  the  intercalary 
formation  of  the  younger  ones, 
soon  withers  (Fig.  69).  Gametes 
are  wanting.  Zoosporangia  are 
developed  from  the  lower  part  of 
a  simple,  few-celled  sporangio- 
phore,  which  is  an  outgrowth  from 
a  surface-cell  and  has  a  large 
club-formed  apical  cell.  The  spo- 
rangia are  aggregated  into  closely 
packed  sori,  which  cover  the  lower 
part  of  the  terminal  leaf,  or  occur 
on  special,  smaller,  lateral,  fertile 
fronds  (Alaria).  Most  of  the 
species  belonging  to  this  order  live 
in  seas  of  moderate  or  cold  tem- 
perature and  OCCUr  in  the  most 
northern  regions  that  have  yet 

'  .          . 

been  explored,  forming  their  organs 

of  reproduction  during  the  cold  and  darkness  of  the  arctic  night. 

Laminaria  is  destitute  of  a  midrib  and  has  only  one  terminal  leaf. 


Jl 


Fm.  69._iflm»n0ria  digitata  (much  re- 

duced  in  8ize)-   B  Portion  of  a  section 

through  a  sorus  ;   sp  sporangia  ;  h  para- 


72  PHjEOSPORE^l. 

L.  digitata  has  a  broad  leaf,  which,  by  the  violence  of  the  waves,  is 
torn  into  a  number  of  palmate  strips  (Fig.  69).  L.  saccharina  has 
a  small,  undivided  leaf.  Alaria  has  a  midrib  and  special  fertile 
fronds.  A.  esculenta  occurs  plentifully  on  the  west  coast  of  Nor- 
way and  on  the  shores  of  Great  Britain.  Chorda  filum,  a  common 
seaweed,  is  thick,  unbranched,  and  attains  a  length  of  several 
metres,  without  any  strong  demarcation  between  stalk  and  leaf. 
Some  attain  quite  a  gigantic  size,  e.g.  Macrocystis  pyrifera,  whose 
thallus  is  said  sometimes  to  be  more  than  300  metres  in  length. 
The  Lessonia-species,  like  the  above,  form  submarine  forests  of 
seaweed  on  the  south  and  south-west  coasts  of  South  America, 
the  Cape,  and  other  localities  in  the  Southern  Hemisphere. 

USES.  The  large  Laminarias,  where  they  occur  in  great  numbers,  are,  like  the 
Fuel,  used  for  various  purposes,  for  example,  in  the  production  of  iodine  and 
soda,  and  as  an  article  of  food  (Laminaria  saccharina,  Alaria  esculenta,  etc). 
Laminaria  saccharina  contains  a  large  quantity  of  sugar  (mannit)  and  is  in 
some  districts  used  in  the  preparation  of  a  kind  of  syrup  ;  in  surgical  operations 
it  is  employed  for  the  distension  of  apertures  and  passages,  as  for  instance  the 
ear-passage.  It  is  by  reason  of  the  anatomical  peculiarities  and  structure  of  the 
cell-walls,  that  they  are  employed  for  this  purpose.  The  cell-walls  are  divided 
into  two  layers,  an  inner  one  which  has  very  little  power  of  swelling,  and  an 
outer  one,  well  developed  and  almost  gelatinous — the  so-called  "  intercellular 
substance  " — which  shrivels  up  when  dried,  but  can  absorb  water  and  swell  to 
about  five  times  its  size.  The  stalks  of  Laminaria  clustoni  are  officinal. 

Order  17.  Cutleriacese.  The  thallus  is  formed  by  the  union 
of  the  originally  free,  band-shaped  shoots.  The  growth  is  inter- 
calary. Sexual  reproduction  by  the  conjugation  of  male  and 
female  gametes.  An  asexual  generation  of  different  appearance, 
which  produces  zoospores,  arises  from  the  germination  of  the 
zygote.  Gutleria,  Zanardinia. 

Sub-Family  2.     Acinetae. 

Branched,  simple  cell-rows  with  intercalary  growth.  The 
organs  of  reproduction  are  partly  uni-  and  partly  multi-cellular  j 
in  the  unicellular  ones  a  cell  without  cilia  is  formed,  which  may  be 
destitute  of  a  cell- wall,  but  has  one  nucleus  (oosphere  ?),  or  which 
has  a  cell-wall  and  contains  several  (generally  four)  nuclei 
(aplanospores  ?);  in  the  multicellular,  monosymmetric  swarm-cells 
with  two  cilia  (spermatozoids  ?)  are  formed.  The  fertilisation  has 
not  been  observed. 

Order  1.     Tilopteridacese.     Haplospora,  Tilopteris. 


CTCLOSPORE2E.  73 

Family  2.     Cyclosporese. 

The  individuals  are  inulticellular,  with  growth  by  an  apical 
cell.  The  thallus — often  bilateral — is  differentiated  into  a  root- 
like  structure  (attachment-disc),  and  stem,  sometimes  also  into 
leaves  (Sargassum).  Sometimes  a  differentiation  occurs  into 
various  tissue-systems,  viz.  an  external  assimilating  tissue,  a  storing 
tissue,  a  mechanical  tissue  of  thickened,  longitudinal,  parenchyma- 
tous,  strengthening  cells,  and  a  conducting  tissue  of  sieve-cells,  or  of 
short  sieve-tubes  with  perforated  walls.  Colouring  material,  as 
in  Phseosporese.  Vegetative  reproduction  can  only  take  place  by 
means  of  detached  portions  of  the  thallus  (Sargassum},  which  are 
kept  floating  by  means  of  bladders  (Fig.  70  A,  a,  Fig.  72).  Zoo- 
spores  are  wanting. 

The  sexual  reproduction  takes  place  by  oogamous  fertilisation. 
The  oog'onia  and  antheridia  are  formed  inside  special  organs 
(conceptacles),  and  are  surrounded  by  paraphyses.  The  concep- 
tacles  (Fig.  70  B,  Fig.  71  b)  are  small,  pear-shaped  or  spherical 
depressions,  produced  by  a  special  ingrowth  of  the  surface  cells 
of  the  thallus,  and  their  mouths  (ostioles)  project  like  small  warts ; 
they  are  either  situated  near  the  end  of  the  ordinary  branches  of 
the  thallus  (Fucus  serratus,  Fig.  71  a)  which  may  be  swollen  on 
this  account  (Fucus  vesiculosus,  Fig.  70  A,  6),  or  on  special  short 
branches  (Ascophyllum,  Sargassum).  The  vertical  section  of  a 
conceptacle  is  seen  in  Fig.  70  B  (see  also  Fig.  71  6)  where,  in 
addition  to  the  paraphyses,  oogonia  only  are  seen  (F.  vesiculosus  is 
dioecious — male  plant,  yellow-brown;  female  plant,  olive-brown); 
but  in  some  species  antheridia,  together  with  oogonia,  are  pro- 
duced in  the  same  conceptacle.  The  oogonia  are  large,  almost 
spherical  cells,  situated  on  a  short  stalk,  in  each  of  which  are 
formed  from  1—8  (in  Fucus,  8 ;  in  Ascophijllum,  4  ;  in  Halidrys,  1 ; 
in  Pelvetia,  2)  rounded,  immotile  oospheres.  The  wall  of  the 
oogonium  raptures,  and  the  oospheres,  still  enclosed  in  the  inner 
membrane,  are  ejected  through  the  mouth  of  the  conceptacle,  and 
float  about  in  the  water,  being  finally  set  free  by  the  bursting  of  the 
inner  membrane.  The  antheridia  are  oblong  cells  (Fig.  70  C,  a), 
many  of  which  are  produced  on  the  same  branched  antheridio- 
phore  (Fig.  70  0) ;  the  numerous  spermatozoids  are  provided 
with  2  cilia  and  are  very  small  (Fig.  70  D,  two  antheridia  sur- 
rounded by  spermatozoids,  one  being  open).  The  spermatozoids, 
still  enclosed  by  the  inner  membrane  of  the  antheridium,  are 


74 


CYCLOSPOREJ;. 


similarly  set  free,  and  fertilisation  takes  place  in  the  water, 
numerous  spermatozoids  collecting  round  the  oosphere  (Fig.  70  E), 
which  is  many  times  larger,  and  by  their  own  motion  causing  it  to 
rotate.  After  fertilisation,  the  oospore  surrounds  itself  with  a 


D  E  F 

FIG.  70. — Fucus  vesiculosus.  A  Portion  of  thallus  with  swimming  bladders  (a)  and 
conceptacles  (b).  J5  Section  of  a  female  conceptacle  ;  h  the  mouth;  p  the  inner  cavity; 
s  oogonia.  C  Antheridiophore;  a  antheridium;  p  sterile  cells.  D  Antheridia  out  of  which 
the  spermatozoids  are  escaping.  E  Fertilisation.  F  Germinating  oospore. 

cell- wall  and  germinates  immediately,  attaching  itself  (Fig.  70  F) 
to  some  object,  and  by  cell-division  grows  into  a  new  plant. 


CYCLOSPOEEJ;. 


75 


Order  1.  Fucaceae.  The  following  species  are  common  on  our 
coasts  :  Fucus  vesictdosus  (Fig.  70)  has  a  thallus  with  an  entire 
margin,  and  with  bladders  arranged  in  pairs  ;  F.  serratus  (Fig.  71) 
without  bladders,  but  with  serrated  margin ;  Ascophyllum  nodosum, 
has  strap-like  shoots,  which  here  and  there  are  swollen  to  form 
bladders  ;  Halidrys  siliquosus  has  its  swimming  bladders  divided 
by  transverse  walls ;  Himanthalia  lorea,  which  is  found  on  the  west 
coast  of  Norway,  and  the  south  coast  of  England,  has  a  small 
perennial,  button-shaped  part,  from  the  centre  of  which  proceeds 
the  long  and  sparsely  branched,  strap-like,  annual  shoot,  which 


FIG.    72. — Sargafsum   bacciferwm. 
portion  of  the  thallus,  natural  size. 


FIG.  71. — Fucus  serratus.  a  Portion  of  a  male  plant  which  has  been  exposed  to  the 
action  of  the  open  air  for  some  time  ;  small  orange-yellow  masses,  formed  by  the  anther- 
idia.are  seen  outside  the  mouths  of  the  male  conceptacles(nat.size).  b  Cross  section  through 
the  end  of  a  branch  of  a  female  plant,  showing  the  female  conceptacles  (  x  4). 

bears  the  conceptacles.  The  Gulf-weed  (Sargassum  lacciferum, 
Fig.  72)  is  well  known  historically  from  the  voyage  of  Columbus  ; 
it  is  met  with  in  large,  floating,  detached  masses  in  all  oceans, 
and  is  found  most  abundantly  in  the  Atlantic,  off  the  Canary 
Islands  and  the  Azores,  and  towards  the  Bermudas.  The- 
stalked,  spherical  air-bladders  are  the  characteristic  feature  of 
this  genus.  The  thallus  is  more  highly  developed  than  in  Fucusr 
and  there  is  a  contrast  between  the  stem  and  leaf-like  parts.  The- 


76  DICTYOTALES. 

portions  which  are  found  floating  are  always  barren,  only  those 
attached  are  fertile. 

USES.  The  Fucaceae,  like  the  Laminariaceae,  are  used  as  manure  (the  best 
kinds  being  Fucus  vesiculosus  and  Ascophyllum  nodosum),  for  burning  to  pro- 
duce kelp,  and  as  food  for  domestic  animals  (Ascophyllum  nodosum  is  especially 
used  for  this  purpose). 

Class  9.    Dictyotales. 

The  plants  in  this  class  are  multicellular,  and  brown,  with 
apical  growth,  new  cells  being  derived  either  from  a  flat  apical 
cell,  or  from  a  border  of  apical  cells.  The  thallus  is  flat,  leaf-  or 
strap-shaped,  attached  by  haptera,  which  are  either  found  only 
at  the  base,  or  on  the  whole  of  the  lower  expansion  of  the  thallus. 
The  cells  are  differentiated  into  the  following  systems  of  tissues  : 
an  external,  small-celled  layer  of  assimilating  cells,  generally  one 
cell  in  thickness,  and  an  internal,  large-celled  layer  of  one  or  only 
a  few  cells  in  thickness,  forming  the  mechanical  and  conducting 
tissues.  All  the  reproductive  cells  are  motionless.  Asexual  re- 
production by  naked,  motionless  spores  (tetraspores)  which  are 
formed  1-4  in  each  tetrasporangium,  the  latter  being  outgrowths 
from  the  surface  cells  of  special,  sexless  individuals.  Zoospores 
are  wanting.  The  sexual  organs  are  of  two  kinds,  oogonia  and 
antheridia,  which  are  formed  from  the  surface  cells,  either  on  the 
same  or  different  individuals.  The  oogonia  are  spherical  or  oval, 
and  are  generally  placed  close  together;  each  contains  one 
oosphere,  which  on  maturity  is  ejected  into  the  surrounding 
water,  and  is  then  naked  and  motionless.  The  antheridia  are 
formed  of  longitudinal  cells,  united  in  groups,  whose  contents 
by  repeated  divisions— transverse  and  longitudinal — are  divided 
into  a  large  number  of  small,  colourless,  motionless  spermatia — 
round  or  elongated — which  are  set  free  by  the  dissolution  of  the 
wall  of  the  antheridium.  The  process  of  fertilisation  has  not 
yet  been  observed. 

The  Dictyotales,  in  having^  tetraspores  and  spermatia,  deviate 
-considerably  from  the  Phseophyceae,  but  may  be  classed  near  to 
the  Tilopteridge,  in  which  there  are  asexual  spores  with  4  cell- 
nuclei,  which  may  be  considered  as  an  indication  of  the  formation 
-of  tetraspores. 

Order  1.  Dictyotacese.  Dictyota  dichotoma  which  has  a  thin,  regularly 
.dichotomously  divided  thallus,  occurs  on  the  coasts  of  the  British  Isles 
Padina  is  found  on  the  south  coast. 


77 


Class  10.     Rhodophyceae    (Red    Seaweeds). 


The  plants  comprised  in  this  class  are  multicellular  ;  they  are 
simple  or  branched  filaments,  or  expansions  consisting  of  1  to 
several  layers  of  cells  ;  the  thallus  may  be  differentiated  (as  in 
many  Floridece),  to  resemble  stem,  root,  and  leaf.  The  cells  con- 
tain a  distinctly  differentiated  nucleus  (sometimes  several),  and 
distinct  chromatophores,  coloured  by  rhodopbyll.  The  chloro- 
phyll of  the  chromatophores  is  generally  masked  by  a  red  colour- 
ing matter  (phycoerythrin),  which  may  be  extracted  in  cold,  fresh 
water  ;  or  rarely  by  phycocyan.  Pyrenoids  occur  in  some.  Starch 
is  never  formed  in  the  chromatophores  themselves,  but  a  modifi- 
cation —  Floridea3  starch  —  may  be  found  in  the  colourless  proto- 
plasm. Asexual  reproduction  by  motile  or  motionless  spores 
(tetraspores)  which  are  devoid  of  cilia  and  of  cell-wall.  Swarm- 
spores  are  never  found. 

Sexual  reproduction  is  wanting,  or  takes  place  by  the  coales- 
cence of  a  spermatium  and  a  more  or  less  developed  female  cell. 
The  spermatia  are  naked  masses  of  protoplasm,  devoid  of  cilia  and 
chromatophores.  The  female  cell  (carpogonium)  is  enclosed  by  a 
cell-wall,  and  after  fertilisation  forms  a  number  of  spores,  either 
with  or  without  cell-  walls  (carpospores),  which  grow  into  new- 
individuals. 

The  RhodophyceaD  may  be  divided  into  two  families  : 

1.  BANGIOIDE^;. 

2.  FLORIDEJE. 

Family  1.     Bangioidese,. 

The  thallus  consists  of  a  branched  or  unbranched  cell-filament, 
formed  of  a  single  row  or  of  many  rows  of  cells,  or  of  an  expan- 
sion, one  or  two  layers  of  cells  in  thickness,  but  without  conspic- 
uous pores  for  the  intercommunication  of  the  cells.  The  growth  of 
the  thallus  is  chiefly  intercalary.  The  star-like  chromatophores 
contain  chlorophyll  and  are  coloured  blue-green  with  phycocyan, 
or  reddish  with  phycoerythrin  ;  all  these  colouring  matters  are 
occasionally  found  in  the  same  cell  (.Bawgrta-species).  Asexual 
reproduction  by  tetraspores,  without  cilia,  but  capable  of  amoeboid 
movements. 

Sexual  reproduction  is  wanting,  or  takes  place  by  the  coalescence 
of  a  spermatium  with  a  carpogonium,  which  is  only  slightly  differ- 
entiated from  the  vegetative  cells,  and  is  devoid  of  a  trichogyne. 


78  FLORIDE^E. 

The  carpospores  are  destitute  of  cell- wall  and  arise  directly  by 
the  division  of  the  fertilised  oosphere.  The  Bangioidege  occur 
chiefly  in  salt  water. 

Order  1.  Goniotrichaceae. — The  thallus  consists  of  a  branched  cell- 
filament  without  rhizoids.  Tetraspores  are  formed  directly  from  the  entire 
contents  of  the  mother-cell,  without  any  preceding  division.  Fertilisation 
unknown.  Asterocystis,  Goniotrichum. 

The  Goniotrichacece,  through  the  blue-green  Asterocystis,  are  allied  to  the 
Myxophyceffi,  and  through  Goniotrichum  to  the  Porphyracece. 

Order  2.  Porphyraceae. — The  thallus  is  formed  of  an  expansion  consisting 
of  a  layer  of  1-2  cells,  which,  at  the  base,  are  attached  to  the  substratum  by 
means  of  a  special  form  of  haptera  (Porphyra,  Diploderma)  ;  or  of  unbranched 
(very  rarely  slightly  branched)  filaments,  attached  at  the  base  by  haptera 
(Bangia) :  or  it  extends  from  a  prostrate  cell-disc  (various  species  of  Frytliro- 
trichia).  Tetraspores  are  formed  after  one  or  more  divisions  of  the  mother-cell, 
either  from  the  whole  or  only  a  part  of  its  contents ;  they  possess  amoeboid 
movements,  or  have  a  jerky,  sliding-forward  motion.  The  antheridia  have 
the  same  appearance  as  the  vegetative  cells,  but  divide  several  times,  and 
several  spermatia  are  formed,  either  simultaneously  from  the  whole  contents 
(Porphyra,  Bangia),  or  the  spermatia  are  successively  formed  from  a  part  of 
the  contents  of  the  antheridium  (Erythrotrichia).  The  carpogonium  is  with- 
out a  trichogyne,  but  the  oosphere  has  a  colourless  spot  which  may  some- 
times rise  a  little  above  the  surface  of  the  thallus,  and  may  be  considered 
as  an  early  stage  in  the  development  of  the  trichogyne.  The  spermatia  form 
a  canal  through  the  membrane  of  the  carpogonium,  and  their  contents  coal- 
esce with  the  oosphere  at  its  colourless  spot.  The  fertilised  oosphere  divides 
on  germination  into  a  number  of  carpospores,  which  are  set  free  as  naked, 
motionless  masses  of  protoplasm,  which  grow  and  give  rise  to  new  individuals 
(alternation  of  generations). 

Family  2.     Florideae. 

The  thallus  has  one  or  more  apical  cells,  grows  principally  by 
apical  growth,  and  may  be  differentiated  into  root,  stem,  and  leaf. 
The  chromatophores  vary  in  form,  but  have  a  red  or  brownish 
colour,  due  to  chlorophyll  and  phycoerythrin.  Asexual  repro- 
duction by  motionless  tetraspores,  which  generally  arise  by  the 
division  into  four  of  the  contents  of  the  tetrasporangium.  The 
carpogonium  has  a  trichogyne,  and  the  carpospores,  which  are 
formed  indirectly  from  the  fertilised  oosphere,  possess  a  cell- 
wall. 

The  thallus  may  assume  very  different  forms.  In  the  simplest 
species  it  is  filamentous  and  formed  of  single,  branched  rows  of 
cells  (Callithamnion,  etc.,  Fig.  73).  Ceramium  has  a  filamentous, 
thallus.  generally  dichotomously  forked  (Fig.  75),  or  sometimes 


FLORIDEJ:. 


79 


pinnately  branched,  which,  at  the  nodes,  or  throughout  its  entire 
length,  is  covered  by  a  layer  of  small  cortical  cells.  Polysiphonia 
(Fig.  74)  has  a  filamentous,  much  branched  thallus,  made  up  of 
a  central  cylindrical  cell,  surrounded  by  a  layer  of  other  cells, 
cortical  cells,  which  in  length  and  position  correspond  to  the 
central  ones.  In  many  of  the  Red  Algee  the  vegetative  organs 
are  differentiated  into  stems  and  leaves,  the  former  having,  as  in 
Chara,  unlimited  growth  in  length,  whilst  the  latter  soon  attain 
their  full  development.  Chondrus  has  a  fleshy,  gelatinous  thallus, 
without  nodes ;  it  is  repeatedly  forked  into  flat  branches  of  vary- 
ing thickness.  Furcellaria  has  a  forked  thallus  with  thick  branches. 


-  PIG.  73.  —  Callithamnion  elegans  :  a  a  plant  with 
tetraspores  ( x  20) ;  b  apex  of  a  branch  with 
tetraspores  ( x  250). 


FIG.  74. — PoZysi'pIioiua  variegata: 
a  a  portion  of  a  male  plant  with 
antberidia;  b  spermatia ;  c  trans- 
verse section  of  thallus. 


and  without  nodes.  The  thallus  of  Delesseria  (Fig.  76)  consists  of 
branches,  often  bearing  leaf-like  structures,  with  a  midrib  and 
lateral  ribs  springing  from  it.  These  ribs  persist  through  the 
winter,  and  at  the  commencement  of  the  succeeding  period  of 
vegetation  the  lateral  ribs  become  the  starting  points  for  new 
leaves.  In  Gorallina  the  thallus  is  pinnately  branched,  and 
divided  into  nodes  and  internodes.  The  name  has  been  given  to 
this  genus  from  the  fact  that  the  thallus  is  incrusted  with  car- 
bonate of  lime  to  such  a  degree  that  it  becomes  very  hard,  and  the 
w.  B.  G 


80 


FLOBIDEJE. 


whole  plant  adopts  a  coral-like  appearance.  Other  genera  which 
are  similarly  incrusted,  and  have  a  leaf-like  or  even  crustaceous 
thallus  (such  as  Melobesia,  Lithothamnioii),  are  included  in  this 
family. 

In  some  instances  the  cells  of  the  thallus  may  be  found  differ- 
entiated into  more  or  less  well  denned  tissues,  so  that  it  is  possible 
to  find  special  assimilating,  mechanical,  and  conducting  tissues, 
the  last  named  in  some  cases  having  the  double  function  of  con- 
ducting and  of  serving  as  a  reservoir  in  which  starch  is  found  as  a 
reserve  material.  The  cells  of  the  Florideae,  which  are  formed  by 
the  division  of  a  mother-cell  into  two  daughter-cells  of  unequal 


FIG.  75. — Ceramium  dinphanum  (nat.  size). 


FIG.  76  — Tlelesseria  sanguined  (about  j). 


size,  have  always  larger  or  smaller  pits  in  the  cell-walls,  and  the 
thin  cell- wall  separating  two  pits  from  each  other  is  perforated  by 
a  number  of  small  holes.  These  pits  are  particularly  developed  in 
the  conducting  tissues,  but  sieve-tubes  are  very  rarely  to  be  found. 

Tetraspores  may  be  wanting  (e.g.  Lemanea)  or  may  often  arise 
on  special,  non-sexual  individuals.  In  some  (e.g.  Batrachosper- 
mum)  only  one  tetraspore  is  formed  in  each  tetrasporangium,  but 
the  number  is  generally  four,  which  may  be  formed  tetrahedrally 
(Fig.  73)  or  by  divisional  walls  perpendicular  to  each  other,  or 
even  in  a  single  row.  The  tetrasporangia  in  some  species  are  free 
(Fig.  73),  but  in  the  majority  they  are  embedded  in  the  thallns. 

The   sexual   reproduction   (discovered  by   Thuret  and    Bornet, 


FLORIDE^. 


81 


1867)  differs  in  the  essential  points  from  that  of  all  other 
plants,  and  approaches  most  nearly  to  the  sexual  reproduction 
of  the  Bangioidece.  The  sexual  cells  are  developed  from  the 
terminal  cells  (never  nodal  cells)  of  the  branched  cell-filaments, 
which  constitute  the  thallus.  The  mother-cells  of  the  spermatia 
(xpermatangia)  are  generally  arranged  in  a  group,  in  the  so-called 
antheridia  (Figs.  74,  77  A,  a).  On  becoming  ripe  the  membrane 
of  the  spermatangium  ruptures  and  the  spermatia  emerge  as 
spherical  or  ovoid,  naked  (a  little  later  they  may  possess  a  cell- 
wall)  masses  of  protoplasm  which  are  not  endowed  with  the  power 
of  motion,  and  hence  are  carried  passively  by  the  current  of  the 


i 


FIG.  77. — A  Lejolisia  mediterranea :  r  haptera ;  s  longitudinal  section  t^rou'jrh  a  cystocnrp  ; 
*p  the  empty  space  left  by  the  liberated  spore  (0-  B-E  Nemalion  multifidum:  a  antheridia; 
b  procarpium  with  trichogyne,  to  which  two  spermsitia  are  adhering. 

water  in  which  they  may  happen  to  be,  to  the  female  cell.  This 
latter  is  analogous  with  the  oogonium  of  the  Green  Alga?.  The 
female  reproductive  organ  is  termed  the  procarpium,  and  consists 
of  two  parts,  a  lower  swollen  portion — the  carpogonium  (Fig.  77  6 
in  A  and  B) — which  contains  the  cell-nucleus,  and  an  upper 
filamentous  prolongation — the  trichoyyne  (Fig.  77  B) — which  is 
homologous  with  the  colourless  receptive  spot  of 'the  oosphere  of 
the  Green  Algae,  and  the  Porphyracecv.  In  the  sexual  reproduc- 
tion of  the  majority  of  the  Floridere,  a  very  important  part  is 
played  by  certain  special  cells,  rich  in  cell -contents — the  auxiliary 


82  FLORIDEJE. 

cells.  These  are  either  dispersed  in  the  interior  of  the  thallus, 
or  are  arranged  together  in  pairs  with  the  cell-filament  which 
bears  the  carpogonium,  and  are  generally  united  with  this  to 
form  an  independent  multicellular  procarpium.  The  spermatia 
attach  themselves  firmly  to  the  trichogyne  and  surround  them- 
selves with  a  cell- wall.  The  dividing  wall  at  the  point  of  contact 
is  perforated,  and  the  nucleus  of  the  spermatium  probably  travels 
through  the  trichogyne  to  the  swollen  part  of  the  procarpium 
— the  carpogonium — and  fuses  with  its  nucleus.  After  fertilisation 
the  trichogyne  withers  (Fig.  77  0),  bat  the  lower  portion  of  the 
procarpium,  constituting  the  fertilised  oosphere,  grows  out  and 
forms  in  various  ways,  first  a  tuft  of  spore-forming  filaments 
known  as  gonimoUasts,  and  finally  the  carpospores.  These  latter 
form  a  new  asexual  generation  (compare  the  germination  of  th& 
oospore  of  CEdogonium  and  Coleochcete}. 

The  gonimoblasts  may  arise  in  three  ways  :  — 

1.  In  the  Nemalionales,  branched  filaments  grow  out  from  the  oosphere  and 
form  an  upright,  compressed  or  expanded  tuft  of  spore-forming  filaments. 

2.  In  the  Cryptonemiales,  several  branched  or  unbranched  filaments  (ooblas- 
tema-filaments)  grow  out  from  the  oosphere,  and  conjugate  in  various  ways  with 
the  auxiliary  cells.     The  gonimoblasts  are  then  formed  from  the  single  cells 
produced  by  the  conjugation. 

3.  In  the  Gigartinaies  and  Rhodymeniales  the  oosphere  conjugates  with  an 
auxiliary  cell  by  means  of  a  short  ooblastem a- filament,  and  from  this  auxiliary 
cell  a  gonimoblast  is  produced. 

The  motionless  carpospores,  which  sometimes  in  the  early  stages  are  naked, 
and  afterwards  invested  with  a  cell-wall,  are  developed  from  the  terminal  cells 
(and  perhaps  also  from  some  of  the  other  cells)  of  the  branches  of  the  gonimo- 
blast. The  gonimoblasts  constitute  sharply  defined  parts  of  the  plant  in  which 
the  carpospores  arise.  These  parts  are  called  cystocarps  and  are  either  naked 
(Fig.  77 -E),  or  surrounded  by  a  covering  (pericarp  or  involucre,  Fig.  77  A^ 
formed  in  different  ways.  On  this  account  the  Florideae  were  formerly  divided 
into  GTYMNOSPORE^E  (Batrachospcrmum,  Nemalion,  Ceramium,  etc.)  and  ANGIO- 
SPORE.E  (Furcellaria,  Lejolisia,  Delesseria,  Melobesia,  etc.). 

The  Floridea3  are  divided  into  four  sub-families  : — 

Sub-Family  I.  Nemalionales.  The  fertilised  oosphere  produces  directly 
the  gonimoblast. 

Order  1.  Lemaneaceae.  Algse  of  brownish  colour  and  living  in  fresh 
water.  They  lack  tetraspores,  and  the  very  sparingly  branched  fertile  filaments, 
composed  of  many  rows  of  cells,  grow  out  from  a  pro-embryo,  which  consists  of 
a  single  row  of  cells  bearing  branches.  Lemanea  fluviatilis,  often  found  OB 
rocks  and  stones  in  quickly  flowing  streams. 


FLOR1DEJ].  »  83 

Order  2.  Helminthocladiaceae.  Tetraspores  are  generally  wanting  (e.g.  in 
Nemalion)  or  arise  one  in  each  tetrasporangium  (e.g.  Batrachospermum)  and  it 
is  only  in  Liagora  that  four  cruciate  tetraspores  are  formed.  Chantransia 
•corymUfera  consists  of  simple,  branched  cell-rows,  and  is  an  independent  species. 
Several  other  Chantransia- forms,  living  in  fresh  water,  are  "proembryos"  of 
species  of  the  genus  Batrachospermum.  The  germinating  carpospore  grows  out 
into  filaments  and  forms  a  so-called  proembryo  which,  if  not  shaded,  attains 
only  a  small  size,  but  when  growing  in  shady  situations  presents  a  much  greater 
development.  These  highly  developed  proembryos  have  been  described  as 
species  of  Chantransia.  The  proembryo  can  reproduce  by  division,  or  by  tetra- 
spores which  are  developed  singly  in  the  sporangia;  in  B.  vagum  and  B.  sporu- 
lans  which  do  not  possess  fully  developed  female  reproductive  organs,  the  pro- 
embryos  serve  almost  entirely  to  reproduce  the  species.  The  young  Batracho- 
spermum-plaut  arises  from  the  end  of  an  upright  filament  of  the  proembryo. 
The  proembryo  is  generally  persistent,  and  continually  produces  new  Batracho- 
spermums.  These  latter  bear  the  sexual  reproductive  organs  and  also  whorls  of 
branches  :  the  central  row  of  cells  is  enclosed  by  cells  growing  from  the  base  of 
the  whorls  of  branches,  and  from  these  cortical  cells  secondary  proembryos  are 
developed.  In  this  alternation  of  shoots  there  is  really  no  alternation  of 
generations,  since  the  proembryo  and  the  shoots  with  the  sexual  reproductive 
organs  are  parts  of  the  same  thallus. 

Several  species  of  Batrachospermum  have  a  bluish-green  or  verdigris  colour. 
Nemalion  multifidum  has  a  brown-red  thallus,  slightly  branched,  which  is 
attached  to  rocks  near  the  water's  edge. 

Order  3.  Chaetangiaceae.  Galaxaura  has  a  thallus  thickly  incrusted  with 
lime. 

Order  4.     Gelidiaceae.     Naccaria,  Gelidium. 

Sub-Family  2.  Gigartinales.  The  fertilised  auxiliary  cell  grows  towards 
the  thallus,  to  produce  the  gonimoblasts.  Procarpia  generally  present. 

Order  5.     Acrotylaceae.     Acrotylus. 

Order  6.  Gigartinaceae.  Gigartina,  Phyllophora,  Ahnfeltia;  Chondrus 
crispus,  with  dark  red,  dichotomously  branched  thallus,  is  common  on  the 
coasts  of  Scandinavia  and  Great  Britain. 

Order  7.  Rhodophyllidaceae.  Rhodophyllis,  Euthora;  Cystoclonium  pur- 
purascens  is  common,  and  sometimes  the  ends  of  its  branches  may  be  modified 
into  tendril-like  haptera. 

Sub-Family  3.  Rhodymeniales.  The  fertilised  auxiliary  cell  forms  the 
gonimoblast  on  the  side  away  from  the  thallus.  Procarpia  are  abundantly 
produced. 

Order  8.     Sphaerococcaceae.     Gracilaria. 

Order  9.  Rhodymeniacese.  Rhodymenia  palmata  is  a  common  species. 
Lomentaria,  Chylocladia,  Plocamium. 

Order  10.  Delesseriaceae.  Delesseria  sanguinca ;  D.  alata  and  D.  sinuosa 
are  handsome  forms  which  are  not  uncommon. 

Order  11.     Bonnemaisoniaceae.     Bonnemaisonia. 

Order  12.  Rhodomelaceae.  Rhodomela,  Odonthalia ;  Polysiphonia,  of 
which  many  species  are  to  be  found  on  the  coasts  of  Great  Britain,  has  a 
filamentous,  richly  branched  thallus  consisting  of  a  central  row  of  cells  sur- 


84  FUNGI. 

rounded  by  a  varying  number  of  cortical  cells  of  similar  size — the  so-called 
"  siphons." 

Order  13.  Ceramiaceae.  Pretty  Algas,  often  branched  dichotomously,  or 
unilaterally  pinnate.  Spcrmothamnion,  Oriffittisia,  Callithamnion,  Ceramiuni, 
Ptilota. 

Sub-Family  4.  Cryptonemiales.  The  cells  formed  by  the  coalescence 
of  the  auxiliary  cells  and  the  ooblastema-filaments,  produce  the  gouimoblasts. 
The  carpogoniiim-jilaments  and  the  auxiliary  cells  are  scattered  singly  in  the- 
thallus. 

Order  14.     Gloiosiphoniaceae.     Gloiopeltis. 

Order  15.     Grateloupiaceae.     Halymenia,  Cryptonemia. 

Order  16.     Dumontiacese.     Dumontia,  Dudresnaya. 

Order  17.  Nemastomaceae.  Furcellaria,  which  has  dichotomously 
branched,  round  shoots,  is  common  on  the  coasts  of  Great  Britain. 

Order  18.     Rhizophyllidaceae.     Polyides,  Rhizophyllis. 

.Order  19.  Squamariaceae.  The  Algas  belonging  to  this  order  form  crust- 
like  coverings  on  stones,  mussel-shells,  and  on  other  Alga?,  but  are  not  them- 
selves incrustated  :  Petrocelis,  Cruoria,  Peyssonellia. 

Order  20.  Corallinaceae.  Partly  crustaceous,  partly  erect,  branched  Algae, 
thickly  iucrusted  with  lime,  so  that  a  few  species  (Lithothamnia,  also  called 
Nullipora)  occur  in  fossilized  condition  from  Jurassic  to  Tertiary  periods. 
Melobesia,  Lithophyllum,  Lithothamnion,  Corallina. 

USES.  "  Carragen  "  is  the  thallus  of  Ghondrus  cricpus  (Irish 
Moss)  and  Gigartina  mamillosa.  It  is  a  common  article  of  food  on 
the  coasts  of  Ireland,  and  swells  to  a  jelly  when  cooked.  It  is 
officinal.  Rhodymenia  palmata  is  generally  eaten  as  food  in 
Ireland  and  in  some  places  on  the  west  coast  of  Norway ;  it  is 
also  used  as  food  for  sheep  and  hence  is  termed  "  Sheep-seaweed." 
Agar-Agar  is  the  jelly  obtained  from  species  of  Gelidium  and 
Gigartina  growing  in  China  and  Japan. 


Sub-Division  III.     FUNGI. 

Mode  of  Life.  The  Fungi  have  no  chlorophyll,  and  are  thus 
unable  in  any  stage  of  their  existence  to  assimilate  carbon ;  they 
must  therefore  live  as  saprophytes  or  parasites.  There  is,  however,, 
no  strong  line  of  demarcation  between  these ;  many  Fungi  com- 
mence as  true  parasites,  but  only  attain  their  full  develop- 
ment upon  or  in  dead  plants  or  animals  (Rhytisma,  Empusa). 
Many  saprophytes  may  occasionally  appear  as  parasites,  and  are 
then  designated  "facultative  parasites"  (Nectria'  cinnabarina* 
Lophodermium  pinastri),  in  contradistinction  to  those  which  only 


FUNGI.  85 

appear  as  parasites,  "  obligate  parasites  "  (Mildew,  Brand-  and  Rust- 
Fungi,  Cordyceps). 

The  parasites  which  live  on  the  surface  of  the  host-plant  are 
termed  epiphytic  (Mildew,  Fusicladium) ;  and  those  living  in  its 
tissues  are  termed  endophytic  (Ustilago,  Peronospora}.  Epizoic 
(Oidium  tonsurans,  Laboulbenia)  and  endozoic  Fungi  (Cordyceps, 
Entomophthora},  are  distinguished,  in  the  same  manner,  as  those 
which  live  on.  the  surface  or  in  the  interior  of  animals.  The  Fungi 
designated  pathogenic  are  especially  those  which  produce  disease 
in  human  beings  and  in  animals. 

Most  of  the  diseases  of  plants  are  attributed  to  the  parasitic 
Fungi.  These  force  their  way  into  the  host-plant  by  piercing  the 
outer  wall  of  the  epidermis,  as  in  the  Potato-disease ;  or  by  grow- 
ing in  through  the  stomata,  e.g.  the  summer  generations  of  the 
Bust  of  Wheat ;  or  they  can  only  penetrate  through  a  wound, 
e.g.  Nectria.  Some  effect  an  entrance  into  the  host-plant  by  the 
secretion  of  a  poison  (toxin)  and  a  ferment,  which  softens  and 
destroys  the  cell-walls  (Sclerotinia) .  Some  Yeast  and  Mould 
Fungi  secrete  ferments  (enzymes),  which,  for  example,  convert 
cane-sugar  into  a  sugar  capable  of  fermentation. 

The  relation  of  the  parasitic  Fungus  to  the  host-plant  is  mainly 
of  two  kinds.  In  the  one  case,  the  cell-contents  are  destroyed, 
the  protoplasm  is  killed,  and  the  cellular  tissue  becomes  discoloured 
and  dies  (Peronospora,  Armillaria  mellea,  Polyporus)  ;  in  the 
other  case,  the  parasite  has  an  irritating  effect  on  the  cellular 
tissue,  whereby  the  affected  organ  grows  more  rapidly  and  be- 
comes larger  than  normal,  producing  hypertrophy.  Such  malfor- 
mations are  termed  Fungi-galls  (Mycocecidia)  ;  in  this  manner 
"  witches'  brooms  "  are  produced  by  ^Eridiuitlj  "  pocket-plum  "  by 
Taphrina,  and  other  deformities  by  Exobasidium  and  Cystopus 
candidus.  This  hypertrophy  may  either  be  produced  by  a  vigorous 
cell-multiplication,  which  is  most  frequently  the  case,  or  by  the 
enlargement  of  the  individual  cells  (Synchytriwm,  Calyptospora). 
The  relation  between  host  and  Fungus  among  the  Lichens  is  of 
a  very  peculiar  nature,  termed  "  symbiosis." 

Vegetative  Organs.  The  vegetative  parts  of  a  Fungus  are 
termed  its  mycelium.1  This  is  formed  of  a  mass  of  long,  cylindrical, 
branched  cells  resembling  threads  (and  hence  termed  hyphai), 
which  have  a  continued  apical  growth.  The  mycelium,  in  its 
early  development,  shows  a  well-marked  difference  between  the 
1  From  the  Greek  ^175  =  Fungus,  hence  "  mycology." 


86  FUNGI. 

two  main  groups  of  true  Fungi :  in  the  Phycomycetes,  or  Algal 
Fungi,  the  mycelium  has  no  transverse  walls,  and  is  therefore 
unicellular,  while  in  the  Mesomycetes  and  Mycomycetes  it  is  pro- 
vided with  dividing  walls,  which  gradually  arise  during  growth,  in 
the  youngest  hyphse  ;  intercalary  transverse  walls  may  also  be 
formed  at  a  later  period.  In  the  hyphaa  of  some  of  the  Higher 
Fungi  (Hymenomycetes),  connections  may  be  formed  between  two 
contiguous  cells  of  the  same  hypha,  by  a  protuberance  growing  out 
from  an  upper  cell  just  above  the  transverse  wall,  and  forming  a 
junction  with  the  cell  below.  These  are  known  as  clamp-connec- 
tions- they  appear  to  be  of  use  in  affording  communication  be- 
tween the  two  cells. 

The  hyphte  of  Fungi,  where  they  come  in  contact  with  one 
another,  often  grow  together,  so  that  H-formed  combinations  (fu- 
sions) are  produced,  which  give  rise  to  very  compact  felted  tissue. 
When  the  hyphae  are  not  only  closely  interwoven,  but  also  united 
and  provided  with  many  transverse  walls,  the  mycelium  assumes 
the  appearance  of  a  tissue  with  isodiametric  cells,  and  is  then 
termed  pseudo-parenchyma.  The  hyphae-walls  are  sometimes  very 
much  thickened,  and  composed  of  several  layers,  and  the  external 
layers,  by  the  absorption  of  water,  may  often  swell  very  much 
and  become  mucilaginous.  In  some  ipstances  the  walls  are  colour- 
less, in  others  coloured,  the  most  .frequent  colour  being  brown. 
The  cell-contents  may  also  be  coloured,  and  in  that  case  are  gene- 
rally yellow;  this  colour  is  chiefly  connected  with  the  fat  (oil) 
which  may  be  found  in  abundance  in  the  Fungi,  whilst  starch  is 
invariably  absent  in  all  the  true  Fungi. 

The  mycelium  assumes  many  different  forms ;  sometimes  it 
appears  as  a  thread-like,  cobwebby,  loose  tissue,  less  frequently  as 
firm  strands,  thin  or  thick  membranes,  horn-like  plates  or  tuber- 
like  bodies.  The  thread-like  mycelium  may,  in  the  parasitic  Fungi, 
be  intercellular  or  intracellular,  according  as  it  only  extends  into 
the  interstices  between  the  cells  or  enters  into  the  cells  proper. 
In  the  first  case  there  are  generally  found  haustoria,  or  organs  of 
suction  (e.g.  among  the  Peronosporacece  ;  Taphrina,  on  the  contrary, 
has  no  haustoria) ;  but  haustoria  are  also  found  among  the 
epiphytic  Fungi  (e.g.  Erysiphacese).  Intracellular  mycelia  are 
found  in  the  Rust-Fungi,  in  Claviccps  purpurea,  Entomophthora,  etc. 
In  spite  of  its  delicate  structure,  this  mycelium  may  live  a  long 
time,  owing  to  the  circumstance  that  it  continues  to  grow  peri- 
pherally, while  the  older  parts  gradually  die  off  ("fairy  rings"). 


FUNGI.  87 

String-like  mycelia  may  be  found,  for  example,  in  Phallus, 
Coprinus,  and  are  formed  of  hyp Jh.se,  which  run  more  or  less 
parallel  to  each  other.  Membrane-like  mycelia  are  chiefly  to  be 
found  in  Fungi  growing  on  tree-stems  (Polyporacese  and  Agari- 
caceaa) ;  they  may  have  a  thickness  varying  from  that  of  the 
finest  tissue-paper  to  that  of  thick  leather,  and  may  extend  for 
several  feet.  The  peculiar  horny  or  leather-like  strands  and  plates 
which,  for  instance,  appear  in  Armillaria  mellea,  are  known  as 
Rhizomorpha  ;  they  may  attain  a  length  of  more  than  fifty  feet. 
The  tuber-like  mycelia  or  sclerotia  play  the  part  of  resting  mycelia, 
since  a  store  of  nourishment  is  accumulated  in  them,  and  after  a 
period  of  rest  they  develope  organs  of  reproduction.  The  sclerotia 
are  hard,  spherical,  or  irregular  bodies,  from  the  size  of  a  cabbage 
seed  to  that  of  a  hand,  internally  white  or  greyish,  with  a  brown 
or  black,  pseudo-parenchymatous,  external  layer.  Sclerotia  only 
occur  in  the  higher  Fungi,  and  are  found  both  in  saprophytes,  e.g. 
Coprinus,  and  in  parasites,  e.g.  Claviceps  (Ergot),  Sclerotinia. 

Reproduction.  SEXUAL  REPRODUCTION  is  found  chiefly  among 
the  lower  Fungi  wrhich  stand  near  to  the  Algse,  the  Algal-Fungi, 
and  takes  place  by  the  same  two  methods  as  in  the  Algae,  namely 
by  conjugation  and  by  the  fertilisation  of  the  egg-cell  in  the 
oogonium.  It  is  also  known  in  some  of  the  Higher  Fungi. 

The  majority  of  Fungi  have  only  ASEXUAL  reproduction.  The 
most  important  methods  of  this  kind  of  reproduction  are  the 
sporangio-fructification  and  the  conidio-fructification. 

In  the  SPORANGIO-FRUCTIFICATION  the  spores  (endospores)  arise 
inside  a  mother-cell,  the  sporangium  (Fig.  80).  Spores  without  a 
cell-wall,  which  move  in  water  by  means  of  cilia  and  hence  are 
known  as  swarmspores  or  zoospores,  are  found  among  the  Oomycetes, 
the  sporangia  in  which  these  are  produced  being*  called  swarm- 
sporangia  or  zoosporangia  (Figs.  86,  87,  91,  94). 

In  the  CONIDIO-FRUCTIFICATION  the  conidia  (exospores)  arise  on 
special  hyphse  (conidiophores),  or  directly  from  the  mycelium. 
When  conidiophores  are  present,  the  conidia  are  developed  upon 
them  terminally  or  laterally,  either  in  a  basipetal  succession 
(in  many  Fungi,  for  example  in  Penicillium,  Fig.  Ill,  Erysiphe, 
Cystopus),  or  acropetally  (in  which  method  the  chains  of  conidia 
are  often  branched ;  examples,  Pleospora  vulgaris,  Hormodendron 
dadosporioides).  All  conidia  are  at  first  unicellular,  sometimes  at 
a  later  stage  they  become  two-celled  or  multicellular  through  the 
formation  of  partition-walls  (Piptocephalis) .  The  conidia  with 


88  FUNGI. 

thick,  brown  cell-walls,  and  contents  rich  in  fats  (resting  conidia}, 
can  withstand  unfavourable  external  conditions  for  a  much  longer 
period  than  conidia  with  thin  walls  and  poor  in  contents. 

The  SPORANGIA  arise  either  from  the  ordinary  cells  of  the 
mycelium  (Protomyces),  or  are  borne  on  special  hyphee.  They  are 
generally  spherical  (Mucor,  Fig.  80 ;  Saprolegniaceee),  egg-,  pear-, 
or  club-shaped  (Ascomycetes),  more  rarely  they  are  cylindrical  or 
spindle-shaped.  While  among  the  Phycornycetes  the  size,  form, 
and  number  of  spores  are  indefinite  in  each  species,  in  the  Ascomy- 
cetes the  sporangia  (asci)  have  a  definite  size,  form,  and  number 
of  spores.  The  spores  of  the  Ascomycetes  are  known  as  asco- 
s  pores. 

The  sporangio-fructification  is  found  under  three  main  forms. 

1.  FREE  SPORANGIOPHORES  which  are  either  single  (Mucor,  Fig. 
78),  or  branched  (Thamnidium) . 

2.  SPORANGIAL- LAYERS.      These    are   produced   by  a   number   of 
sessile  or  shortly-stalked  sporangia,  being  formed  close  together 
like  a  palisade  (Taphrina,  Fig.  105). 

3.  SPORANGIOCARPS.     These  consist  usually  of   many  sporangia 
enclosed  in  a  covering,  they  are  found  only  in  the  Carpoasci,  and 
are   also   known  as  ascocarps.     The  parts  of    an  ascocarp  are  the 
covering  (peridium),  and  the  hymenium,  which  is  in  contact  with 
the  inner  wall  of  the  peridium,  and  is  generally  made  up  of  asci, 
and  sterile,  slender  hyphae.     The  latter  either  penetrate  between 
the  asci  and  are    branched   and  multicellular  (paraphyses,   Figs. 
103  d,  123,  125,  129),  or  clothe  those  parts  of  the  inner  wall  which 
bear  no  asci    (periphyses  ;  among  many  peronocarpic  Ascomycetes, 
e.g.  Chcefomium,  Sordaria,  Stictosphcera-  hoffmanni) .     The  ascocarps 
are  produced  directly  from  the  mycelium,  or  from  a  stroma,  that 
is  a   vegetative   body  of    various   forms,  in  which  they  may  be 
embedded  (Figs.  116  £,  (7). 

Among  the  conidio-fructifications  there  are,  in  the  same  way, 
three  divisions. 

1.  FREE  CONIDIOPHORES  (Fig.  109).   The  form  of  the  conidiophores, 
the  shape,   and  number  of  its  spores   are  various.     In  the  most 
highly  developed  Fungi,  the   Basidiomycetes,  there  are,  however, 
special  more  highly  developed  conidiophores,  the  basidia,  which 
have  a  definite  form  and  spores  of  a  definite  shape  and  number. 
The  conidia  borne  on  basidia  are  called  basidiospores . 

2.  CONIDIAL-LAYERS.       (a)   The    SIMPLEST   case   of   this    is    found 
when  the  conidiophores  arise  directly  from  the  mycelium,  parallel 


FUNGL  89 

to  one  another,  and  form  a  flat  body  (e.g.  Exobasidium  vaccinii, 
Hypochnus  ;  among  the  Phycomycetes,  Empusa  muscce  and  Cystopus). 
(6)  In  a  HIGHER  form  the  conidial-layers  are  thick,  felted  threads 
(stroma)  inserted  between  the  mycelium  and  the  hymenium  (i.e. 
the  region  of  the  conidiophores).  Examples  are  found  in  a  section 
of  the  Pyrenomycetes  (Fig.  122).  (c)  The  HIGHEST  form  has  the 
has  id  tal -layer,  that  is  a  conidial-layer  with  more  highly  developed 
conidiophores  (basidia).  The  basidial-layer,  with  stroma,  and 
the  hymenium  (region  of  the  basidia),  forms  the  basidio-fructifica- 
tion,  which  is  branched  in  the  Clavariaceaa,  and  hat-shaped  in 
other  Hymenomycetes  (in  these  groups  the  hymenium  is  confined 
to  the  lower  side  of  the  pileus). 

The  hymenium  of  the  conidial-layer  and  basidial-layer  is  com- 
posed entirely  of  conidiophores,  or  of  conidiophores  and  sterile 
hypha3  (paraphyses)  which  are  probably  always  unicellular. 
Paraphyses  are  found  in  Entomophthora  radicans,  and  in  certain 
Basidiomycetes  (e.g.  Corticium). 

3.  CONTDIOCARPS  (pycnidio).  A  special  covering  surrounds  the 
conidia-forming  elements.  The  inner  side  of  this  covering 
(peridium)  bears  the  hymenium,  i.e.  those  elements  from  which 
the  conidia  are  abstricted.  The  conidiocarps  arise  either  imme- 
diately from  the  hypha3  or  from  a  stroma  in  which  they  are 
generally  embedded.  Conidiocarps  are  entirely  wanting  in  the 
Phycomycetes.  On  the  other  hand  they  are  found  among  the 
Ascomycetes  and'  Basidiomycetes,  and  in  the  latter  group  the 
conidiocarps  contain  more  highly  differentiated  conidiophores 
(basidia)  and  are  known  as  basidiocarps.  Conidiocarps  with  simple 
conidiophores,  are  found  only  among  the  Basidiomycetes,  in  the 
Uredinaceee,  and  in  Craterocolla  cerasi.  In  the  Ascomycetes  (Figs. 
120  d,  e  ;  117  a,  b  ;  123  a  ;  124  b)  the  conidiocarps  are  visible,  as 
points,  to  the  naked  eye,  while  the  basidiocarps  of  the  Basidiomy- 
cetes (Figs.  170,  171,  173-176,  178-180)  vary  from  the  size  of  a 
pea  to  that  of  a  child's  head.  The  "  spermogonia "  of  the 
Ascomycetes  and  Lichenes,  are  conidiocarps  with  small  conidia 
(microconidia)  which  germinate  sometimes  more  slowly  than  other 
conidia.  There  is  good  evidence  for  considering  these  as  male 
reproductive  cells,  called  sperruatia. 

The  conidia  of  the  Fungi  are  not  primitive  structures.  The 
comparison  of  the  sporangia  and  conidia  among  the  Zygomycetes, 
and  among  the  species  of  the  genus  Peronospora  shows,  that  the 
conidia  are  aberrant  formations,  and  that  they  have  arisen  through 


90  FUNGI. 

the  degeneration  of  the  sporangium,  which,  by  the  reduction  of  its 
spores  to  one,  has  itself  become  a  spore. 

In  the  genera  Thamnidium  and  Ch&tocladium  the  gradual  diminution  of  the 
sporangia,  and  the  reduction  of  the  number  of  spores  can  be  distinctly  followed. 
In  Thamnidium  the  number  of  spores  is  often  reduced  to  one,  which  is  free  in 
the  sporangium.  In  Chatocladium  however  the  sporangia  are  typically  one-spored, 
the  spore  is  always  united  with  the  sporangium,  and  the  two  become  a  single 
body,  the  so-called  conidium,  which  is  in  reality  a  closed  sporangium.  How 
close  is  the  connection  between  the  sporangia  and  conidia  of  Thamnidium 
and  Chtftocladium,  is  seen  from  the  fact  that,  in  the  conidial  stage  of 
Chatocladium  the  same  whorl-form  of  branching  appears  as  in  the  sporangial 
stages  of  Thamnidium  chtftocladioides,  and  also,  that  the  conidia  of  Ch.  fresen- 
ianum  throw  off  the  former  sporangium-wall  (exosporium),  while  Ch.  jonesii 
germinates  without  shedding  its  exosporium.  The  Phycomycetes  have  doubt- 
less sprung  from  Water-Alges  and  inherit  the  sporangia  from  them.  On  this 
supposition,  as  the  Phycomycetes  assumed  a  terrestrial  mode  of  life,  the  sporangia 
would  become  adapted  to  the  distribution  of  the  spores  by  means  of  the  air, 
the  sporangia  would  become  small,  contain  dust-like  spores,  and  would 
eventually  become  closed-sporangia,  i.e.  conidia.  The  conidia  are  a  terrestrial 
method  for  the  multiplication  of  Fungi.  In  the  Hemiasci  and  the  Ascomycetes 
the  sporangia  are  still  preserved,  but  in  every  instance  they  are  adapted  to 
terrestrial  spore-distribution,  their  spores  being  set  free  on  the  destruction  of  the 
sporangium-wall  (generally  shot  out)  and  distributed  through  the  air.  For 
further  examples  of  spore-distribution  see  below,  p.  91-93. 

The  reproduction  of  Fungi  is  accomplished  not  only  by  spores 
and  conidia,  but  also  sometimes  by  chlamydospores.  These  are 
developed  in  joints  of  the  hyphae  by  the  aggregation  of  the  proto- 
plasm into  masses  which  become  surrounded  by  a  cell-wall  and 
take  on  a  resting  condition  in  the  form  of  a  spore,  and  are  able 
to  germinate  and  produce  carpophores.  In  the  formation  of  the 
chlamydospores  the  hyphaa  accumulate  reserve  materials  at  the 
expense  of  the  neighbouring  cells  ;  in  the  undivided  hyphee  of  the 
Phycomycetes  transverse  walls  are  formed,  and  finally  the  chlamy- 
dospores are  set  free  by  the  decay  of  the  empty  cells  connecting 
them  with  the  mycelium.  One  must  distinguish  between  oidia 
and  true  chlamydospores.  The  former  are  merely  spore-like  cells 
which  are  cut  off  from  the  ends  of  hyphae  (Figs.  162,  184),  and 
which  serve  for  propagation  in  the  same  manner  as  spores.  In 
Chlamydomucor  racemosus  the  chlamydospores  grow  out  into  the 
air  and  form  differentiated  carpophores.  In  the  Autobasidiomy- 
cetes  they  only  germinate  vegetatively,  and  not  with  the  forma- 
tion of  fructifications.  From  Chlamydomucor  up  to  the  Auto- 
basidiomycetes  the  successive  development  of  the  fructification, 


FUNGI.  91 

which  is  interrupted  by  the  formation  of  the  chlamydospores, 
degenerates  more  and  more.  Among  certain  Ustilagineoe  the 
chlamydospores  (brand-spores)  no  longer  germinate  with  the  pro- 
duction of  fructifications.  In  the  Uredinaceae,  only  one  of  the  three 
chlamydospore-forms  has  the  property  of  producing  fructifications 
on  germination ;  the  other  forms  only  germinate  vegetatively,  like 
ordinary  spores,  and  in  the  same  manner  as  the  chlamydospores 
of  the  Autobasidiomycetes.  In  the  Hemibasidii,  and  the  Uredin- 
acese,  in  Protomyces,  the  chlamydospores  are  the  chief  means  of 
reproduction.  They  are  found  also  among  the  Ascomycetes. 

The  sporangia  and  the  conidia  of  the  Fungi  have  their  common 
origin  in  the  sporangia  of  the  Phycomycetes.  The  asci  (and 
the  Ascomycetes  which  are  characterised  by  these  bodies)  are 
descended  from  the  sporangia-forming,  lower  Fungi  ;  the  basidia 
(and  the  Basidiornycetes)  from  those  which  bear  conidia.  The 
sporangia  of  the  Phycomycetes"  are  the  primitive  form  and  the  start- 
ing point  for  all  the  reproductive  forms  of  the  Fungi.  The  chlamy- 
dospores appear  besides  in  all  classes  of  Fungi  as  supplementary 
forms  of  reproduction,  and  are  of  no  importance  in  determining 
relationships.  Although  the  expression  "  fruit "  must  essentially 
be  applied  to  true  Phanerogams,  yet,  through  usage,  the  term 
"fruit-forms,"  is  employed  to  designate  the  forms  or  means  of 
reproduction  of  Fungi,  and  the  organs  of  reproduction  are  known 
as  organs  of  fructification,  the  sporangiophores  and  conidiophores 
as  fruit-bearers  (carpophores},  and  the  sporangiocarps,  conidiocarps, 
and  basidiocarps  as  "-fruit-bodies." 

The  majority  of  Fungi  have  more  than  one  method  of  reproduction,  often  on 
various  hosts  (Uredinaceas).  Species  with  one,  two,  or  more  than  two  methods  of 
reproduction  are  spoken  of  as  having  monomorphic,  dimorphic,  or  pleornorphic 
fructification.  Monomorphic,  e.g.  the  Tuberaceae  ;  dimorphic,  Mucor,  Pipto- 
cephalis,  SaprolegniaceaB,  Penicillium  crustaceum  ;  pleomorphic,  Puccinia 
graminis,  Capnodium  salicinum  (in  the  last  species  there  are  five  methods  of 
reproduction :  yeast-like  conidia,  free  conidiophores,  conidiocarps  with  small 
and  large  conidia,  and  ascocarps). 

The  liberation  and  distribution  of  the  spores  and 
conidia.  The  spores  and  conidia,  on  account  of  their  small  size 
and  lightness,  are  spread  far  and  wide  by  currents  in  the  air,  but 
in  addition  to  this  method,  insects  and  other  animals  frequently 
assist  in  disseminating  them.  The  liberation  of  the  conidia  is 
occasionally  effected  by  the  complete  shrinking  away  of  the 
conidiophore,  but  more  frequently  by  abstriction  from  the  conidio- 
phores, either -by  their  gradually  tapering  to  a  point,  or  by  the 


92  FUNGI. 

dissolution  of  a  cross-wall  (generally  of  a  mucilaginous  nature). 
The  individual  links  of  conidia- chains  are  detached  from  one 
another  in  the  same  way,  or  often  by  means  of  small,  intercalary 
cells,  which  are  formed  at  the  base  of  the  individual  links,  and 
becoming  slimy,  dissolve  upon  the  maturity  of  the  spores. 
Special  contrivances  for  ejecting  the  spores  and  conidia  may  often 
be  found.  In  Peronospora  the  cylindrical  fruit-hyphee  in  the  dry 
condition  become  strap-shaped  and  also  twisted.  These  are  very 
hygroscopic,  and  the  changes  of  form  take  place  so  suddenly,  that 
the  spores  are  violently  detached  and  shot  away.  In  Empusa  a 
peculiar  squirting  mechanism  may  be  found  (Fig.  85).  Each  club- 
shaped  hypha  which  projects  from  the  body  of  the  fly,  bears  a 
conidium  at  its  apex  ;  a  vacuole,  which  grows  gradually  larger, 
is  formed  in  the  slimy  contents  of  the  hypha,  and  the  pressure 
thereby  eventually  becomes  so  great  that  the  hypha  bursts  at  its 
apex,  and  the  conidium  is  shot  into  tne  air.  By  a  similar  mechan- 
ism, the  spores  of  many  of  the  Agaricacese  are  cast  away  from 
the  parent-plants.  In  the  case  of  Pilobolus  (Fig.  84)  the  entire 
sporangium  is  thrown  for  some  distance  into  the  air  by  a  similar 
contrivance,  the  basal  region  of  the  sporangium  having,  by  the 
absorption  of  water,  been  transformed  into  a  slimy  layer  which 
is  readily  detached.  Sphcerobolus,  a  Gasteromycete,  has  a  small, 
spherical  fruit-body  (basidiocarp),  the  covering  of  which,  when 
ripe,  suddenly  bursts,  and  the  basidiospores  contained  in  it  are 
forcibly  ejected. 

The  spores  which  are  enclosed  in  asci  are,  in  some  instances,  set 
free  from  the  mother-cell  (ascus)  prior  to  their  complete  develop- 
ment (Elaphomyces,  Eurotiuni).  In  the  case  of  the  majority  of  the 
Pyrenomycetes  and  Truffles,  the  asci  swell  by  the  absorption  of 
water  into  a  slimy  mass,  which  gradually  disappears,  so  that  the 
spores  lie  free  in  the  fruit-body ;  they  either  remain  there  till  the 
fruit-body  decays,  as  in  those  which  have  no  aperture  (Perisporiaceas, 
Tuberaceee),  or  the  slimy  mass,  by  its  growth,  is  forced  out  through 
the  aperture  of  the  sporocarp,  taking  the  spores  with  it  (Nectria). 
The  ejection  of  the  spores  by  mechanical  means  takes  place  in  a 
number  of  Ascomycetes,  and  should  many  spores  be  simultane- 
ously ejected,  a  dust-cloud  may  be  seen  with  the  naked  eye  to 
arise  in  the  air  from  the  fruit-body.  This  is  the  case  in  the  larger 
species  of  Peziza,  Helvella,  Rhytisma,  when  suddenly  exposed  to  a 
damp  current  of  air.  A  distinction  is  drawn  between  a  simultaneous 
ejection  of  all  the  spores  contained  in  the  ascus,  and  an  ejection  at 


FUNGI.  93 

intervals  (successive),  when  only  one  spore  at  a  time  is  thrown  out. 
The  first  of  these  methods  is  the  most  frequent,  and  is  brought 
about  by  the  ascus  being  lined  with  a  layer  of  protoplasm,  which 
absorbs  water  to  such  a  degree  that  the  elastic  walls  are  extended 
at  times  to  double  their  original  size.  The  spores  are  forced  up 
against  the  free  end  of  the  ascus,  a  circular  rupture  is  made  at 
this  point,  and  the  elastic  walls  contract,  so  that  the  fluid  with 
the  spores  is  ejected.  Special  means  may  in  some  instances  be 
found  to  keep  the  spores  together,  and  compel  their  simultaneous 
ejection.  Thus,  a  tough  slime  may  surround  all  the  spores  (Sac- 
cobolus},  or  a  chain- apparatus,  similarly  formed  of  tough  slime; 
or  there  may  be  a  hooked  appendage  from  each  end  of  the  spores 
which  hooks  into  the  appendage  of  the  next  spore  (Sordaria).  The 
paraphyses  occurring  between  the  asci  in  many  Ascomycetes,  also 
play  a  part  in  the  distribution  of  the  spores,  by  reason  of  the 
pressure  they  exercise.  The  asci  in  some  of  the  Pyrenomycetes, 
which  are  provided  with  jar-shaped  fruit-bodies,  elongate  to  such 
an  extent  that,  without  becoming  detached  from  their  bases,  they 
reach  the  mouth  of  the  fruit-body  one  at  a  time,  burst  and  disperse 
their  spores,  and  so  make  room  for  those  succeeding.  An  ejection 
of  the  spores  at  intervals  from  the  ascus  is  rarer.  It  takes  place, 
for  instance,  in  Pleospora,  whose  asci  have  a  double  wall.  The 
external  wall,  by  absorption  of  water,  at  last  becomes  ruptured, 
and  the  internal  and  more  elastic  membrane  forces  itself  out  in 
the  course  of  a  few  seconds  to  one  of  two  or  three  times  greater 
length  and  thickness,  so  that  one  spore  after  another  is  forcibly 
ejected  from  a  narrow  aperture  at  the  end  of  the  ascus. 

Germination  of  spores  (conidia  and  chlamydospores).  In 
many  spores  may  be  found  one  or  more  germ-pores,  i.e.  thinner 
places,  either  in  the  inner  membrane  (uredospores.  Sordaria) 
or  in  the  external  membrane  (teleutospores  in  Rust-Fungi), 
through  which  the  germination  takes  place.  Generally  this  does 
not  occur  till  the  spores  have  been  set  free  :  in  some  Ascomycetes 
germination  commences  inside  the  ascus  (Taphrina,  Sclerotinia) . 
The  different  ways  in  which  the  spores  germinate  maybe  classified 
into  three  groups. 

I.  THE  ORDINARY  GERMINATION  occurs  by  the  spore  emitting  a 
germ-tube,  which  immediately  developes  into  a  mycelium.  In 
spores  with  a  double  wall  it  is  only  the  inner  membrane  which 
forms  the  germ-tube.  In  swarmspores  a  single  wall  is  formed 
after  the  withdrawal  of  the  cilia,  and  this,  by  direct  elongation, 


94  FUNGI. 

becomes  the  germ-tube.  The  protoplasm  accumulated  in  the 
spore  enters  the  hypha,  which,  in  pure  water,  can  only  grow  as 
long  as  the  reserve  nourishment  lasts. 

2.  GERMINATION  WITH  PROMYCELIUM  differs  only  by  the  circum- 
stance that  the  hypha  developed  from  the  germ-tube  has  a  Tery 
limited  growth,  and  hence  it  does  not  immediately  develope  into  a 
mycelium,  but  produces  conidia  (Rust-  and  Brand-Fungi).     This 
promycelium  must  only  be  regarded  as  an  advanced  development 
of  a  conidiophore  or  basidium. 

3.  THE  YEAST-FORMATION  of  conidia  consists  in    the   production 
of  outgrowths,  very  much  constricted  at  their  bases,  from  one  or 
more  places.     Each  of  the  conidia  formed  in  this  manner  may 
again  germinate  in  the  same  way.     When  sufficient  nourishment  is 
present,  a  branched  chain  of  such  conidia  is  formed,  and  these  are 
iiiially  detached  from  one  another.     Yeast-like  buddings  from  the 
conidia  are   produced  in  various  Fungi,  e.g.  Ascoidea,  Protomyces, 
Ustilaginece,  Ascomycetes,  Tremellacese,  etc.     In  the  Ustilaginese 
these    conidia    are    an   important   element   in    the    development. 
The  budding  conidia  of  Exobasidium  forms  a  "  mould "   on  the 
nutritive    solution.     The    yeast-like  conidia    are   not   to    be   con- 
founded with  the  "  Mucor-yeast "  (comp.  Mucoraceae).     For  Sac- 
charomijces  see  Appendix  to  the  Fungi,  page  176. 

In  a  compound  spore  (i.e.  when  a  mass  of  spores  are  associated 
together)  each  spore  germinates  on  its  own  account.  There  are 
sometimes,  however,  certain  among  them  which  do  not  germinate, 
but  yield  their  contents  to  those  which  do. 

The  length  of  time  for  which  conidia  can  retain  their  power  of 
germination  is  shortest  (being  only  a  few  weeks)  in  those  having 
thin  walls  and  containing  a  large  supply  of  water  (Peronosporaceae, 
Uredinaceae).  In  many  spores  a  resting  period  is  absolutely  neces- 
sary before  they  are  able  to  germinate  (resting  spores).  It  has 
been  observed  in  some  spores  and  conidia,  that  the  faculty  of 
germinating  may  be  preserved  for  several  years  if  the  conditions 
necessary  for  germination  remain  absent  (Ustilagineae,  Eurotium, 
Penicillium). 

The  optimum,  minimum  and  maximum  temperatures  required 
for  the  germination  of  the  spores  has  been  decided  in  the  case  of 
a  good  many  Fungi.  A  large  portion  of  the  most  common  Fungi 
have  their  optimum  at  20°C.,  minimum  at  1-2°C,  maximum  at 
40°C.  In  the  case  of  pathogenic  Fungi  the  optimum  is  adapted  to 
the  temperature  of  the  blood.  Fungi  living  in  manure,  whoso 


ZYGOMICETES.  95 

spores  are  often  adapted  to  germinate  in  the  alimentary  canals  of 
warm-blooded  animals,  have  an  optimum  corresponding  to  the 
temperature  of  these  animals,  but  with  a  little  margin. 

Systematic  Division. — The  lowest  class  of  the  Fungi  is  that 
of  the  PHYCOMYCETES,  which  have  an  unicellular  mycelium,  sexual 
and  asexual  reproduction,  and  have  doubtless  sprung  from  spo- 
rangia-bearing, lower  Green  Algse.  From  the  Phycomycetes  (and 
certainly  from  the  Zygomycetes)  spring  two  well  defined  branches, 
each  with  numerous  distinct  species;  to  the  one  branch  belong  the 
HEMIASCI  and  the  ASCOMYCETES,  to  the  other  the  HEMIBASIDII  and 
the  BASIDIOMYCETES.  Ascomycetes  and  Basidiomycetes  may  be 
united  under  the  title  of  MYCOMYCETES  or  HIGHER  FUNGI.  The 
Hemiasci  and  the  Hemibasidii  constitute  the  class  of  MESOMYCETES. 
The  Hemiasci  are  an  intermediate  form  between  Zygomycetes  and 
Ascomycetes ;  the  Hemibasidii  a  similar  group  between  the  Zygo- 
mycetes and  Basidiomycetes.  Mesomycetes  and  Mycomycetes  have 
only  asexual  reproduction  ;  sexual  reproduction  is  wanting.  Their 
mycelium  is  multicellular. 

Up  to  the  present  time  about  39,000  species  have  been  described. 

Review  of  the  divisions  of  the  Fungi : — 

class  L— Phycomycetes  (Algal-Fungi). 
Sub-Class  1.    Zygomycetes. 
Sub-Class  2.    Oomycetes. 

Family  1.     ENTOMOPHTHORALES. 
Family  2.     CHYTRIDIALES. 
Family  3.     MYCOSIPHONALES. 

class  ii.   Mesomycetes. 

Sub-Class  1.    Hemiasci. 

Sub-Class  2.    Hemibasidii  (Brand-Fungi). 

Class  in.— Mycomycetes  (Higher  Fungi). 

Sub-Class  i.    Ascomycetes. 
Series  1.     Exoasci. 
Series  2.     Carpoasci. 

Family  1.     GYMNOASCALES.     -\ 

Family  2.     PERISPORIALES.      £  Angiocarpic  Exoasci. 

Family  3.     PYRENOMYCETES.  J 

Family  4.     HYSTERIALES.     1    TT      • 

J  \  Herm-angiocarpic  Exoasci. 

Family  5.     DISCOMYCETES.  ) 

Family  6.     HELVELLALES.     Gymnocarpic  (?)  Exoasci. 
Additional :  ASCOLICHENES.     Lichen-forming  Ascomycetes. 
W.B.  H 


96  ZYGOMYCETES. 

Sub-Class  2.    Basidiomycetes. 
Series  1.— Protobasidiomycetes.       Partly    gymnocarpic,    partly 

angiocarpic. 

Series  2.     Autobasidiomycetes. 
Family  1.    DACRYOMYCETES.    Gymnocarpic. 
Family  2.     HYMENOMYCETES.    Partly  gymnocarpic,   partly  hemi- 

angiocarpic. 

Family  3.    PHALLOIDEJE.     Hemiangiocarpic. 
Family  4.     GASTEROMYCETES.     Angiocarpic. 
Additional :  BASIDIOLICHENES.    Lichen-forming  Basidiomycetes. 
Additional  to   the  Fungi :    FUNGI  IMPEBFECTI.       Incompletely   known 
(Saccliaromyces,  Oidium-iorms,  etc.). 


Class  1.    Phycomycetes  (Algal-Fungi).1 

This  group  resembles  Vaucheria  and  the  other  Siphonese  among 
the  Alg83. 

ORGANS  OF  NUTRITION.  The  mycelium  is  formed  of  a  single  cell, 
often  thread-like  and  abundantly  branched  (Fig.  78).  Vegetative 
propagation  by  chlamydospores  and  oidia.  Asexual  reproduction 
by  endospores  (sometimes  swarmspores)  and  conidia.  Sexual  re- 
production by  conjugation  of  two  hyphae  as  in  the  Conjugate,  or 
by  fertilisation  of  an  egg-cell  in  an  oogonium.  On  this  account 
the  class  of  the  Phycomycetes  is  divided  into  two  sub-classes : 
ZYGOMYCETES  and  OOMYCETES. 

Sub-Class  I.    Zygomycetes. 

Sexual  reproduction  takes  place  by  zygospores,  which  function 
as  resting-spores,  and  arise  in  consequence  of  conjugation  (Fig.  81); 
in  the  majority  of  species  these  are  rarely  found,  and  only  under 
special  conditions.  The  most  common  method  of  reproduction  is  by 
endospores,  by  acrogenous  conidia,  by  chlamydospores,  or  by  oidia. 
Swarmspores  are  wanting.  Parasites  and  saprophytes  (order  6  and  7). 
The  zygospores  are  generally  produced  when  the  formation  of 
sporangia  has  ceased;  e.g.  by  the  suppression  of  the  sporangial- 
hyphce  (Mucor  mucedo},  or  by  the  diminution  of  oxygen;  Pilobolus 
cnjstallinus  forms  zygospores,  when  the  sporangia  are  infected 
with  saprophytic  Piptocephalis  or  Pleotrachelus. 

A.     Asexual  reproduction  only  by  sporangia. 

Order  1.  Mucoraceae.  The  spherical  sporangia  contain  many 
spores.  The  zygospore  is  formed  between  two  unicellular  branches 
(gametes). 

1  Also  termed  Water-Fungi  (Wasserpilzen). 


ZYGOMYCETES.  97 

The  unicellular  mycelium  (Fig.  78)  of  the  Mucoraceae  branches 
abundantly,  and  lives,  generally,  as  a  saprophyte  on  all  sorts  of 
dead  organic  remains.  Some  of  these  Fungi  are  known  to  be  cap- 
able of  producing  alcoholic  fermentation,  in  common  with  the  Sac- 
charomyces.  This  applies  especially  to  Chlamydomucor  racemosus 
(Mucor  racemosus},  when  grown  in  a  saccharine  solution,  and  de- 
prived of  oxygen;  the  mycelium,  under  sucli  conditions,  becomes 
divided  by  transverse  walls  into  a  large  number  of  small  cells. 


FIG.  78. — Mucormucedo.  A  mycelium  which  has  sprung  from  one  spore,  -whose  position 
is  marked  by  the  *  :  a,  b,  c  are  three  sporangia  in  different  stages  of  development ;  a  is 
the  youngest  one,  as  yet  only  a  short,  thick,  erect  branch ;  b  is  commencing  to  form  a 
sporangium  which  is  larger  in  c,  but  not  yet  separated  from  its  stalk. 

Many  of  these  swell  out  into  spherical  or  club-shaped  cells,  and 
when  detached  from  one  another  become  chlamydospores,  which 
abstrict  new  cells  of  similar  nature  (Fig.  79).  These  chlamydo- 
spores were  formerly  erroneously  termed  "  mucor-yeast,"  but  they 
must  not  be  confounded  with  the  yeast-conidia  (page  94).  They 
are  shortened  hyphae,  and  are  not  conidia  of  definite  size,  shape, 
and  point  of  budding.  Oidia  are  also  found  in  Chlamydomucor. 


98 


ZYGOMYCETES. 


,  The  Mucoracese,  in  addition  to  the  chlamydospores  and  oidia, 
have  a  more  normal  and  ordinary  method  of  reproduction;  viz.r 
by  spores  which  are  formed  without  any  sexual  act.  Mucor  has- 
round  sporangia ;  from  the  mycelium  one  or  more  long  branches,, 
sometimes  several  centimetres  in  length,  grow  vertically  into  the 
air ;  the  apex  swells  (Figs.  78,  80)  into  a  sphere  which  soon 
becomes  separated  from  its  stalk  by  a  transverse  wall ;  in  the 

interior  of  this  sphere  (spor- 
angium) a  number1  of  spores  are 
formed  which  eventually  are  set 
free  by  the  rupture  of  the  wall. 
The  transverse  wall  protrudes 
into  the  sporangium  and  forms 
the  well-known  columella  (Fig. 
80  d,  e).  The  formation  of 
spores  takes  place  in  various 
ways  among  the  different 
genera. 

SEXUAL  EEPEODUCTION  by  conjugation  takes  place  in  the  follow- 
ing manner.  The  ends  of  two  hyphte  meet  (Fig.  81)  and  become 
more  or  less  club-shaped ;  the  ends  of  each  of  these  are  cut  off 
by  a  cell- wall,  and  two  new  small  cells  (Fig.  81  A)  are  thus 
.formed,  these  coalesce  and  give  rise  to  a  new  cell  which  becomes 
the  very  thick- walled  zygote  (zygospore),  and  germinates  after  a» 


PIG.  79.— Chlamydospores  of  CTiZamydo* 
mucor  racemosus  ( x  375  times.) 


FIG.  80.— JJfitcor  mucedo :  a  a  spore  commencing  to  germinate  (  x  300  times);  b  a  germi- 
nating spore  which  has  formed  a  germ-tube  from  each  end  ( x  300  times)  ;  c  the  apex  of  a 
young  sporangium  before  the  formation  of  spores  has  commenced;  the  stalk  is  protruded 
in  the  sp  orangium  in  the  form  of  a  column  :  on  the  wall  of  the  sporangium  is  found  a  very 
fine  incrustation  of  lime  in  the  form  of  thorn-like  projections  ;  d  a  sporangium  in  which 
the  formation  of  spores  has  commenced  ;  e  a  sporangium,  the  wall  of  which  is  ruptured,, 
leaving  a  remnant  attached  to  the  base  of  the  columella  as  a  small  collar.  A  few  spores  are- 
seen  still  adhering  to  the  columella. 


ZYGOMYCETES. 


99 


period  of  rest,  producing  a  new  hypha,  which  bears  a  sporangium 
(Fig.  81  JS). 

Mucor  mucedo,  Pin- 
mould,  resembles  some- 
what in  appearance 
Penicillium  crustaceum 
and  is  found  growing 
upon  various  organic 
materials  (bread,  jam, 
•dung,  etc.). 

Pilobolus  (Figs.  83, 
84)  grows  on  manure. 
Its  sporangium  (Fig.  84 
a")  is  formed  during  the 
night  and  by  a  peculiar 
mechanism  (page  92)  is 
shot  away  from  the  plant 
in  the  course  of  the  day. 
This  generally  takes 
place  in  the  summer, 
between  eight  and  ten 
a.m.  The  sporangium 
is  shot  away  to  a  height 
which  may  be  300  times 
greater  than  that  of  the 
plant  itself,  and  by  its 
stickiness  it  becomes 
attached  to  portions  of 
plants,  etc.,  which  are 
in  the  vicinity.  If  these 
are  eaten  by  animals, 
the  spores  pass  into  the 
alimentary  canal  and 
are  later  on,  sometimes 
even  in  a  germinating 
condition,  passed  out 
with  the  excrement,  in 
which  they  form  new 
mycelia. 

Phycomyces  nitens 
("  Oil-mould ")  is  the 
largest  of  the  Mould  Fungi ;  its  sporangiophores  may  attain  the  height  of  10-30  c.  m. 

Order  2.  Rhizopaceae.  Ehizopus  nigricans  (Mucor  stolonifer)  which  lives 
on  decaying  fruits  containing  sugar,  on  bread,  etc.,  has,  at  the  base  of  the 
sporangiophores,  tufts  of  rhizoids,  i  e.  hyphae,  which  function  as  organs  of 
attachment.  From  these,  "  runners  "  are  produced  which  in  a  similar  manner 
develope  sporangiophores  and  rhizoids. 

Order  3.     Thamnidiaceae.     On  the  same  sporangiophore,  in  addition  to  a 


FIGS.  81,  82.  — afucor  mucedo:  A-C  stapes  in  the  formation  of 
the  zygote ;  D  zygote ;  E  germination  of  zygote :  the  exospore 
has  burst,  and  the  endospore  grown  into  a  bypha  bearing 
a  sporangium. 


100 


OOMYCETES. 


large,  terminal,  many-spored  sporangium,  many  smaller,  lateral  sporangia  are 
formed  with  a  few  spores.     Thamnidium. 

B.  Asexual  reproduction  by  sporangia  and  conidia. 

Order  4.     Choanephoraceae.     Choanephora  with  creeping  endophytic  my- 
celium, and  perpendicular  sporangiophores. 

Order  5.     Mortierellaceae.     Mortierella  polycephala  produces  on  the  same 
mycelium    conidia    and    sporangiophores.      M. 
rostafinskii  has  a  long   stalked  sporangiophore, 
which  is  surrounded  at  its  base  by  a  covering  of 
numerous  felted  hyphae. 


FIG.  83.—  PiZobolus.  Mycelium  (a,  a), 
with  a  sporangiophore  (A)  and  the 
fundament  of  another  ( B). 


FIG.  81.— Pilololus.  Sporangium  (a")  with 
stalk  (a-c),  which  is  covered  by  many 
small  drops  of  water  pressed  out  by  tur- 
gescence. 


C.  Asexual  reproduction  only  by  conidia. 

Order  6.  Chsetocladiacese.  The  conidia  are  abstricted  singly  and  aero- 
genously.  Chcetocladium  is  a  parasite  on  the  larger  Mucoraceas. 

Order  7.  Piptocephalidacese.  The  conidia  are  formed  acrogenously  and 
in  a  series,  by  transverse  divisions.  The  zygospore  arises  at  the  summit  of  the 
conjugating  hyphse,  which  are  curved  so  as  to  resemble  a  pair  of  tongs.  'Pipto- 
cephalis  and  Syncephalis  live  parasitically  on  the  larger  Mucorace®. 

Sub-Class  2.    Oomycetes. 

Sexual  reproduction  is  oogamous  with  the  formation  of  brown, 
thick-walled  oospores  which  germinate  after  a  period  of  rest. 
Asexual  reproduction  by  conidia  and  swarmspores.  Parasites, 
seldom  saprophytes. 

The  oospores  are  large  spores  which  are  formed  from  the  egg- 
cell  (oosphere)  of  the  oogonium  (oosporangium,  Fig.  89,  95).  A 
branch  of  the  mycelium  attaches  itself  to  the  oogonium  and  forms 
at  its  apex  the  so-called  "  antheridium  "  (pollinodium1)  :  this  sends 
one  or  more  slender  prolongations  (fertilising  tubes)  through  the 
wall  of  the  oogonium  to  the  egg-cell. 

1  Antheridium  is  preferred  in  this  sub-class  as  keeping  amore  uniform  term  (Kn). 


OOMYCETES. 


101 


A  fertilisation,  a  passage  of  the  contents  of  the  antheridium  to  the  egg-cell, 
has  as  yet  only  been  observed  in  Pythium;  in  Phytophthora  only  one  small 
mass  of  protoplasm  passes  through  the  fertilising  tube  to  the  egg-cell;  in  Pero- 


FIG.  85. — Empusn  muscce  (Fly-mould).  I.  A  fly  killed  by  the  fungus,  surrounded  by  a  white 
layer  of  conidia.  II.  The  conidiophores  (t)  projecting  from  the  body  of  the  fly.  Some  of 
the  conidia,  a  few  of  which  have  developed  secondary  conidia,  are  attached  to  the  hairs 
(mag.  80  times).  III.  A  perfect  hypha.  IV.  A  hypha  in  the  act  of  ejecting  a  conidinm 
(c),  enveloped  in  a  sticky  slime  (g).  V.  A  conidium  which  has  developed  a  secondary 
conidium(sc).  VI.  A  branched  hypha  produced  by  cultivation.  VII.  A  secondary  con- 
idium which  has  produced  a  small  mycelium  (m).  VIII.  A  conidinm  germinating  on  the 
fly's  body.  IX.  Mycelium.  X.  Conidia  germinating  like  yeast  in  the  fatty  tissue  of  the 
fly.  (III.-VII.  and  IX.  magnifled  300  times  ;  vm.  and  X.  magnified  500  times.) 


102  OOMYCETES. 

nospora  and  the  Saprolegniaceae'no  protoplasm  can  be  observed  to  pass  through 
the  fertilising  tube,  so  that  in  these  instances  parthenogenesis  takes  place ; 
Saprolegnia  thuretii,  etc.,  have  generally  even  no  antheridia,  but  nevertheless 
form  normal  oospores.  Fertilisation  of  the  egg-cell  by  means  of  self-motile 
spermatozoids  is  only  found  in  Monoblepharis  sphcerica. 

A.  Asexual  reproduction  by  conidia  only. 

Family  1.     Entomophthorales. 

The  mycelium  is  richly  branched.  The  family  is  a  transitional 
step  to  the  conidia-bearing  Zygomycetes,  since  the  oospores  of  many 
members  of  this  family  arise,  and  are  formed,  like  zygospores. 

Order  1.  Entomophthoraceae.  Mycelium  abundantly  de- 
veloped. This  most  frequently  lives  parasitically  in  living  insects, 
causing  their  death.  The  conidiophores  forming  the  conidial- 
layer  project  from  the  skin,  and  abstrict  a  proportionately  large 
conidium  which  is  ejected  with  considerable  force,  and  by  this 
means  transferred  to  other  insects.  These  become  infected  by 
the  entrance  of  the  germ-tube  into  their  bodies.  The  spherical, 
brown  resting-spores  develope  inside  the  bodies  of  insects  and 
germinate  by  emitting  a  germ- tube. 

GENERA  :  Empusa  has  a  good  many  species  which  are  parasitic  on  flies, 
moths,  grasshoppers,  plant-lice.  The  conidia  emit  a  germ-tube  which  pierces 
the  skin  of  the  insect ;  a  number  of  secondary  conidia  are  then  produced  inside 
its  body,  by  division  or  by  gemmation  similar  to  that  taking  place  in  yeast,  each 
of  which  grows  and  becomes  a  long  unbranched  hypha,  and  these  eventually 
fill  up  the  body  of  the  animal,  causing  distension  and  death.  Each  of  these 
hyphae  projects  through  the  skin,  and  abstricts  a  conidium,  which  is  ejected 
by  a  squirting  contrivance.  The  best  known  species  is  E.  muscce  (Fig.  85), 
which  makes  its  appearance  epidemically  towards  autumn  on  the  common 
house-fly,  and  shows  itself  by  the  dead  flies  which  are  found  on  the  windows 
and  walls  attached  by  their  probosces,  distended  wings,  and  legs.  They  have 
swollen  abdomen,  broad  white  belts  of  hyphse  between  the  abdominal  rings,  and 
are  surrounded  by  a  circle  of  whitish  dust  formed  by  the  ejected  conidia. — 
Entomophthora  sends  out,  at  definite  places,  from  the  mycelium  hidden  in  the 
insect's  body,  bundles  of  hyphae,  which  serve  the  purpose  of  holding  fast  the 
dead  insects,  the  ramifications  attaching  themselves  to  the  substratum :  the 
conidiophores  are  branched,  the  conidia  are  ejected  by  the  divisional  walls  be- 
tween the  hyphaa  and  the  conidia  dividing  into  two  layers,  those  which  ter- 
minate the  hyphae  suddenly  expanding  and  throwing  the  conidia  into  the  air. 
E.  radicans  makes  its  appearance  epidemically  on  caterpillars. 

B.  Asexual  reproduction  by  zoospores  or  conidia. 

Family  2.     Chytridiales. 

In  this  family  the  mycelium  is  very  sparsely  developed  or  is 
wanting.  The  entire  plant  consists  principally  or  entirely  of  a 


OOMYCETES.  103 

single  zoosporangium  whose  zoospores  have  generally  one  cilium. 
The  resting-spores  arise  either  directly  from  the  zoosporangium, 
which,  instead  of  forming  zoospores,  surrounds  itself  by  a  thick 
cell-wall ;  or  they  are  formed  by  the  conjugation  of  two  cells  (in 
which  case  they  are  spoken  of  as  oospores).  Microscopic  Fungi, 
parasitic  on  water  plants  (especially  Algee)  or  small  aquatic  ani- 
mals, seldom  on  land  plants. 

Order  1.  Olpidiaceae.  Without  mycelium.  Swarmspores 
and  resting-spores. 

In  the  Olpidiez,  the  swarmspores,  probably,  most  frequently  form  themselves 
into  a  plasmodium  (naked  mass  of  protoplasm)  which  may  become  a  single 
zoosporangium  or  a  resting  sporangium.  Olpidium  trifolii  occurs  in  Trifolium 
repens. — In  the  Synchytriece  the  plasmodium  emerging  from  the  swarmspores 
breaks  up  either  at  once,  or  after  a  period  of  rest,  into  smaller  plasmodia,  each 
of  which  will  become  a  zoosporangium.  Synchytrium  anemones  is  found  on 
Anemone  nemorosa ;  S.  mercurialis  on  Mercurialis  perennis ;  S.  aureum  on  many 
plants,  particularly  Lysintachia  nunimularia. 


FIG.  86. — Chytridium  lagenula.    Zoo-  FIG.  87.— Obelidium  mucronatum : 

sporangium  a  before,  b  after  the  libera-  m  mycelium ;  s  swarmspores. 

tion  of  the  swarmspores. 

Order  2.  Rhizidiaceae.  Mycelium  present.  Zoospores  and 
resting-spores. 

Chytridium  (Fig.  86).  Obelidium  (Fig.  87)  is  bicellular ;  the  one  cell  is  the 
mycelium,  the  other  the  zoosporangium ;  found  on  insects.  The  species  of 
Cladochytriiim,  are  intercellular  parasites  on  marsh  plants.  Physoderma. 

Order  3.  Zygochytriacese.  Mycelium  present.  Zoospores 
and  oospores.  The  latter  are  the  product  of  the  conjugation  of 
two  cells  (Fig.  88). 

Polyphagus  euglence  on  Euglena  viridis.  Uroplilyctis  pulposa  on  species  of 
Chenopodinm.  (Edomyces  leproides  on  the  Beet-rooc. 


104  OOMYCETES. 

Family  3.     Mycosiphonales. 

The  mycelium  is  bladder-like  or  branched.  Zoospores.  Sexual 
reproduction  by  oospores,  which  are  produced  in  oogonia.  The 
latter  are  fertilised,  in  some  forms,  by  the  antheridium. 

Order  1.  Ancylistacese.  The  entire  bladder-like  mycelium  is  used  for  the 
construction  of  zoosporangia,  oogonia,  or  antheridia.  Lagenedium  is  parasitic 
on  Spirogyra,  etc. 

Order  2.  Peronosporaceae.  Almost  entirely  parasites.  The 
unicellular,  often  very  long  and  abundantly  branched  mycelium 
lives  in  the  intercellular  spaces  of  living  plants,  especially  in 
the  green  portions,  and  these  are  more  or  less  destroyed  and  de- 
formed in  consequence.  Special  small  branches  (suction-organs, 


FIG.  88. — Polyphagus  euglence.    A  with  smooth,  B  with  thorny  oospores ;  m  and /the  two 
conjugating  cells. 

"  haustoria ")  are  pushed  into  the  cells  in  order  to  abstract 
nourishment  from  them.  Both  oospores  and  conidia  germinate 
either  immediately,  or  they  develope  into  sporangia  with  swarm- 
spores,  having  always  two  cilia.  Only  one  oospore  is  formed  in 
each  oogonium  ;  its  contents  (Fig.  89)  divide  into  a  centrally 
placed  egg-cell  and  the  "  periplasm  "  surrounding  it;  this  is  of  a 
paler  colour  and  on  the  maturity  of  the  oospore  forms  its  thick, 
brown,  external  covering. 

The  Potato-fungus  (Phytophthora  infestans)  is  of  great  interest. 
Its  thallus  winters  in  the  Potato-tuber;  other  organs  for  pass- 
ing the  winter,  such  as  oospores,  are  not  known.  When  the  tuber 
germinates,  the  Fungus-hyphse  penetrate  the  young  shoot  and 
keep  pace  with  the  aerial  growth  and  development  of  the  plant. 
The  conidiophores  emerge  through  the  stomata,  especially  on  the 
under  side  of  the  leaves ;  they  branch  like  a  tree  (Fig.  90),  and 


OOMYCETES. 


105 


appear  to  the  naked  eye  as  a  fine  mould  on  the  surface  of  the 
plant.  The  disease  soon  makes  itself  known  by  the  brown  colour- 
ing of  those  parts  of  the  plant  which  are  attacked,  and  by  their 
withering.  An  ovoid  conidium  arises  at  first  by  the  formation  of 
a  dividing-  wall  at  the  apex  of  each  branch  of  the  conidiophore 


FIG.  89. — Peronospora.  aJsiiiearutn.    My 
celiam  with  egg-cell  and  antheridium. 


infestans  (strongly  magnified).    Cross  section  through  a  small 
rtion  of  a  Potato-leaf  (the  under  side  turned  upwards) :  a  the  mycelium ;  b  b  two  conidio- 
hores  projecting  through  a  stoma;  c  conidia;  e  the  spongy  tissue  of  the  leaf;    g  the 
pifiannis. 


106 


OOMYCETES. 


FIG.  91. — PhytopMliora  infestans :  a-c  conidia  de- 
tached ;  in  c  the  swarm-cells  are  leaving  the  mother- 
cell;  d  two  free-swimming  swarm-cells. 


(Fig.  90  c  c),  and  immediately  underneath  it  another  is  formed, 
which  pushes  the  first  to  one  side,  and  so  on.  These  conidia  some- 
times germinate  directly,  and  form  a  mycelium,  but  most  frequently 
their  protoplasm  divides  into  many  small  masses,  each  of  which 
becomes  a  pear-shaped  zoospore  provided  with  two  cilia  (Fig.  91). 
Water  is  required  for  their  germination,  and  when  the  ripe  conidia 

are  placed  in  a  drop  of 
water  the  swarm-cells  are 
formed  in  the  course  of 
about  five  hours.  They 
swarm  about  in  rain  and 
dewdrops  in  the  Potato- 
fields,  and  are  carried  with 
the  water  to  the  Potato- 
plants  and  to  the  tubers 
in  the  soil.  The  wind  also 
very  easily  conveys  the  conidia  to  healthy  Potato  fields  and  infects 
them.  The  enormous  quantity  of  conidia  and  swarm-cells  that 
may  be  formed  in  the  course  of  a  summer  explains  the  rapid 
spreading  of  the  disease ;  and  the  preceding  makes  it  clear  why 
wet  summers  are  favourable  to  its  existence.  When  the  swarm- 
cells  germinate,  they  round  off,  and  then  surround  themselves 
with  a  cell-wall  which  grows  out  into  the  germ-tube,  and  pian's 
through  the  epidermis  of  the  host-plant  (Fig.  92).  Having  entered 

the  host,  a  new  mycelium 
is  formed.  The  potato 
disease,  since  1845,  has 
been  rampant  in  Europe  ; 
it  has,  no  doubt,  been  in- 
troduced from  America, 
which,  it  must  be  remem- 
bered, is  the  home  of  the 
Potato-plant. 

The  conidia  exhibit  various 
characters  which  are  employed 
for  the  separation  of  the  genera. 
Pythium  is  the  most  simple 
form.  The  contents  of  the  ter- 
minally-formed conidia  emerge  as  a  spherical  mass  and  divide  into  swarmspores. 
P.  de  Baryanum  lives  in  the  seedlings  of  many  different  Flowering-plants, 
which  it  completely  destroys. — Phytophthora  is  distinguished  by  the  circum- 
stance that  the  sparsely-branched  conidiophores  bear,  sympodially,  chains  of 


FIG.  92.— PUytopUliora  infestans.  Cross  section 
through  a  portion  of  a  Potato-stalk.  Two  germinat- 
ing conidia  (a,  b)  piercing  the  epidermis,  and  the 
mycelium  penetrating  the  cells. 


OOMYCETES. 


107 


conidia.  Besides  the  Potato-fungus  (see  above),  Ph.fayi  belongs  to  this  group  ; 
it  developes  oospores  very  abundantly,  and  does  great  harm  to  seedlings  of  the 
Beech,  Sycamore,  and  Pine  trees. — Peronospora  generally  has  conidiophores 
which  are  repeatedly  forked,  and  bear  a  conidium  on  each  of  the  most  extreme 
ramifications.  Many  do  great  harm  to  their  host-plants.  P.  viticola,  on  Vines, 
and  P.  nivea,  on  umbelliferous  plants,  have  swarmspores,  which  are  absent  in 
the  following  species  of  this  genus  :  P.  sparsa,  on  Eoses ;  P.  gangliformis,  on 
composites  ;  P.  alsinearum,  on  Stitchwort ;  P.  parasitica,  on  cruciferous  plants  ; 
P.  vicice,  on  Vetches  and  Peas;  P.  schachtii,  on  Beets;  P.  violacca,  on  the 


FIG.  93.— A  fly  overgrown 
with  Saprolegnia. 


FIG.  94.— Formation,  of  swarmspores  in  a  Saprolegnia : 
a  germinating  swarmspores. 


flowers  of  Scabiosa ;  P.  radii,  on  the  ray-florets  of  Matricaria. — Cystopus  (Albugo) 
has  the  conidia  developed  in  chains,  which  form  a  cohesive  white  layer  under- 
neath the  epidermis  of  the  host-plant.  Cystopus  candidus,  on  cruciferous  plants, 
especially  Shepherd's  Purse  and  Brassica ;  the  germination  commences  on  the 
cotyledons,  and  from  this  point  the  mycelium  developes  together  with  the  host- 
plant  ;  C.  cubicus,  on  the  leaves  of  Composite. 

Order  3.     Saprolegniacese,  Water-Fungi  which  live  as  sapro- 
phytes on  organic  remains  lying  in  water,  for  instance,  on  dead 
flies  (Fig.  93),  worms,  remains  of  plants ;  but  they  may  also  make 
their  appearance  on  living  animals,  being 
frequently    found,    for    example,    on    the 
young  trout  in  rearing  establishments. 

The  thallus  is  a  single,  long  and  branched 
cell.  It  has  one  portion  which  serves  as 
root,  and  lives  in  the  substratum,  where  it 
ramifies  abundantly  for  the  purpose  of  ab- 
sorbing nourishment ;  and  another  portion 
projecting  freely  in  the  water,  and  sending 
out  hyphje  on  all  sides  (Fig.  93).  The 
asexual  reproduction  takes  place  by  swarm- 
spores (Fig.  94),  which  are  developed  in 
large  sporangia;  these  swarmspores  gener- 
ally possess  two  cilia,  and  on  germination  FlG-  95.-Oogonium  with 

two  antheridia,  Achlyarace- 

grow  into  new  plants.     The   entire  proto-      mesa. 


108  MESOMYCETES. 

plasm  in  the  oogonium  is  formed  into  one  or  more  oospheres, 
without  any  surrounding  "  periplasm."  The  oospheres  may  not 
be  fertilised  (p.  100),  and  then  develope  parfchenogenetically. 

Genera  :  Saprolegnia,  whose  swarmspores  disperse  immediately  after  having 
left  the  sporangium.  S.  ferax  is  the  cause  of  a  disease  in  fish  ("Salmon 
disease  ")  and  in  the  crayfish. — Achlya,  whose  swarmspores  accumulate  in  a 
hollow  ball  before  the  mouth  of  the  sporangium. — Leptomitus  has  strongly  in- 
dented hyphae,  causing  a  "  linked  "  appearance.  L.  lacteus  is  frequent  in  the 
waste  matter  from  sugar  factories. — Monoblepharis  deviates  from  the  others  by 
the  greater  development  of  its  fertilising  process ;  the  oosphere,  situated  in  an 
open  oogonium,  becoming  fertilised  by  self-motile  spermatozoids,  which  are 
provided  with  a  cilium  at  the  posterior  end. 

Class  2.    Mesomycetes. 

The  Mesomycetes  are  intermediate  forms  between  the  Phy- 
comycetes  and  the  Higher  Fungi.  In  the  vegetative  organs,  and  in 
the  multicellular  hyphae,  they  resemble  the  Higher  Fungi;  the 
methods  of  reproduction,  however,  show  the  characters  of  the  Phy- 
comycetes,  namely  sporangia  and  conidiophores  of  varying  size 
and  with  varying  number  of  spores ;  definite  and  typically  formed 
asci  and  basidia  are  not  present.  Sexual  reproduction  is  wanting. 
The  HEMIASCI  are  transitional  between  the  Phycomycetes  and  the 
Ascomycetes,  the  HEMIBASIDII  (Brand-Fungi)  form  the  transition 
to  the  Basidiomycetes. 

Sub-Class  1.     Hemiasci. 

The  Hemiasci  are  Fungi  with  sporangia  which,  although  resem- 
bling asci,  yet  have  not,  however,  a  definite  form  and  a  definite 
number  of  spores.  Besides  endospores,  conidia,  chlamydospores 
and  oidia  are  found. 

Order  1.  Ascoideaceae.  Ascoidea  rubescens  forms  irregular,  reddish-brown 
masses  in  the  sap  issuing  from  felled  Beeches.  It  has  free  sporangia,  which 
resemble  asci  in  their  structure,  in  the  development  and  ejection,  and  in  the 
definite  shape  and  size  of  the  spores.  The  formation  of  the  sporaugia  takes 
place  when  the  nutriment  is  nearly  exhausted,  and  resembles  that  of  the  conidia, 
since  they  are  developed  from  the  end  of  a  hypha  which  enlarges,  and  the  swelling 
becomes  separated  by  a  transverse  wall.  Within  the  sporangia  numerous  spores  of 
a  cap-like  form  are  developed,  which  are  set  free  through  an  opening  at  the  apex. 
Sporangia  are  formed  successively  at  the  apex  of  the  same  hypha,  the  second 
commencing  to  develope  as  the  first  is  dehiscing.  Conidia  and  sporangia  are  not 
formed  simultaneously  ;  the  former  may  be  considered  as  closed  sporangia. 

Order  2.  Protomycetaceae.  Protomyces  pacJiydcrmus  causes  hard  swellings 
•on  the  stems  a  ad  leaf -stalks  of  the  Cichoriese  (Taraxacum,  etc.).  These  swell- 


HEMIBASIDII. 


109 


ings  consist  of  chlamydospores  (resting-spores),  which  germinate  and  become 
free,  ascus-like  sporangia,  with  numerous  small  spores.  In  nutritive  solutions 
the  chlamydospores  form  conidia  with  yeast-like  buddings.  P.  macrosporus  on 
jEgopodium,  and  other  Umbelliferae. 

Order  3.  Thelebolaceae.  Thelebolus  stercoreus,  is  found  on  the  dung  of 
deer,  hares,  and  rabbits,  and  has  closed  sporangia,  which  resemble  asci  in  their 
shape  and  regular  construction,  and  in  the  ejection  of  spores.  The  covering 
encloses  only  one  sporangium,  even  where  the  sporangia  arise  close  together. 

This  order,  by  reason  of  the  covering  of  the  sporangia,  forms  the 
transition  from  the  Hemiasci  to  the  Carpoasci,  while  the  two  first 
supply  an  intermediate  step  to  the  Exoasci. 


Sub-Class  2.     Hemibasidii,  Brand-Fungi. 

The  Brand-Fungi  (also  known  as  USTILAGINE^)  are  Fungi  with 
basidia-like  conidiophores,  which,  however,  have  not  yet  advanced  to 
a  definite  form  or  number  of  conidia.  They  are  true  parasites, 
whose  mycelium  spreads  itself  in  the  intercellular  spaces  of  Flower- 


FIG.  96. — Entyloma  ranunculi.  1.  Cross  section  of  a  portion  of  a  leaf  of  Ficaria  permeated 
by  the  mycelium ;  a  bundle  of  hyphee  with  conidia  emerging  from  a  stoma  ;  in  one  of  the 
cells  are  found  four  brand-spores.  2.  A  brand-spore  developed  in  the  middle  of  a  hypha. 

ing  plants.  The  mycelium  is  colourless,  quickly  perishable,  has 
transverse  walls  at  some  distance  from  each  other  (Fig.  96),  and 
sends  out  haustoria  into  the  cells  of  the  host-plant. 

It  most  frequently  happens  that  the  germ-tube  enters  the  host- 
plant  at  its  most  tender  age,  that  is,  during  the  germination  of  the 
seed  ;  the  mycelium  then  wanders  about  in  the  tissues  of  the  shoot 


110 


HEMTBASID1I. 


during  its  growth,  until  it  reaches  that  part  of  the  plant  where 
the  spores  are  to  be  formed.  The  spore-formation  takes  place  in 
the  same  way  in  all  those  species  whose  brand-spores  are  deve- 
loped in  the  floral  parts  of  the  host-plant.  Many  Brand-Fungi  have, 
however,  a  more  local  occurrence,  and  the  mycelium  is  restricted 
to  a  smaller  area  of  the  leaf  or  stem.  Those  organs  of  the  host- 
plant  in  which  the  brand-spores  are  developed  often  become 
strongly  hypertrophied.  In  perennial  plants  the  mycelium  winters 
very  often  in  the  rhizome. 

The  brand-spores  are  the  winter  resting-spores  of  the  Brand- 
Fungi.  They  arise  in  the  tissues  of  the  host-plant,  which  is  often 
destroyed,  and  become  free  through  the  rupture  of  the  epidermis ; 


FIG.  97. — Doassansia  alismatis.  1.  A  fruit-body,  formed  by  a  covering  of  oblong  hyphae, 
which  encloses  a  mass  of  brand-spores,  and  is  embedded  in  the  leaf-tissue  of  the  Lost- 
plant  ;  20  times  natural  size.  2.  A  germinating  brand-spore,  500  times  natural  size.  3. 
Three  connected  resting-spores,  400  times  natural  size.  4.  Two  conidia  grown  together, 
GOO  times  natural  size. 

they  are  thick-walled,  generally  brown  or  violet,  and  very  often 
possess  warts',  spines,  or  reticulate  markings.  Fruit-bodies,  that  is 
enclosed  organs  of  reproduction,  are  found  in  few  genera  (Sphace- 
lotheca,  Graphiola ;  Doassansia,  Fig.  97).  In  Tolyposporium,  Tiibur- 
cinia,  Thecaphora  (Fig.  102),  etc.,  the  brand-spores  are  united  into 
a  ball  of  spores.  On  germination  the  brand-spores  behave  as 
chlamydospores,  namely,  as  the  foundation  of  conidiophores,  by 
emitting  a  short  germ-tube,  i.e.  a  conidiophore  ("promycelium"). 
The  TJsTiLAGiNACEJ]  (Fig.  99,  2)  have  a  short  transversely  divided  co- 
nidiophore, with  Laterally  developed  conidia  (comp.  the  basidia  of 
the  Protobasidiomycetes).  The  conidiophores  of  the  TILLETIACEJ; 
are  undivided  (unicellular  promycelia),  and  bear  the  conidia  ter- 
minally, and  so  resemble  the  basidia  of  the  Autobasidiomycetes. 


HEMIBASIDII. 


Ill 


In    Tilletia,   Entyloma,  Neovossia,  Tuburcinia,  the   brand-spores   germinate 
and   form  basidia-like  conidiophores  with  spindle-shaped  conidia ;  their  my- 


FIG.  98,—Tubttrcinia.  1.  T.  trientalie.  Hyphse,  some  of  which  bear  conidia  at  the  apex, 
forcing  themselves  out  between  the  epidermal  cells  on  the  under  side  of  the  leaf;  320  times 
natural  size.  2.  T.  trientalis.  A  ball  of  spores  in  which  some  of  the  individual  brand- 
spores  are  about  to  germinate  ;  520  times  natural  size.  3.  T.  primulicola  :  various  forms 
of  conidia  (500  times  natural  size). 

celium,  on  the  other  hand,  produces  later  only  single,  sickle-shaped  conidia,  so 
that  two  kinds  of  conidia  are  found,  as  in  a  few  Basidiomycetes.  In  some 
species,  e.g.  Ustilago  hordei,  the 
brand-spores  only  germinate 
vegetatively  and  form  a  myce- 
lium. In  nutritive  solutions  (so- 
lutions of  dung,  etc.)  where  they 
live  as  saprophytes,  the  brand- 
spores  of  many  species  emit 
germ-tubes,  and  on  these,  yeast- 
like  conidia  are  produced  by 
repeated  budding,  which  grow 
into  mycelia  only  when  the 
nutritive  solution  is  exhausted. 
These  conidia  have  not  the 
power  of  producing  alcoholic 
fermentation .  The  very  numer- 
ous conidia,  which  are  found 
in  the  dung  of  herbivorous  ani- 
mals, are  probably  the  yeast- 
conidia  of  Brand-Fungi.  The 
brand-spores,  which  are  eaten 
by  animals  with  the  grain  and 
hay,  pass  into  the  dung  and 
without  doubt  give  rise  to  a 
very  rich  multiplication  of 
yeast-conidia. 

The     conidia      (also     called 
w.  R. 


FIG.  99.  —  Ustnago.  1.  Formation  of  brand- 
spores.  2.  Germinating  brand-spore  of  U.  peren- 
nans.  3.  Germinating  brand-spore  of  U.  cardui 
(after  Brefeld).  4.  U.  filiformis.  a  A  brand-spore 
with  developed  basidium  ;  b  another,  with  a  coni- 
dium;  c  with  two  conidia;  d  with  two  conidia 
placed  diametrically  opposite  to  each  other ;  e,  de- 
tached conidia  which  are  growing  into  hyphse. 

I 


112 


HEMIBASID1I. 


"sporidia")  of  many  species  unite   generally  into   an   H-form  (Figs.  97,  4; 
100  h  ;  101,  4).     This  union  in  pairs  does  not,  however,  take  place  with  a  view 


FIG  100.— Tilletia  tritici :  a  an  ear  of  Wheat  in  which  all  the  grains  are  attacked  by  Stink- 
brand  ;  b  a  blighted  corn  surrounded  by  the  chnff ;  c  a  blighted  corn  grown  together  with  a 
stamen;  d  the  same  cutacross  ;  e  a  brand-spore ;  /,  g,  7i  germinating  brand-spores ;  i  germi- 
nating conidia ;  j  the  mycelium  ;  7c-fc  brand-spore-l'orming  mycelium-threads,  (c-h  magni- 
fied 400  times ;  i-fc  300  times.) 


HEMIBASIDII.  113 

to  germination,  there  is  no  fusion  of  nuclei,  and  therefore  in  this  "  fusion  "  there 
is  no  sexual  act. 

Order  1.  Ustilaginacese.  Conidiophores  with  transverse  walls  and  lateral 
conidia. —  Ustilago  (Fig.  99)  generally  developes  its  spores  in  the  floral  organs 
of  its  host-plant,  the  ovary  or  anthers,  where  they  arise  from  hyphae,  and  form 
&  slimy  mass  which  when  mature  becomes  a  black  dust. 

To  this  order  belong  U.  avence,  parasitic  on  Oats,  V.  hordei  and  U.  nuda 
{U.  jenseni),  on  Barley;  these  are  the  usual  cause  of  "  Smut"  on  cereals.  U. 
Jiypodytes  on  straw  of  Elymus  and  Agropyrum.  U.  filiformis  in  the  leaves  of 
Glyceria.  U.  caricis  transforms  the  fruits  of  various  species  of  Car  ex  into 
black,  dusty  balls.  U.  violacea  developes  its  violet  spore-powder  in  the  anthers 
of  the  Caryophyllaceae.  U.  trugoppgonin,  transforms  entire  inflorescences  of 
Tragopogon  into  a  black-violet  mass.  Among  the  largest  are  U.  grandis,  which 
•causes  the  large  swollen  nodes  in  the  stem  of  Phragmites,  and  the  Maize  Blight, 
U.  tnm/dis,  which  produces  outgrowths  about  the  size  of  a  hand  on  the  spadix  of 
the  Maize. 

Order  2.  Tilletiaceae.  •Conidiophores  undivided,  generally  several  conidia 
arise  at  their  apices. — Tilletia  tritici,  the  Stinkbiand  on  Wheat  (Fig.  100).  The 
mycelium  lives  in  Wheat-plants,  producing  its  spores  in  the  ovary  after  the 
whole  interior  of  this  body  has  been  destroyed  by  the  mycelium,  with  the  ex- 
ception of  the  external  layer  of  the  wall  of  the  ovary,  which  remains  essentially 
unaltered  and  encloses  the  closely  packed,  firm  mass  of  spores  (Fig.  100  d). 
The  grains  of  Wheat  thus  attacked  are  shorter  and  thicker  than  the  sound  ones, 
and  the  ears  show  the  presence  of  this  Fungus  by  their  erect  position,  and 
the  wide  separation  of  the  chaff  (Fig.  100  a).  The  unpleasant  odour  of  the 
ovary  prior  to  the  ripening  of  the  spores,  has  given  the  name  "  Stinkbrand," 
and,  in  like  manner,  its  hardness  when  it  encloses  the  ripe  spores,  is  the  reason 
of  its  beiug  also  called  "  Stonebrand."  On  account  of  this  hardness,  the  dis- 
•eased  grains  are  readily  harvested  together  with  the  healthy  ones,  which  become 
infected  by  the  spores  at  the  threshing.  T.  lavis  (T.  fastens)  also  occurs  on 
Wheat  and  has  smooth  brand-spores. 

Entyloma  (Fi^.  96),  a  genus  with  numerous  species,  which  appear  in 
spots  on  the  leaves  of  the  host-plant,  and  Tuburcinia  (Fig.  98),  which  makes 
its  appearance  on  the  Primu- 
laceae,  produce  white  conidia- 
spots  on  the  surface  of  the 
host-plant.  The  first-named 
has  single  spores,  the  latter 
has  its  spores  closely  massed 
together.— Urocystis  (Fig.  101) 
has  its  spores  surrounded  by 
a  number  of  small  and  lighter 
coloured  barren  spores.  U. 
occulta,  Eye-stem  Blight,  Fi&.lOl.— Urocystis.  1,  U.coralloides.  A  spore-ball, 

forms     its      spores     in     long        magnified  450  times.     2-4,  U.  anemones:  2-3,  brand- 
streaks    in    the    stems    and       sP°res  which  are  about  to  terminate  (magnified  450 
.,.•«,  T    i  times).    4,  Conidia,  the  two  in  a  state  of  fusion,  a 

leaves  of  the  Eye,  and  does       third  with  yacuolea  and  division-wall,  magnified  600 
•considerable  damage.     U.  ce-      times. 


114  ASCOMYCETES. 

pulcR  on  Onions.  U.  liola  forms  large  dark-violet  swellings  in  the  leaf-stalk 
and  stems  of  Violets.—  Thecaphora  (Fig.  102)  appears  in  seedlings  of  Con- 
volvulus and  Astragalus. 

As  a  means  of  protection  against  the  Smut -Fungi  which  make  their  appear- 
ance on  the  different  cereals,  a  submersion  of  the  grains  in  a  solution  of  blue 
vitriol  (J%)  for  twelve  hours,  or  better  still,  submerging  for  five  minutes  in 
water  heated  to  53-55°  C  (Jensen's  method)  is  employed. 


Fro.  10?.— Tliecaplora.  1,  T.  convolvuli,  a  ball  of  spores, one  of  the  brand-spores  has  emitted 
a  fceptate  branched  conidiophore  ( x  520).    2,  T.  affmis,  a  ball  of  spores  (x  520). 

Class  3.    Mycomycetes,  Higher  Fungi. 

The  MYCOMYCETES  are  not  entirely  aquatic  in  habit  ;  they  have- 
hyphse  with  transverse  walls',  sexual  reproduction  in  only  few 
cases.  The  asexual  reproduction  takes  place  in  very  different 
ways;  by  endospores  (in  asci),  conidia,  basidiospores,  chlaniy- 
dospores,  and  oidia.  Swarmspores  are  never  found. 

Two  chief  methods  of  reproduction  may  be  distinguished,  and 
hence  the  class  may  be  divided  into  two  large  sub-classes  : — the 
ASCOMTCETES  (with  asci),  and  the  BASIDIOMYCETES  (with  basidia). 

Sub-Class  1.    Ascomycetes. 

The  main  characteristic  which  distinguishes  the  Ascomycetes  is- 
the  ascus ;  a  name  given  to  a  sporangium  of  a  definite  shape  and 
size,  and  containing  a  definite  number  of  spores.  The  shape  is. 


ASCOMYCETES. 


115 


generally  club-like  or  spherical,  the  number  of  spores  8  (in  some 
2,  4,  16  or  more),  see  Figs.  103,  105,  108,  110,  113,  116,  120,  121, 
123,  129. 

In  the  lowest  forms,  the  EXOASCI,  the  ascus  springs  directly  from 
the  mycelium  without  the  formation  of  a  fruit-body  (i.e.  asco- 
carp).  In  the  higher  forms,  which  contain  many  species,  the 
CARPOASCI,  the  asci  are  united  and  form  ascocarps  which  may  be 
more  or  less  enclosed  (angiocarpic,  hemiangiocarpio,  and  probably 
gymnocarpic). 


FIG.  103,— Endogenous  formation  of  spores  in  Peziza  confluens.  In  the  youngest  asci 
there  is  only  one  nucleus  (b,  e) ;  this  divides  into  two  (/);  and  the  division  is  repeated  so 
that  there  are  4  nuclei  in  c  and  8  in  g.  These  surround  themselves  with  protoplasm  and  a 
cell-wall  (7i,  i).  The  protoplasm  of  the  mother-cell  is  not  entirely  used  up. 

The  hyphse  of  the  Mycelium  in  some  remain  free,  in  others 
they  are  felted  together  and  form  thick  strands  or  flat,  cushion-like 
bodies  (compare  in  particular  the  stromata  of  the  Pyrenomycetes). 
Some  species  form  sclerotia  (Figs.  116,  128). 

Asexual  reproduction  by  means  of  conidia  is  known  in  many 
species  as  the  principal  means  of  reproduction,  and  the  one 
which  affords  the  most  rapid  means  of  distribution.  The  conidia 
may  be  produced  on  conidiophores  (Fig.  109),  in  conidial -layers 
(Fig.  122),  and  often  in  conidiocarps  (pycnidia,  Figs.  120  d,e; 


116 


ASCOMYCETES 


123  a  :  124  &).     The  conidiophores  bear  no  analogy  to  the  basidia. 
[The  question  of  the   occurrence  of  sexual  reproduction  among 
the  Ascomycetes  has  been  much  disputed.     Eepently  Harper  has 
proved  the  fusion  of  a  male  and  female  nucleus  in  Sphcerotheca 
(see  page  120),  and  also  in  Pyronema,  a 
Fungus  closely  allied  to  Peziza.     In  Py- 
0  ronema  both  the  oogonium  and  antheri- 
dium     contain     several     nuclei.        The 
**  former  is  a  large  cylindrical  cell  having 
the  apex  prolonged   into    a    trichogyne? 
which  is  also  multinucleate.      The  latter 
is    a  cylindrical    cell    which    originates 
below  the    oogonium,    its    apex    coming 
into    close    touch    with    the    tip   of    the 
trichogyne.       The    intervening   wall    is 
absorbed   at  the   point   of    contact,  &nd 
the    male    nuclei     then    pass    into    the 
trichogyne,   and      finally,    entering    the 
FIG.  102A.— Sphcerotheca  castag-  oogonium  by  the  breaking  down  of  the 

net:   A    Ooeronium  (o)  and  anthe-  ..  '    i    •  • 

ridium  (a);jBthetrneantheridium  separating  wall,  the  male  nuclei  conjugate 
in  pairs    with  the  female  nuclei   of   the 
oogonium.       The    oogoniumis    now  cut 
a  new  cell-wall    formed   at  the 

Harperf  enclosing  base  of  the  trichogyne,  and  sends  forth 
ascogenous  hyphee  containing  the  conju- 


Fio.  102s. — Pyronema  confluent :  A  Rosette  of  three  pairs  of  sexual  cells  drawn  from 
surface  view;  B  section  ot!  an  oogonium,  ascogenous  liyphse  containing  nuclei  growing 
out  from  the  ooifonium ;  new  partition  wall  has  formed  at  tlie  base  of  the  trichogyne;  C 
semi-diagramatic  drawing  of  a  section  of  an  ascocarp  in  which  the  first  asci  are  ripening. 
a  Antheridium ;  og  oogonium;  as/  ascogenous  hypnte ;  ascasci;  t  trichogyne. 


EXOASCI.  116  A 

gated  nuclei.     These  hyphee  branch,  and  ultimately  bear  the  asci  at 
their   extremities,   the  paraphyses  arising  beneath  the  ascogone.] 

In  many  species  the  ascospores  germinate  and  form  conidia  immediately 
(Nectria  cinnabarina,  Sclerotinia,  Taphrina,  etc.),  sometimes  while  they  are 
still  in  the  ascus  and  before  their  ejection  (Taphrina,  Fig.  105  a).  In  many  in- 
stances the  conidia  by  means  of  continued  budding  can,  for  a  longer  or  shorter 
time,  produce  yeast-conidia,  e.g.  Taphrina.  In  many  other  cases  the  conidia 
arise  from  the  germ-tubes  of  the  ascospores,  or  at  any  part  of  the  mycelium. 
The  unripe  asci  of  Taphrina,  when  placed  in  water,  develope  conidia  at  their 
apices.  The  ticlerotinia-  species  produce  numerous  conidia  whose  germination 
has  never  been  observed.  The  formation  of  conidia  and  asci  sometimes  takes 
place  on  the  same  fruit-body.  In  Heterospharia  patella  the  conidia  and  asci 
are  developed  successively  in  the  same  fruit-body  ;  in  the  ascocarps  of  Derma- 
tea  Jrungula  and  Sclerotinia  sclerotiorum  the  formation  of  conidia  may  take 
place.  The  ascocarps  frequently  arise  from  the  conidial-layers  (Nectria  cin- 
uabarina,  etc.).  This  relationship  of  the  two  forms  of  reproduction  to  each 
other  may  be  explained  by  considering  that  both  have  descended  phylogene- 
tically  from  sporangia. 

Sometimes  chlamydospores  and  oidia  also  appear  in  the  Ascomy- 
cetes  ;  on  germination,  however,  they  do  not,  as  in  Protomyces, 
form  sporangia,  and  on  this  account  cannot  be  distinctly  distin- 
guished from  conidia. 

The  asci  are  morphologically  the  highest  form  of  reproduction 
and  are  always  found  at  the  close  of  the  development  of  these 
Fungi  ;  the  accessory  forms  of  reproduction  are  first  developed,  but 
a  well-defined  alternation  of  generations  does  not  occur. 

In  the  Ascomycetes  there  are  more  than  11,000  described  species,  which  can 
be  classed  as  follows  : — 

Series  1.    EXOASCI.     Only  one  order. 
,,      2.     CARPOASCI. 

Family  1.     Gymnoascales,  -\ 

„        2.     Perisporiales,    >  Angiocarpic  Carpoasci. 
,,        3.     Pyrenomycetes,) 

„       4.     Hysteriales,    |  Hemiangiocarpic  Carpoasci. 
N        5.    Discomycetes,  ) 

„        6.     Helvellales,  Gymnocarpic  (?)  Carpoasci. 
Additional  Ascolichenes :  Lichen-forming  Ascomycetes. 


Series  1.     Exoasci. 

Ascomycetes  with  FREE  ASCI;  sometimes  also  conidia,  chlamydo- 
spores and  oidia.  One  order. 

Order.  Taphrinaceae.  Of  the  genera  belonging  to  this  order, 
Taphrina,  Endomyces,  and  Ascocorticium,  the  first  is  most  important. 


. 


EXOASCI. 


117 


Endomyces  decipiens  is  a  parasite  in  the  fruit-body  of  Armillaria  mellea ; 
E.  magmisii  lives  in  the  gelatinous,  fermenting  exudations  of  Oak-trees ;  Asco- 
corticium  albidum,  is  found  under  the  bark  of  the  Fir-tree.  Endomyces  has 
chlamydospores  and  oidia. 

The  species  of  Taphrina  are  parasites,  whose  free  asci  may  be 
found  in  great  numbers,  generally  closely  pressed  together,  on 
the  parts  of  plants  which  they  have  att  icked.  The  asci  are  de- 
veloped directly  from  the  ascogenous  cells  of  a  fertile,  generally 
sub-cuticular,  hypha,  which  arises  from  the  sterile  mycelium.  The 
latter  arises  from  the  germinating  ascospore,  and  may  hibernate 
in  the  tissues  of  its  host,  particularly  in  the  winter  buds,  and  then 
with  the  commencement  of  the  next  period  of  vegetation  it  con- 


FIG.  104.  —  Taphrina  (Exoascus) 
pruni.  Yeast-like  budding  of  a 
germinating  spore  (  x  600). 


FIG.  105.— Taphrina  betulina:    a  ascus  filled 
with  conidia;  b  germinating  spores  (x600). 


tinues  its  growth  side  by  side  with  that  of  its  host.  The  hyphge 
ramify  in  the  intercellular  spaces  or  beneath  the  cuticle,  but  have 
no  haustoria.  The  ascospores  (Fig.  105  A)  and  unripe  asci  may 
produce  conidia. 

Very  remarkable  appearances,  and  swellings  of  the  attacked  tissues,  are  pro- 
duced when  the  mycelium  is  perennial;  for  example,  the  "  Witches'  -brooms  " 
and  "  Pockets."  The  hard,  hollow,  stoneless  plums,  known  as  "  Pocket  "  or 
"  Bladder  "  Plums,  are  produced  by  considerable  changes  in  the  tissues  of  the 
fruit ;  these  are  caused  particularly  by  T.  pruni  on  several  species  of  Prunus. 
The  "  Witches'-brooms,"  on  the  contrary,  are  deformations  of  entire  twigs  or 
branches,  and  often  attain  a  very  large  size.  They  occur  on  Alnus  incana, 
caused  by  T.  epiphylla  ;  on  Carpinu*  betulus,  by  T.  carpini ;  on  Cherjy-trees, 
by  T.  cerasi ;  on  Plum-trees,  by  T.  insitities ;  on  Birches,  by  T.  tuvyida  and 
T.  betulina.  T.  deformam  attacks  the  leaves  of  the  Peach,  and  causes  them 
to  curl. 


118  CARPOASCI. 

When  a  perennial  mycelium  is  wanting,  the  infection  is  confined  as  a  rule  to 
white  or  yellow  spots  on  the  leaves,  e.fj.  the  commonest,  T.  sadebcckii,  on  Ainu* 
gliititw*at  and  T.  aurea  on  species  oiPopulus.  T.  <alni  incanae  (Fig.  106)  causes 
considerable  hypertrophies  on  the  pistillate  catkins  of  the  Alder,  which  may  be 
compared  to  the  "  pockets  "  of  Prunus. 


FIG.  106. — TapTirina  alni  incance  on  the  Aliler  (nat.  size). 

Series  2.     Carpoasci. 

The  Carpoasci  are  Ascomycetes,  whose  asci  are  enclosed  in  fruit- 
bodies,  i.e.  ascocarps.  The  accessory  melans  of  reproduction  are 
free  conidiophores  (Fig.  109),  conidial-layers  (Fig.  122),  conidio- 
carps  (Fig.  120  D,  E,  etc.),  chlamydospores  and  oidia. 

For  the  different  methods  of  distributing  the  ascospores,  see  p. 
92. 

Of  the  six  families  of  the  Carpoasci,  the  first  three — Gymnoa- 
scales,  Perisporiales,  and  Pyrenomycetes — are  ANGIOCARPIC  (that  is, 
the  ascocarp  remains  closed  throughout  its  existence,  and  does 
not  dehisce  when  ripe)  ;  the  fourth  and  fifth  families  (Hysteriales 
and  Discomycetes),  011  the  other  hand,  are  HEMIANGIOCARPIC  (the 
ascocarp,  here  also  called  an  apothecium,  is  closed  in  the  early 
stages,  but  opens  at  the  commencement  of  ripening  and  exposes 
a  hymenium  of  crowded  asci)  ;  the  family  of  Helvellales  has 
probably  GYMNOCARPIC  (or  hemiangiocarpic)  fruit-bodies. 

Family  1.     Gymnoascales. 

The  ascocarps  are  surrounded  by  a  spongy  and  incomplete  en- 
velope. One  order,  poor  in  species. 


CAEPOASCT.  119 

Order  Gymnoascaceae. — The  ascocarps  are  borne  sometimes  solitarily,  or 
sometimes  coiled  together.  Gymnoascus  'reessii  forms  small  bodies  about  1 
mm.  in  diameter  on  old  horse-dung,  which  at  first  are  white  and  afterwards 
•orange-red. — Ctenomyces  serratus  lives  on  the  old  feathers  in  birds'  nests. 

Family  2.     Perisporiales. 

The  ascocarps  are  surrounded  by  a  complete  envelope  without  any 
opening :  the  fruit-bodies  are  cleistocarpic ;  the  spores  are  only 
liberated  after  the  disintegration  of  the  fruit-bodies.  Paraphyses 
are  wanting.  The  two  first  orders  have  in  addition  the  means 
of  reproduction  by  conidia. 

Order  1.  Erysiphacese,  Mildews.  The  Fungi  belonging 
to  this  order  are  epiphytic  parasites,  whose  mycelium,  somewhat 
resembling  a  cobweb,  may  be  seen  on  the  leaves  and  other 
green  portions  of  plants  (see  Figs.  107,  108).  The  hyphre  ramify 


FIG.  107.— Erysiphe  cichoriacearum :  a  mycelium-threads ;  l>  antheridium ;  c  oogoniam ; 
d  and  e  young  ascocarps. 

in  all  directions  upon  the  surface  of  their  host,  and  emit  haustoria 
which  penetrate  the  epidermal  cells,  and  thus  derive  the  necessary 
nutriment.  The  Mildew-Fungi  thus  belong  to  the  obligate  para- 
sites, and  during  their  growth  dwarf  and  destroy  the  portions 
of  their  host  on  which  they  live.  The  reproduction  takes  place  in 
the  first  instance  by  abstriction  of  conidio-chains  from  the  end  of 
special  branches  (Fig.  108  c,  a  hypha  is  seen  in  the  act  of  detaching 
a  conidium).  The  conidia  may  germinate  immediately,  and  thus 
quickly  reproduce  their  species.  When  present  in  large  numbers 
they  appear  as  a  white  ineal  covering  the  surface  of  the  plant  on 
which  the  fungus  is  found.  Later  on  appear  the  dark  brown, 
spheroid  ascocarps  (Fig.  108  a)  which,  although  small,  are  gene- 
rally just  visible  to  the  naked  eye  as  black  specks. 

A  characteristic  feature  of  the  Mildew-Fungi  is  the  thin,  pseudo- 


120 


CAEPOASCI. 


parenchymatous  covering  of  the  ascocarp,  enclosing  one  (Podo- 
sphcvra  and  Sphcerotheca ;  compare  Thelebolus  among  the  Hemi- 
asci)  or  a  few  asci  (Fig.  108  c),  which  do  not  form  any  hymenium, 
but  are  irregularly  placed.  The  cells  of  the  ascocarp-envelope 
are  often  prolonged  into  hair-like  appendages.  The  ascocarps  are 
developed  from  the  mycelium  at  places  where  two  hyphre  cross 
each  other  (Fig.  107).  At  these  places  two  short  and  erect  hyphse 
are  produced  side  by  side.  The  one  from  the  lower  hypha  (Fig. 
107  c)  assumes  an  ellipsoidal  shape,  and  is  cut  off  by  a  transverse 
wall  to  form  the  oogonium  •  -while  the  other  (6)  arches  over,  and 


FIG.  108. — Erysiplie  comm  urn's.  A  small  portion  of  a  leaf  with  this  Fungus  growing  upon, 
it  (ccrasi'ierably  magnified).  The  hypbae  b  and  d  do  not  belong  to  this  Fungus,  but  are  re- 
productive organs  of  a  pyrenomycetous  Fungus  parasitic  upon  it  (Cicinnobolas) . 

from  its  apex  the  true  antheridium  is  cut  off  (Fig.  102A).  The 
nucleus  from  the  antheridium  passes  into  the  oogonium,  and  unites 
with  its  nucleus.  After  fertilization  the  oogonium  becomes  multi- 
cellular,  and  is  known  as  the  ascogone.  In  Sphcerotheca  the  binu- 
cleate  penultimate  cell  produces  the  ascus,  but  in  other  genera  it 
branches  and  gives  rise  to  numerous  asci.  Hyphal  branches  from 
the  base  of  the  ascogone  surround  it  and  form  the  envelope  of  the 
ascocarp  (perithecia). 

Many  plants,  both  cultivated  and  wild,  are  attacked  by  various 


CARPOASCI. 


121 


species  of  Mildew.     A  common  means  of  prevention  against  their 
attacks  is  to  dust  the  diseased  parts  with  sulphur. 

Sphccrotheca  pannosa  occurs  on  the  leaves  of  Roses,  and  on  the  fruit  of 
Peaches  and  Apricots.  S.  castagnei  on  Humulus,  Cucumis,  etc. — Erysiphe  tuckeri 
grows  on  the  leaves  and  fruit  of  the  Vine  ;  it  spins  its  hyphae  over  the  bunches 
of  grapes,  curtails  their  growth,  and  causes  them  to  burst,  and  to  become  de- 


FIG.  109. — Eurotium  glaucum  :  a  portion  of  mycelium  lying  horizontally  ;  ft  vertically- 
placed  conidiophore ;  the  mycelium  gives  rise  to  another  branch  near  a  ;  the  conidia  are 
abstricted  from  short  flask-shaped  cells ;  b  a  ripe  conidiutn;  c,  d  germinating  conidia; 

spirally-twisted  hypha,  commencement  of  an  ascocarp  ;  /  a  stage  later  ;  g  still  later,  tlie 
hypha  at  the  base  of  the  coil  has  given  off  branches  which  are  applied  to  it;  7i,  i  sections  of 
young  ascocarps. 

cayed  and  rotten  (Grape-disease).  The  Fungus  was  first  noticed  in  England 
in  1845,  and  later  was  found  in  all  countries  whese  grapes  are  grown.  It  is  only 
known  in  the  conidial  form  ("  Oidium  tuckeri").  Many  other  species  of  Ery- 
siphe are  found  on  herbaceous  plants. — Microsphcera  has  appendages  which  are 
repeatedly  forked  at  their  extremities.  M.  grossularice  on  Eibes  grossularia. — 


122 


CAEPOASCI. 


Uncinula  has  appendages  with  spirally-coiled  extremities  ;  on  Salix  and  Acer. 
— Phyllactin<a  has  a  circle  of  bristle-like  appendages  with  dilated  bases.  P. 
guttata  on  Corylus,  Fraxinus,  Fagus,  etc. 

Order  2.  Perisporiaceae,  Moulds  and  Mildews.  A  group  of 
Fungi  widely  distributed  and  found  in  all  situations.  Usually  they 
have  a  well-developed  surface  mycelium,  and  small,  round,  seldom 
conspicuous  ascocarps,  containing  ovoid,  pulley-like  spores.  They 
are  partly  saprophytic,  partly  parasitic,  in  the  latter  condition 
having  a  brown  mycelium. 

Eurotium  glaucum  (  =  E.  herbariorum,  Figs.  109,  110)  and  E. 
repens  live  on  dead  organic  matter,  preserved  fruits,  etc.  The 
conidial  forms  of  both  species  are  known  as  "  Moulds  "  (Fig.  109), 
and  formerly  were  described  under  the  name  "Asperyillus  glaucus" 


o 


FIG.  110.— Euvottum  glaucum  :  a  longitudinal  section  of  a  half.ripe  nscocarp,  bounded 
externally  by  a  well-defined  layer  of  cells,  enclosing  asci  in  various  stages  of  develop- 
ment; b  a  semi- ripe,  c  nri  almost  ripe  ascus;  d  and  e  spores  seen  from  tbe  edge  and 
side  j  /germinating  spore  twenty-two  hours  after  been  sown  in  plum  juice. 

The  conidia  for  some  time  remain  attached  to  each  other  in 
chains  (Fig.  109  a) ;  they  are  abstricted  from  sterigmata  arranged 
radially  on  the  spherical,  swollen  end  of  the  conidiophore.  The 
small  yellow  or  brownish  ascocarps  are  frequently  found  in 
herbaria,  especially  when  the  specimens  have  been  insufficiently 
dried.  Euaspergillus  futnigatus  and  others  are  pathogenic,  causing 
mycosis  in  warm-blooded  animals. 

Penicillium  crustaceum  (P.  glaucum,  Figs.  Ill,  112)  is  an  ex- 
ceedingly common  "Mould."  Its  mycelium  appears  very  fre- 
quently on  any  organic  matter  which  is  permitted  to  remain 
untouched,  and  soon  covers  it  with  a  dense  mass  of  blue-green 


CARPOASC1.  123 

conidiophores.  These  branch  at  their  summits  and  bear  flask- 
shaped  cells  from  which,  the  conidia  are  abstricted.  The  ascocarps 
which,  both  in  size  and  colour,  resemble  grains  of  sand,  have  only 


FIG.  111. — Penicillium  crustaceum :  a  conidia  (x  300);  b  germination  of  conidia;  c  small: 
portion  of  mycelium,  produced  from  a  conidium  at  *,  with  five  conidiophores ;  d  young 
conidiophore  (x  630),  a  flask-shaped  cell  is  abstricting  a  conidium;  e  the  same  conidio- 
phore  after  9-10  hours. 


FIG.  112.—  Penicillium  crustaceum:  a  two  spirally-coiled  byphse  arise  from  the  my- 
celium, from  one  of  which  (ascogone)  the  asci  are  produced  ;  fc  a  further  step  in  the- 
development  of  the  ascocarp  ;  the  branching  ascogone  is  surrounded  by  sterile  hyphae ; 
c  section  of  young  ascocarp ;  the  larger  hyphaa  in  the  centre  are  the  ascogenous. 
hypha?;  these  are  enclosed  by  a  pseudo-parenchyma  of  sterile  hyphas  (x  390);  d  series 
of  ripe  asci  with  spores ;  e  four  ascopores  seen  laterally ;  /  germinating  ascosporea 
(x  800). 


124  CARPOASCI. 

been  obtained  in  luxuriant  cultivation  with  a  limited   supply  of 
oxygen. 

Capnodium  salicinum  (Fumago  salicina,  Cladosporium  fumarjo),  a  common 
Mildew,  forms  dark  overgrowths  on  the  leaves  and  branches  of  'various  shrubs 
(Poplars,  Elms,  Willows)  and  on  Hops.  The  conidia  appear  in  various  forms, 
as  on  conidiophores,  in  conidiocarps  with  large  multicellular  conidia,  and  in 
conidiocarps  with  small  unicellular  conidia ;  in  nutritive  solutions  yeast-like 
conidia  are  also  developed. — Apiosporium  pinophilum  produces  mildew  on  the 
leaves  of  Abies  alba  and  Picea  excelsa?  (The  conidial-forms  were  formerly 
described  as  "  Antennaria  pinophila  "). 

Order  3.  Tuberacese,  Truffles.  The  Fungi  belonging  to 
this  order  are  entirely  subterranean.  The  mycelium  is  filamen- 
ious,  and  partly  parasitic  upon  the  roots  of  plants,  especially  trees, 
in  its  neighbourhood  ;  ^it  is  then  known  as  Mycorhiza.  The  f ruit- 
»body  is  relatively  large,  in  some  cases  about  the  size  of  a  hen's 


FIG.  113.— Tuber  melanosporam :  a  fruit-body  (nat.  size),  a  portion  having  been  re- 
moved to  show  the  internal  structure  ;  b  an  ascus  with  ascospores. 

egg.  Internally  it  is  traversed  by  a  number  of  winding  passages 
(Fig.  113  a),  the  walls  of  which  are  coated  with  the  asci.  The  asci 
«(&)  contain  only  a  small  number  of  spores,  and  these  are  set  free 
by  the  putrefaction  of  the  fruit-body.  Conidia  are  unknown. 

Tuber  melanosporum,  T.  brumale,  T.  cestivum,  and  other  species  are  edible. 
Terfezia  leonis  and  Clioiromyces  mceandriformis  are  also  edible.  The  Truffles 
are  always  found  in  woods  and  under  trees,  and  disappear  when  these  are  de- 
stroyed. France  and  Italy  produce  the  best  and  the  largest  number  of  Truffles, 
which  are  hunted  by  specially  trained  dogs  and  pigs. 

In  Elaphomyces  (Stag-Truffle)  the  fruit-body  has  a  corky  external  layer,  and 
is  inedible.  Some  of  the  species  are  found  in  this  country.  E.  granulatus  is 
parasitic  on  the  roots  of  the  Fir. 


PYRENOMYCETES.  125 

Family  3.     Pyrenomycetes. 

In  this  family  the  hymenium  is  enclosed  in  small  fruit-bodies, 
perithecia  (Fig.  120  &),  which  appear  to  the  naked  eye  as  small  dots. 
In  shape  they  resemble  a  globe  or  a  flask  with  a  narrow  mouth, 
through  which  the  spores  are  ejected  (peronocarpic  ascocarps). 
Different  kinds  of  reproduction — conidia,  pycnidia  (chiefly  with 
microconidia),  chlamydospores,  and  perithecia — are  found  in  the 
same  species.  The  various  stages  in  the  life-history  of  these 
Fungi  are  so  dissimilar,  that  formally  they  were  considered  to  be 
different  genera.  Ergot  furnishes  a  very  good  example. 

This  family  may  be  subdivided  into  3  sub-families. 


PIG.  114.— 'A  email  portion  of  an  ovary  attacked  FIG.  115. — An  ovary  with  the  conidial 

with  Claviceps  purpurea  (Sphacelia).  stage  of  Claviceps  purpurea  (Sphacelia). 

Sub-Family  1.     Hypocreales. 

The  perithecia  are  pale,  fleshy,  brightly  coloured,  and  generally 
aggregated  on  a  stroma.  Conidia  and  chlamydospores  occur  very 
frequently.  Only  one  order. 

Order.  Hypocreacese.  In  this  order  the  majority  are 
parasites  upon  Flowering-plants  (Nectria,  Polystigma,  Epichloe, 
Claviceps) ;  but  some  are  parasites  upon  Fungi  (Hypomyces,  Melano- 
spora),  or  upon  insects  (Cordyceps). 

The  most  important  member  of  this  order  is  the  ERGOT  {Clavi- 
ceps purpurea,  Figs.  114,  115,  116).  This  Fungus  is  found  in  the 
flowers  of  many  species  of  Grasses,  especially  the  Rye,  attacking 
and  destroying  the  ovaries.  In  the  FIRST  or  coxfloiAL  STAGE  of  the 
attack,  the  ovaries  are  found  covered  with  a  white,  irregularly 


126 


PYRENOMYCETES. 


folded  mycelium  (Fig.  114  m,  Fig.  115),  formed  of  numerou 
hyphse  woven  together  and  penetrating  the  wall  of  the  ovary. 
From  these  a  number  of  hyphaB  (Fig.  114  a)  project  into  the  air 
and  abstrict  from  their  apices  the  conidia  (6)  which  serve  as  repro- 
ductive organs.  The  mycelium  also  secretes  a  sticky,  stinking 
fluid  (honey-dew)  in  which  the  conidia  are  embedded  in  great 
numbers.  The  honey-dew  exudes  from  the  bases  of  the  glumes, 
and  is  greedily  sought  by  flies,  which  thus  carry  the  conidia  to- 
other ovaries.  In  this  manner  fresh  ears  are  infected,  which  might 


, 


Pis.  116.— Chviceps  purpnrea.  A  Sclerotium  with  stromata  (cl)  (x  by  2).  B  Stroma- 
divided  longitudinally  to  show  the  perithecia  (cp).  C  A  perithecium  with  the  surrounding- 
hyphge(Jiy).  D  An  ascus  ruptured,  with  the  eight  filamentous  ascospores  emerging. 

escape  were  the  conidia  only  distributed  by  the  wind.  This  stage 
formerly  was  regarded  as  an  independent  Fungus,  known  as  Spha- 
ceha  segetum  (Fig.  115).  On  germination,  the  conidia  produce 
either  a  new  mycelium  (Fig.  114  d,  c),  or  new  conidia.  The  SECOND 
or  SCLEROTIUM  STAGE  is  the  one  in  which  the  Fungus  passes  the 
winter.  The  mycelium  penetrates  deeper  and  deeper  into  the 
attacked  ovaries,  their  tissues  are  destroyed  and  replaced  by  the 
hyphas,  which  gradually  become  more  and  more  felted  together. 


PYRENOMYCETES. 


127 


A.  firm,  pseado-parenchymatous  mass  of  hyphoe  is  thus  formed  at 
the  base  of  the  loosely-woven  Sphacelia,  which  is  in  part  trans- 
formed into  the  hard  sclerotium,  and  the  remainder  thrown  off. 
A  dark,  hard,  poisonous  body,  longer  than  the  natural  grain,  is 
thus  formed ;  these  bodies  are  known  as  Ergots,  and  were  formerly 
considered  to  be  a  distinct  species, — Sclerotium  clavus  ("  Secale 
cornutum,"  Ergot,  Fig.  116  A,  c).  The  THIRD  STAGE,  described 
as  Claviceps  purpurea,  is  developed  in  the  following  spring  from 
the  germinating  sclerotium,  which  produces  dark-red  stromata 
with  short  stalks.  In  the  stroma  numerous  perithecia  with  asci 
and  ascospores  are  produced.  The  latter  may  infect  young  flowers 
of  the  cereals,  in  which  the  disease  is  then  developed  as  before. 


FIG.  117.— Nectria  cinnnlarina  :  a  branch  of  Acer  pseudoplatanus,  with  conidial-layers 
and  perithecia  (nat.  size) ;  b  a  conidial -layer  (Tubercularia  vulgaris) ;  c,  a  mass  of  peri- 
thecia.  (6  and  c  x  8.) 

Several  species  of  the  genus  Nectria,  with  blood-red  perithecia,  are  found  as 
dangerous  parasites,  especially  N.  ditissima,  which  causes  "  Canker "  in  the 
Beech,  Ash,  and  Apple,  etc. ;  N.  cucurbitula,  which  appears  on  Pine-trees,  and 
N.  ciimabarina  (Fig.  117),  whose  conidial  form  was.formerly  named  Tubercularia 
vulgaris. — Folystiyma  rubrum  forms  shining  red  spots  on  the  green  leaves  of 
Prunus-species.—Epichloe  typhina  is  parasitic  on  the  sheaths  of  Grasses,  on  which 
it  first  forms  a  white  conidial-layer,  later  on  a  yellow  layer  of  perithecia. — Cor- 
dyceps  (Chrysalis  Fungus,  Figs.  118,  119)  lives  in  and  destroys  insects,  and  after 
compassing  their  death  produces  the  club-formed,  generally  yellow,  stromata, 
one  part  of  which  bears  conidia  (Isaria)  and  another  perithecia.  C.  militaris 
(Fig,  118)  on  the  chrysalides  and  caterpillars  of  moths,  is  the  most  common. 

W.B.  K 


128 


PYKENOMYCETES. 


The  so-called  Botrytls  bassiana,  which  produces  the  disease  known  by  the  name 
of  "  Muscardine,"  in  silkworms,  is  probably  a  conidial  form  belonging  to 
Cordyceps. 


Fi«.  118.—  Cordyccps  militaris.  I  Stromata  with 
conidiophores  (ft  aria  farinosa).  II  A  larva,  with 
stromata,  bearing  perithecia.  Ill  A  spore. 


Fio.  119.— Cordyceps  robertii  on  the  larva  of  Hepialus  virescens  :  a  stalk  of  stroma  ; 
perithecia. 


PYRENOMYCETES. 


129 


Sub-Family  2.    Sphaeriales. 

To  this  sub-family  belong  the  majority  of  the  Pyrenomycefces. 
The  perithecia  are  of  afirm  consistence  (tough,  leathery,  woody  or 
carbonaceous),  and  of  a  dark  colour.  Their  covering  is  quite  dis- 
tinct from  the  stroma  when  this  structure  is  present.  The  stromata 
are  sometimes  very  large,  and  may  be  either  cushion-like,  crus- 
taceous,  upright  and  club-like,  or  branched  bodies.  In  general, 
small,  inconspicuous  Fungi,  living  on  dead  vegetable  matter, 
sometimes  parasites.  Free  conidiophores  and  conidiocarps  are 
known  in  many  species,  and  in  several,  chlamydospore-like  forms 
of  reproduction.  Orders  3-18  constitute  the  Sphasriacese  of  older 
systematists. 


I 


FIG.  120.— Strtc7:eria  obduccns  :  a  a  portion  of  an  Ash-branch  with  the  bark  partly  thrown 
off;  on  the  wood  are  numerous  black  perithecia  (x  20);  b  longitudinal  section  through  a 
perithecium ;  c  a  spore :  d  longitudinal  section  through  a  pycnidium  whose  ascospores 
are  being  ejected  ;  e  portion  of  the  same,  with  hyphse  and  spores. 

Order  1.  Sordariaceae.  —  Fungi  living  on  dung  with  fragile  perithecia, 
•either  aerial  or  buried  in  the  substratum.  The  dark  brown  or  black  spores  have 
either  a  mucilaginous  envelope  (Sordaria,  etc.)  or  mucilaginous  appendages  (Podo- 
spora),  by  means  of  which  their  expulsion  and  distribution  are  promoted. 

Order  2.  Chsetomiaceae.  Perithecia  fragile,  free,  bearing  on  the  summit  a 
tuft  of  hairs.  Chcetomium,  on  decaying  vegetable  matter. 

Orders  3-7.  PcritUecia  scattered  or  aggregated,  situated  from  the  commence- 
ment on  the  surf  ace  of  the  substratum.  Stroma  wanting. 

Order  3.  Trichosphaeriacese.  Trichosph^ria  parasitica  (Fig.  121),  on 
Abies  alba  ;  Herpotrichia  nigra  on  Picea  excelsa  and  Pinus  montana. 


130 


PYRENOMYCETES. 


Order  4.  Melanommaceae.  Eosellinia  quercina  lives  in  the  roots  of  1-3- 
year-old  Oaks,  and  destroys  the  plants. 

Order  5.     Ceratostomaceae. 

Order  6.  Amphisphaeriaceae.  Strickeria  obducens  (Fig.  120)  has  brick- 
like  spores,  and  lives  aggregated  on  the  hard  branches  of  Fraxinus. 

Order  7.     Lophiostomaceae. 

Order  8.  Cucurbitariaceae.  Perithecia  tufted,  at  first  embedded,  then 
breaking  through,  often  situated  upon  an  indistinct  stroma. 

Orders  9-13.  The  perithecia  remain  embedded,  and  are  only  liberated  by  the 
casting  off  of  the  covering  layers  of  the  substratum.  Stroma  loanting. 

Order  9.  Sphaerellaceae.  The  species  of  Sphcerella  have  colourless,, 
bicellular  spores.  They  live  upon  the  leaves  of  many  plants,  and  develope- 
spherical  perithecia  upon  the  fallen  leaves. 

Order  10.  Pleosporaceas.  The  conidial-forms  of  Pleoapora  herbarnm  and 
P.  vulgaris  form  a  black  covering  on  various  plants,  known  as  "smuts." — 


a 


\l 


FIG.  121. — Trichospharia  parasitica  -.  a  a  twig  of  Abies  alba  with  epiphytic  mycelium ; 
b  a  leaf  with  mycelium  and  sporangia  (magnified);  c  a  sporangium  (x  60);  d  an  ascus 
with  spores  ( x  550). 

Venturia  ditricha  occurs  on  the  underside  of  dry  Birch  leaves,  and  perhaps  to 
this  belongs  the  conidial-form,  Fusicladium  pirinum,  which  causes  the  "  Bust- 
spots  "  on  Apples  and  Pears. 

Order  11.     Massariacese. 

Order  12.     Clypeosphaeriaceae. 

Order  13.  Gnomoniaceaa.  Perithecia,  with  peak-like  apfrture.  Gnomonia 
erythrostoma  in  the  leaves  of  Prunus  avium,  which  turn  brown  and  do  no  fall 
in  autumn. 

Orders  14-18.  Stroma  generally  well  developed,  The  perithecia  are  embedded, 
in  the  stroma,  but  ichen  this  is  rudimentary,  in  the  substratum. 

Order  14.     Valsaceae.     VaJsa. 

Order  15.     Diatrypaceae.     Diatrype.     ' 

Order  16.     Melanconidaceae. 

Order  17.     Melogrammataceae. 


PYRENOMYCETES. 


131 


Order  18.  Xylariacese.  This  order  is  the  most  highly  de- 
veloped of  the  Sphseriales.  The  stroma  arises  on  the  surface  of 
the  substratum,  which  is  generally  dead  or  decorticated  wood ; 
it  is  well-developed,  crustaceous,  hemispherical  or  upright.  Iii 
the  younger  conditions  it  is  covered  with  a  layer  of  conidia,  and 
later  on  it  bears  the  perithecia,  arranged  in  a  layer  immediately 
beneath  its  surface.  The  ascospores  are  of  a  dark  colour.  Often 
also  there  are  free  conidiophores. 


FIG.  122. — JTylaria  hypoxylon  (nat.  size)  on  a  tree  stump  :  a  younger,  b  an  older  stroma, 
both  of  which,  with  the  exception  of  the  black  lower  portion,  are  covered  with  white 
conidia  ;  n,  spot  where  the  perithecia  are  developed ;  c  an  old  stroma  with  upper  part 
fallen  off ;  d,  e  large  branched  stromata  ;  fc  conidia. 

Hypoxylon  and  Ustulina  have  a  cushion-like  or  crustaceous  stroma. — Xylaria 
has  a  club-shaped  or  branched  stroma,  often  several  centimetres  high.  X. 
hypoxylon  (Fig.  122)  and  X.  polymorpha  occur  on  old  tree  stumps. — Poronia 
grows  on  old  horse  dung,  and  has  a  conical  stroma. 

Sub-Family  3.     Dothideales. 

The  perithecia  are  always  embedded  in  a  black  stroma,  and  are 
not  distinctly  separated  from  it.  The  accessory  forms  of  reproduc- 
tion are :  conidiophores,  conidiocarps,  and  yeast-like  conidia.  The 
majority  are  parasites.  One  order. 

Order  Dothideacese.  Phyllachora  graminis  produces  scab-like  patches  on 
the  leaves  of  the  Grasses. — Scirrhia  rimosa  grows  on  the  leaf-sheathes  of 
Phragmites. — Rhopographus  pteridis  on  Pteridium  aquilinum. 


132 


D1SCOMYCETES. 


Family  4.     Hysteriales. 

This  family,  like  the  following,  has  hemiangiocarpic  ascocarps 
(apothecia).  These  are  closed  in  the  early  stages,  but  when  ripe 
open  in  a  valvular  manner  by  a  longitudinal  fissure  ;  they  are  black, 
oblong,  and  often  twisted.  Some  species  are  parasites,  especially 
upon  the  Coniferae. 

Order  1.  Hysteriaceae.  Hysterium  pulicare  upon  the  ruptured  bark  cf 
many  trees. 

Order  2.  Hypodermaceae.  The  species  of  Lophodermium  live  upon  tie 
leaves  of  Conifers,  and  are  the  cause  of  their  falling  off  (blight).  L.  pinastri 


FIG.  124.— Three  leaves  of 
the  Red-pine  with  Lopho- 
dermium macrosporuin :  a 
under  side  of  the  leaves  with 
apothecia  ;  b  a  leaf  from 
upper  side  with  pycnidia. 
(x  about  2.) 


a  b 

FIG.  125. — Lophodermium 
pinastri:  a  leaves  of  Pinus 
sylvestris  with  apothecia 
(nat.  size);  b  two parapLyses 
and  an  ascus  with  filamen- 
tous spores. 


FIG.  123. — Lophodermium 
(Hypodermd)  nerviscquium  : 
a  two  leaves  of  Abies  aZba 
seen  from  above  with  pyc- 
nidia; b  a  leaf  seen  from 
the  underside  with  apothe- 
cia; c  an  ascus  with  asco- 
spores.  (x  500.) 

(Fig.  125),  on  the  leaves  of  Pinus  and  Picea ;  the  leaves  become  red-brown 
and  fall  off ;  at  first  conidiocarps  are  formed,  and  later  apotbecia  ;  L.  nervise- 
quium  (Fig.  123),  on  Abies  alba;  L.  rnacrosporum  (Fig.  124),  on  Picea  exceha,\ 
L.  bnichysporum,  on  Pinus  strobus. 

Order  3.     Dichaenaceae. 

Order  4.     Acrospermaceae. 

Family  5.     Discomycetes. 

The   ascocarps  (apotJiecia)  are  at  first  closed,  and  only  open  at 
the    time   of    their    ripening,    not    valvularly,    but    more  or    less- 


DISCOMYCETES.  133 

like  a  saucer  or  cup,  so  that  the  hymenium  lies  exposed  on  their 
upper  surface.  In  the  first  three  sub-families,  and  generally  also 
in  the  fourth,  the  apothecia  are  formed  inside  the  substratum. 
The  apothecia  are,  in  contrast  to  the  Pyrenomycetes,  light  and 
brightly  coloured,  and  their  size  varies  very  much,  and  may  be 
several  centimetres  in  diameter.  Paraphyses  are  often  present 
between  the  asci ;  they  often  contain  colouring  matter,  and  give  to 
the  disc  its  characteristic  colour.  The  tissue  on  which  the  asci 
are  borne  is  known  as  the  kypotliecium .  The  shape  and  colour  of 
the  spores  is  not  so  varied  as  in  the  Pyrenomycetes.  The  acces- 
sory forms  of  reproduction  are  conidia  (sometimes  of  two  forms), 
chlamydospores,  and  oidia.  The  family  is  divided  into  5  sub- 
families. 

Sub-Family  1.     Phacidiales. 

The  apothecia  are  developed  in  the  interior  of  the  substratum, 
which  they  break  through,  and  in  general  dehisce  apically.  The 
envelope  is  tough  and  black.  Hypothecium  inconspicuous ; 
hymenium  flat. 

Order  1.  Euphacidiaceae.  Phacidnim  abietinum,  on  the  leaves  of  Abies 
alba. — Rhytisma;  the  pycnidia  are  found  in  the  summer  on  the  green  leaves, 
while  the  apothecia  are  developed  oil  the  fallen  leaves  and  dehisce  in  the 
following  spring.  R.  acerinum  causes  black  spots  on  the  leaves  of  the  Syca- 
more, and  R.  salicinuin  on  Willows. 

Order  .2.     Pseudophacidiaceae. 

Sub-Family  2.    Stictidales. 

The  apothecia  when  ripe  break  through  the  substratum  which 
forms  a  border  round  them.  Hymenium  generally  saucer-shaped. 

Order  1.     Stictidaceae.     Stictis. 
Order  2.     Ostropaceae.     Ostropa. 

Sub-Family  3.    Tryblidiales. 

The  apothecia  are  embedded  in  the  substratum  in  the  early 
stages,  and  then  are  raised  high  above  it.  Hypothecium  thick. 
Hymenium  cup-shaped. 

Order  1.     Tryblidiaceae.     Tryblidium. 

Order  2.  Heterosphaeriaceae.  Heterospluzria  patella  on  the  dead  stalks 
of  Umbellifers. 


134 


DCSCOMYCETES. 


Sub-Family  4.     Dermateales. 

The  apotliecia  in  the  early  stages  are  embedded  in  tlie  sub- 
stratum and  then  break  through  it,  or  are  from  the  first  situated 
on  the  surface  of  the  substratum.  Hypothecium  thick. 

Order  1.     Cenangiaceae.     Cenangium. 

Order  2.     Dermateaceae.    Dermatea. 

Order  3.     Patellariaceae.    Patellea,  Biatorella,  Patellaria. 

Order  4.     Caliciaceae.     Calicium,  Coniocybe,  etc. ,  on  the  bark  of  trees. 

Order  5.  Arthoniaceae.  Arthonia  on  the  bark  of  several  trees.  Celidium 
stictarum  on  the  apotliecia  of  Sticta  pulmonaria. 

Order  6.  Bulgariaceae.  Apothecia  gelatinous  under  moist  conditions,  and 
horny  when  dried. — Galloria  fusarimdes  ;  the  red  apothecia  break  out  in  the 
spring  on  the  dried  stalks  of  Urtica  dioica  ;  a  gelatinous  reproductive  form  of 
the  Fungus  is  found  before  the  apothecia,  which  consists  of  oidia  (formerly 
described  as  "  Dacryomyces  urticcB  "). — Bulgaria  inquinans  on  the  living  or 
fallen  trunks  of  Oaks  and  Beeches. 


Sub-Family  5.     Pezizales. 

The  apothecia  are  developed  on  the  surface  of  the  substratum  and 
are  waxy  or  fleshy ;  at  the  commencement  closed,  and  covered  with 
a  saucer-  or  cup-shaped,  seldom  flat,  hymenium.  The  hypothecium 
is  generally  well  developed.  This  sub-family  is  the  richest  in 


FIG.  126. — Botrytis  cinerea :  a,  slightly 
magnified  ;  b  more  highly  magnified ;  c 
germinating  conidium. 


FIG.  127. — Sclerotinia  fuclccliania :  a  sole- 
rotium  with  conidiophores ;  b  with  apo- 
thecia ;  c  section  through  sclerotium  and 
apothecium ;  d  ascus  with  eight  asco- 
spores.  ( x  390.) 


DISCOMYCETES. 


135 


species  of  the  Discomycetes  and  contains  forms  of  very  different 
habit.  They  grow  upon  dead  wood,  upon  the  ground,  and  upon 
dung.  A  few  are  parasites. 

Order  1.  Helotiaceae.  Apothecia  with  waxy  envelope  of 
colourless,  or  yellowish  prosenchymatous  cells. — Chlorosplenium 
lemginosum  is  found  on  decaying  wood  (particularly  Oak  and  Birch) 
to  which  it  gives  a  green  colour.  Sclerotinia  has  sclerotia  which  are  do 
veloped  upon  the  host-plant  and  from  which,  after  a  period  of  rest,  the  long 
brown-stalked  apothecia  arise.  S.  ciborioides  (S.  trifoliorum,  Fig.  128)  is 
parasitic  on  Clover ;  S.  sclerotiorum,  on  Daucus-roots,  Phaseolus,  etc.  ;  S.  bac- 
carum,  on  the  berries  of  Vaccinium  myrtillus;  "  Botrytia  cinfrea  "  is  a  common 
parasite  and  is  probably  the  conidial  form  of  S.  fuclteliana  (Fig.  127). — Helotium 
Jicrbarum  lives  on  dry  plant  stems. — Dasyscypha  vnllJtommii  (Fig.  129)  produces 
Larch -canker  on  the  bark  of  the  Larch. 


FIG.   128.  —  Sclerotinia  ciborioides:    a  sclerotium  FIG. 129.  —  Uafytcypliawillkommii: 

with  three  apothecia  slightly  magnified ;   b  ascus        o  portion  of  bark  of  Larix  decidua 
with  erjht  ascospores;  c  germinating  ascospore.  with  sessile,  cup-shaped  apothecia 

(nat.  size);  b  tvro  paraphyses  on 
.  either  side  of  an  ascns  with  eight 
ascospures. 

Order  2.     Mollisiacese.     Mollisia  cinerea,  principally  on  decaying  wood. 

Order  3.  Pezizaceae.  This  order  contains  the  largest  and 
morphologically  the  highest  forms  of  the  Discomycetes.  Apothecia 
fleshy,  and  in  the  later  conditions  generally  saucer- shaped. 

Peziza,  with  sessile  apothecia,  growing  on  the  ground  ;  P.  cochleata  is  brown, 
and  coiled  like  a  snail-shell ;  P.  coccinea  is  scarlet ;  P.  aurantia  occurs  as  an 
orange- coloured  expansion  on  the  ground. 

Order  4.      Ascobolaceae.       Apothecia  fleshy  ;    in   the  later  stages  flat  or 


136 


HELVELLALES. 


convex.  The  asci  are,  comparatively  speaking,  large,  and  often  contain  a  great 
number  of  spores  which  escape  by  the  casting  off  of  a  lid  on  the  summit  of  the 
ascus.  Generally  living  on  dung. — Ascnbolus  furfuraceus,  etc. 

Family  6.     Helvellales. 

These  Fungi  have  the  appearance  of  clubs,  bells,  or  mushrooms, 
consisting  of  an  upright  stalk  bearing  a  large  and  fleshy  head,  on 
the  exterior  surface  of  which  the  hyinenium  is  spread.  The  asco- 

carps  are  probably  gymnocarpic  from 
the  beginning,  and  on  this  account 
these  plants  are  placed  in  a  separate 
family.  The  development  of  the 
nscocarps  is  unknown.  The  Morchella 
(Morell)  grows  on  the  ground ;  some 
species  are  edible.  1  order. 

Order.  Helvellaceae.  SpnUndca  is  yellow 
and  club-shaped,  and  forms  "fairy  rings  "  in 
\voods. — Geoylossvm  (Earth-tongue)  projects 
above  the  ground  as  a  black  tongue,  or  as  a 
club-shaped  body.  Several  species  are  found 
in  meadows  and  on  heaths. — Helvetia  has  a 
stalk,  bearing  an  irregularly  folded  head,  on 
the  external  surface  of  which  is  the  bypothe- 
cium.— Morchella  (Morell,  Fig.  130),  the  stalk 
bears  on  its  summit  the  conical  or  spherical 
head,  the  external  surface  of  which  is  reticu- 
late and  bears  the  asci. — Mitiula.  Verpa. 


FIG.  ISO.— Mnrchelln  esculenta:  a 
an  entire  specimen,  about  one  half 
natural  size ;  b  longitudinal  section 
through  the  head. 


APPENDIX  TO  THE  ASCOMYCETES  : 
Family  7.     Ascolichenes  (Lichen-forming  Ascomycetes). 

The  Lichens  were  formerly  classed  among  the  Thallophyta  as  a 
group  quite  distinct  from  the  Alga3  and  Fungi.  Investigations  dur- 
ing the  last  twenty-five  years,  however,  have  conclusively  proved 
that  the  Lichens  are  Fungi  which  reproduce  in  the  same  manner  as 
the  Ascomycetes,  or,  more  rarely,  the  Basidiomycetes,  and  have 
entered  into  a  peculiar  symbiotic  relation  with  Algce,  especially 
the  Cyanophyceae  and  Protococcoideae,  with  which  they  associate, 
and  without  which  they  would  be  unable  to  exist.  The  Fungus 
forms  the  largest  portion  of  the  Lichen,  enclosing  the  Alga  with 
which  it  may  be  said  to  be  commensal.  The  Fungus  especially 
produces  reproductive  bodies  and  absorbs  the  inorganic  nourish- 
ment through  the  rhizoids,  whilst  the  Alga  supplies  it  with  the 
organic  materials.  In  consequence  of  this  the  Lichens,  in  con- 


ASCOLTCHENES. 


137 


tradistinction  to  other  Fungi,  need  light  for  the  development  of 
their  nutritive  organs,  and  are  therefore,  in  any  case  internally,  of 
a  more  or  less  greenish  colour.  The  form  and  condition  of  the 
thallus  is  unusual  among  the  Fungi,  and  they  can  grow  upon  rocks 
and  in  other  places  where  no  dead  organic  matter,  such  as  would 
be  required  by  other  Fungi,  is  obtainable. 

Two  cellular  forms  are  therefore  to  be  found  in  each  Lichen: 
1.    The  cells  which  belong  to  the  Fungus.     These  are  generally 
septate,  branched    J/ypJice  without    any  trace  of  chlorophyll.     In 


Fie.  131.— Transverse  section  through  the  thallus  of  Stictafuliginosa  (x500) :  r-r  rhizoid- 
strands  which  arise  from  the  under  side  ;  g-g  gonidial  layer  ;  m  medullary  layer;  o  upper, 
it  lower  cortex. 

the  thallus  of  the  majority  of  Lichens  there  may  be  found  a 
medullary  layer  (Fig.  131  m)  of  loosely-woven  hyphae,  between 
which  there  are  large  air  chambers ;  and  an  external  layer  (cortex) 
(Fig.  131  o,  u)  formed  of  closely-woven  hyphse  without  any 
intercellular  spaces.  In  some  Lichens  (Collemacea3)  the  hypht© 
wind  about  in  the  thallus,  being  equally  distributed  throughout, 
without  forming  any  decided  strata.  These  Lichens  moreover  be- 


138  ASCOLICHENES. 

come  gelatinous  when  exposed  to  moisture  (Fig.  132),  on  account 
of  the  swelling  of  the  walls  of  the  Algae.  The  hyphse  contain 
protoplasm  with  drops  of  oil,  but  never  starch  ;  their  walls  easily 
swell  when  exposed  "to  damp  after  having  been  dried,  and  in  some 
{e.g.  Cetraria  islandica)  they  become  gelatinous  when  cooked. 
Certain  strata  of  hyphie  become  blue  on  treatment  with  iodine 
alone,  from  which  it  is  inferred  that  the  wall  is  allied,  in  its 
chemical  nature,  to  starch. 

2.     The  enclosed  Algae,  termed  "gonidia."     Some  belonging  to 
the  Cyaiiophyceae,    Protococcoideae,    (especially  Pleurococcus)  and 


FIG.  132.— Collema  microphyllum.     Transverse  section  through  the  thallua;  a  extremity 
of  tvichogyne  with  spermatia  attached  ;  g  A'ostoo  chains;  h  hyphse. 

Chroococcaceae,  are  spherical  and  are  found  isolated,  or  in  irregular 
groups  of  cells  (Fig.  131  g}  ;  some  belonging  to  Nostoc  (Fig. 
132  </),  Lyngbyaceaa,  etc.,  are  placed  in  cell-rows.  Each  Lichen, 
as  a  rule,  has  only  one  definite  Algal-form  for  its  gonidium. 

The  gonidia  either  lie  together  in  a  certain  stratum  between 
the  cortex  and  the  medullary  layer  (Fig.  131  g},  or  are  scattered 
irregularly  throughout  the  entire  thallus  (Fig.  132).  The  thallus 
is  in  the  first  instance  termed  "  heteromerous,"  in  the  second  in- 
stance, "homoiomerous."  The  Fungal-hyphae  embrace  the  gonidia 
and  apply  themselves  closely  to,  or  even  penetrate  them,  and 
hence  it  has  been  difficult  to  decide  whether  the  one  cellular  form 
does  or  does  not  develope  from  the  other  (Figs.  134,  135). 


ASCOLICHENES. 


139 


This  theory  regarding  the  symbiosis  of  Fungi  and  Algse  to  form  a  Lichen  is 
termed  the  Schwendenerian  theory,  after  the  first  scientist  who  advanced  it  with 
any  weight.  It  had  heen  already  indicated  by  De  Bary,  and  further  arguments 
in  its  support  have  at  a  later  time  been  adduced  by  Bornet,  Stahl,  Treub, 
Frank,  Bonnier,  Alfr.  Moller  and  others. 


Fro.   131.— Yosfoc  Uchenoides,  which  is  attached 
by  a  g-erminating  thread  (M  of  Collema  glaucescens. 


FIG.  133.  —  Epliele  pubefccns. 
The  apex  of  a  branch  of  the 
thallus  with  two  lateral  branches 
(s)  :  h  its  hyphse;  g  the  apical 
goniclinm  of  the  main  branch. 


FIG.  135.— A  Germinating  spore  of  Physcia  parie- 
tina  with  Profococcus  vivid's.  B  Synalissa  sympliorea 
with  Gl wocapsa.  C  Cladonia  furcata  with  Proto- 

COt'CUS. 


The  thallus  of  the  Lichen  appears  mainly  under  three  forms : — 
1.     The   CRUSTACEOUS,  which   adheres  firmly  to  the  substratum 
(bark,  stone)  throughout  its  entire  surface,   without  being  raised 
into   any  free  patches  or  lobes.       It  has,  in  many  instances,  no- 
definite    outline,   and    hyphal-branches    from    it   often    penetrate 


140 


ASCOLICHENES. 


deeply  into  the  substratum.     It  grows  at  the  circumference  and 
sometimes  dies  away  in  the  centre  (Figs.  138,  139,  140). 

2.     The  FOLIACEOUS.     This  also   lies  flat  upon  the  substratum, 
but  is  not  firmly  attached  to  it  and  has  a  definite  outline.  It  grows 


FIG.  136.—  Portion  of  ».  hymenium  :  d  a  thin 
stratum  on  which  the  asci  (s)  are  situated. 


Fio.  137.—  Spoies  of,  a  Cladonia,  Lecanora  and  Pertusaria;  b  Bceomyces  •  c  Sphinctrina: 
d,  e,f  various  species  of  Parmelia  ;  g,h  Verrucaria  in  its  younger  and  older  condition;  t, 
fc  species  of  Leplogium. 

at  the  margin,  and  raises  itself  a  little  by  free  outgrowths  and 
lobes  (Fig.  141).  The  rhizoid-strands  spring  out  from  its  whitish 
under  surface  (Fig.  131,  r). 

3.  The  FRUTICOSE,  which  is  attached  to  its  substratum  at  a 
small  point  from  which  it  projects  freely,  either  erect  or  pen- 
dulous. It  is  more  or  less  tufted,  in  the  form  of  a  bush  (Figs.  14:.', 


FIG.  13d. — Lecanora  subfusca : 
a  the  bark  on  which  it  is  situ- 
ated ;  I  the  thallus  ;  s  the 
ascocarp;  s'  anascocarp. 


Fi«.  139.  —  Grapliis 
(two  species). 


FIG.  140.— Pei-tusaria  com- 
munis. 


143).      These  three    thallus-forms  gradually  pass  over  by  many 
intermediate  forms  into  one  another. 

The    Lichens,   like    other    Ascomycetes,    have   very    variously 


ASCOLICHENES. 


141 


constructed  ascospores  (Fig.  137),  which  are  enclosed  in  asci 
(Fig.  136),  usually  surrounded  by  paraphyses  attached  together. 
They  also  possess  spermogonia  (Fig.  141)  containing  numerous 
spermatia.  These  are  by  some  considered  as  pycnidia  and  micro- 
conidia,  Moller  having  shown  that  the  microconidia  under  certain 
conditions  are  able  to  germinate  and  produce  a  mycelium  with 
new  conidia,  just  as  in  other  Ascomycetes.  This,  however,  does 
not  disprove  the  sexual  nature  of  these  spores. 

[The  development  of  the  ascocarps  (apothecia)  from  carpogonia, 
as  in  the  Floridese,  which  was  first  shown  by  Stahl,  has  lately 
been  more  fully  established.  Collema  may  be  taken  as  a  type  of 


FIG.  140A. — Collema  crispum:  A 
Carpogonium  (c)  with  trichogyne 
(t)  (x  405);  J5  apex  of  the  tri- 
chogyne with  spermatium  (s) 
affixed  ( x  1125).  (After  E.  Baur.) 


the  origin  of  these  structures  in  the  Lichens.  The  carpogonium 
is  a  multicellular  filament  terminating  in  a  trichogyne  which 
projects  slightly  above  the  surface  of  the  thallus,  while  the  lower 
part  is  spirally  coiled  and  embedded  in  the  tissue  (Figs.  132 


142 


ASCOLICHENES. 


140A).  The  cells  of  the  carpogonium  contain  one  nucleus,  and 
communicate  with  one  another  by  means  of  pits  in  the  cell-walls. 
The  spermatia,  developed  in  the  spermogonia,  become  affixed  to 
the  tip  of  the  trichogyiie  and  empty  their  contents  into  it.  After 
this  conjugation  has  taken  place  the  trichogyne  withers,  the  coiled 
cells  now  divide  and  constitute  an  ascogoiie,  from  which  the  asco- 
genous  hyphge  arise.  The  paraphyses  proceed  from  two  or  three 
cells  at  the  base  of  the  ascogone.] 

VEGETATIVE  REPRODUCTION  takes  place  by  soredia,  which  to  the 
naked  eye  appear  as  whitish  powder  on  the  surface  of  the  thallus. 
They  are  small  round  bodies,  formed  by  one  or  a  group  of  gonidia, 
which  are  surrounded  by  a  mass  of  felted  hyphse.  After  the 
rupture  of  the  cortex  they  are  set  free,  and  readily  carried  by  the 
wind  to  other  places,  where  under  favourable  circumstances  they 
establish  a  new  thallus. 


FIG.  141. — A  A  portion  of  the  thallus  of  Parmelia  parietina  with  ascocarps  (a)  and  sper- 
mogonia (6).  B  A  portion  of  the  thallua  of  Celraria  islandica  with  spermosjonia  at  t».e 
end  of  small  lobes.  C  A  lobe  with  spermogonia  and  ejected  gpermatia.  (Magnified.) 

GEOGRAPHICAL  DISTRIBUTION.  The  Lichens  are  the  most  hardy 
plants,  and  are  the  first  to  appear  on  hitherto  bare  rocks  which 
they  gradually  disintegrate,  and  hence  prepare  the  way  for  the 
growth  of  other  plants.  They  are  to  be  found  from  the  Polar 
regions  to  the  Equator;  from  the  highest  snow-free  mountain- 
peaks  down  to  the  level  of  the  sea  ;  on  the  stems  of  trees ;  on  rocks, 
soil,  some  even  on  inundated  places ;  on  stones  in  woodland  streams, 
and  on  beaches;  but  they  are  never  found  upon  rotten  organic 
remains.  Some  grow  gregariously  in  enormous  masses,  and  form 
wide-stretching  carpets,  e.g.  Reindeer  Moss  (Cladonia  rangiferina), 
species  of  Cetraria  and  other  fruticose  Lichens. 

USES.     On  account  of  the  cell-wall  being  composed  of  Lichen- 


ASCOLICHENES.  142A 

starch  (Lichenin),  the  Iceland-Lichen  and  Manna- Lichen  (Lecanora 
esculenta)  are  used  as  food ;  the  latter  grows  on  stones,  in  the 
deserts  of  Asia  and  North  Africa,  and  is  often  torn  loose  in  large 
masses  and  carried  away  by  the  wind.  The  Reindeer- Lichen  is 
not  only  the  principal  food  of  the  reindeer,  but  it  is  also  used  in 
the  manufacture  of  Danish  brandy.  Cetraria  islandica  (Lichen 
islandicus)  is  OFFICINAL.  Colouring  materials  (lacmus,  orseille, 
persio)  are  made  from  several  species,  especially  from  Roccella 
tinctoria  (from  the  rocky  coasts  of  the  Mediterranean).  Parmelia 
saxatilis  and  particularly  Lecanora  tartarea  are  used  for  colouring 
purposes  in  the  Northern  countries. 

About  2,000  species  of  Lichens  have  been  described.  If  we 
disregard  the  Basidiolichenes,  which  will  be  considered  on  page 
176,  the  remaining  Lichens  (Ascolichenes)  may  be  divided  into 
the  two  following  orders  according  to  the  structure  of  the  fruit- 
bodies  : — 

Order  1.  Pyrenolichenes.  The  ascocarps  (apothecia)  are 
spherical  or  flask-shaped,  as  in  the  Pyrenomycetes,  more  rarely 
linear  (Graphis}. 

According  to  the  nature  of  the  thallus,  these  Lichens  may  be  divided  into  : — 

a.  Thallus  homoiomerous,  but  not  gelatinous,  branching  according  to  the 
mode  of  growth  of   the  Algae  :    Ephebe  (Fig.  133),   with  Algae   of  the  genus 
Stigonema. 

b.  Thallus  homoiomerous,  gelatinous  :  Lichina. 

c.  Thallus  heteromerous,  crustaceous :   Verrucaria,  Pyrenula;   Graphis  (Fig. 
139),  which  may  be  considered  as  Hysteriaceae  with  gonidia;  several  species  of 
Graphis  are  common  on  bark. 

d.  Thallus  heteromerous,  foliaceous  :  Endocarpon. 

e.  Thallus  heteromerous,  fruticose  :    Sphcerophorus. 

Order  2.  Discolichenes.  These,  as  in  the  Discomycetes, 
have  open  apotkecia,  which,  as  a  rule,  are  cupular,  more  rarely 
hemispherical  (Oladonia). 

According  to  the  nature  of  the  thallus,  these  Lichens  may  be  divided  into  : — 

a.  Thallus  homoiomerous,  but  not  gelatinous,  branching  according  to  the 
mode  of  growth  of  the  Algae  :  Ccenogonium. 

b.  Thallus  homoiomerous,  gelatinous :  Collema  (Fig.  132),  with  Algae  of  the 
genus  Nostoc ;  Leptogium. 

c.  Thallus  heteromerous,  crustaceous:  Pertusaria  (Fig.  140),  Lecidea,  with 
apothecia  open  from  the  beginning  ;  Lecanora,  with   apothecia,  which  in  the 
beginning  are  closed,  later  on   open,  but  with  a  rim  formed  by  the  thallus 
(Fig.  138) ;  Bceomyces,  whose  apothecia  are  borne  on  a  stem  formed  by  the 
thallus. 


ASCOLICHENES. 


143 


d.  Thallus  heteromerous,  foliaceous :   Parmelia  (P.  saxatilis ;    P.  parietina, 
Wall-Lichen,   Fig.  141,   is    yellow,  very  frequent  on   tree-stems,    stone-walls, 
tiles);  Physcia  (P.   ciliaris, 

frequent  on  tree-stems)  ; 
Sticta  (S.  pulmonacea,  Lung- 
Lichen,  on  tree-stems) ; 
Peltigera,  especially  on  the 
Moss  among  trees  ;  Umbili- 
caria,  on  rocks. 

e.  Thallus  heteromerous, 
fruticose  :    Cetraria   (G.  it- 
landica,    "Iceland     Moss," 
with    an    olive-brown,   flat, 
furrowed,    fringed    thallus, 
on  heaths  ;  C.  nivalis,  white, 
in      the      Polar      regions ; 
Evernia,     Ramalina,   Usnra 
(U.  barbata,  Beard-Lichen, 
Fig.  143)  ;  Roccella,  Stereo- 
caulon,  Cladonia,  of  which 
the  genus    C.    rangiferina, 
Reindeer- Moss  (Fig.  142)  is 

important ;  Cladonia  has  two  kinds  of  thallus,  one  scaly  and  leaf-like,  the  other 
erect,  which  bears  the  apothecia  and  may  be  fruticose  (Fig.  142),  or  cupular 
(Fig.  144) ;  they  grow  in  soil  in  forests  and  on  heaths. 


FIG  142. — Cladonia  rangiferina :  s  ascocarp 


FIG.  143.— Usnea  barbata  :  s  ascocarp.     (Slightly 
magnified.) 


FIG.  144.  —  Clndonia  pyxidata. 


\V.  R. 


144 


BASIDIOMYCETES. 


Sub-Class  2.    Basidiomycetes. 

This  sub-class  embraces  the  most  highly  developed  Fungi,  with 
large  "  fruit-bodies,"  which  in  ordinary  language  we  shortly  term 
Funguses,  Toadstools,  or  Mushrooms. 

They  have  no  sporangia,  but  reproduce  only  by  means  of 
basidiospores,  conidia,  chlamydospores  and  oidia.  The  chief 
characteristic  of  this  sub-class  is  the  basidiiim  (Fig.  145),  i.e.  the 
conidiophore,  which  has  a  distinctive  form,  and  bears  a  definite 
number  (generally  4)  of  characteristically  shaped  conidia  (basidio- 
spores, Fig  145  c,  d,  e). 


FIG-.  145. — Development  of  spores  in  Corticlum. 

The  summit  of  each  basidium  is  produced  generally  into  four 
conical  points  (sterigmata,  Fig.  145  6),  from  each  of  which  a 
basidiospore  is  abstracted.  The  basidia  may  be  classified  into 
three  principal  groups,  each  of  which  accompanies  a  distinctive 
conidiophore  :  1,  the  long,  filamentous,  transversely  divided  basidia, 
with  lateral  sterigmata  and  spores,  found  in  the  Uredinaceas  (Figs. 
146  D,  153),  Auriculariacea3  (Fig.  160  £),  and  Pilacraceae ;  2,  the 
spherical,  longitudinally  divided  basidia  of  the  TremellaceaB  (Figs. 
160  c  d-,  161  iii.  iv.) ;  and  3,  the  ovoid,  or  cylindrical,  undivided 
basidia  of  the  Autobasidiomycetes  (Figs.  145,  163,  etc.)  ;  the  two 
last-  have  apical  sterigmata  and  spores. 

The  first  two  groups  are  the  septate  basidia  (protobasi'iia),  of  the  Protobasidio- 
mycetes-,  while  the  unseptate  basidia  (autobasidia)  of  the  Autobasidiomycetes 
are  the  third  group.  On  the  formation  of  the  basidiospores,  the  nucleus 


BASTDIOMYCETES.  145 

of  the  basidium  divides  into  four  nuclei,  each  of  which  is  transferred  to  a 
«pore. 

In  addition  to  the  basidia,  simple  conldiophores  are  also  found. 
In  the  Protobasidiomycetes,  the  simple  conidia  are  very  generally 
found  as  accessory  methods  of  reproduction  in  conjunction  with 
the  basidiospores ;  but  less  frequently  in  the  Autobasidiomycetes, 
e.g.  among  the  Dacryomycetes,  Tomentellaceae,  Heterobasidion 
annosum. 

The  simple  conidiophores  vary  in  size,  and  in  the  number  and  shape  of 
the  conidia  ;  they,  however,  resemble  the  basidia,  and  are  doubtless  an  early 
stage  in  the  development  of  the  definitely  formed  basidia. 

Finally,  well-defined  clilamydospores,  formed  in  various  ways, 
appear  in  the  Basidiornycetes  as  supplementary  reproductive  bodies 
(compare  p.  90).  Among  the  Protobasidiomycetes,  chlamydospores 
are  at  present  only  found  among  the  Uredinaceae,  but  in  various 
forms;  in  the  majority  of  families  of  the  Autobasidiomycetes 
oidia  frequently  occur  (Fig.  162),  but  genuine  chlamydospores 
seldom. 

In  the  same  species  several  of  the  known  forms  of  reproduc- 
tion may  be  distinguished. 

The  mycelium  is  generally  composed  of  white,  branched  strands, 
consisting  of  numerous  felted  hyphas ;  in  some,  sclerotia  are 
found. — The  great  majority  are  saprophytes ;  some  (particularly 
all  the  Uredinacese),  are  parasites. 

DIVISIONS  OF  THE  BASIDIOMYCETES. 

Series  1.     PROTOBASIDIOMYCETES  :  partly  gymnocarpic,  partly  angiocarpic. 
„     2.    AUTOBASIDIOMYCETES. 

Family  1.     DACRYOMYCETES  :  gymnocarpic. 

,,      2.    HYMENOMYCETES:  partly  gymnocarpic,  partly  hemian- 

giocarpic. 

„      3.     PHALLOIDE.^  :  hemiangiocarpic. 
„       4.     GASTEBOMYCETES  :  angiocarpic. 
Appended.     BASIDIOLICHENES  :  Lichen-forming  basidiomycetes. 

Series  I.     Protobasidiomycetes. 

To  this  series  belong  the  lowest  of  the  Basidiomycetes.  The 
basidia  appear  in  two  principal  forms  (1  and  2  on  page  144)  and 
are  divided  into  four  cells,  either  transversely  or  longitudinally, 
each  division  forming  a  sterigma  which  abstricts  a  basidiospore. 
The  first  three  orders,  Ur^dinaceae,  Auriculariacese,  and  Tremel- 


146  BASIDIOMYCETES. 

lacese  have  gymnocarpic  fruit-bodies,  while  those  of  the 
Pilacraceae,  on  the  contrary,  are  angiocarpic. 

Order  1.  Uredinaceae  (Rusts).  All  the  Rust-Fungi  are 
parasites,  their  mycelium  living  in  the  interior  of  the  stems  and 
leaves  of  their  hosts,  causing  red,  brown,  or  black  spots — hence 
their  name — and  malformations,  sometimes  of  considerable  size. 

The  Rust-Fungi  are  gymnocarpic  and  destitute  of  a  hymenium ; 
for  these  reasons  they  are  regarded  as  the  simplest  order  of  the 
Basidiomycetes.  They  are  entirely  parasitic,  and  their  filamentous, 
branched  mycelium  ramifies  in  the  intercellular  spaces  of  its  host, 
and  often  protrudes  haustoria  into  the  cells.  The  mycelium  is 
perennial  should  it  enter  a  woody  tissue  ;  it  may  also  hibernate  in 
the  rhizomes  of  perennial  herbs  and  permeate  the  shoots  springing 
from  them,  but  in  the  majority  of  the  Rust-Fungi  the  mycelium 
has  a  very  limited  growth.  The  chief  means  of  reproduction  of 
the  Rust-Fungi  are  the  chlamydospores,  which  in  the  more  highly 
developed  species  occur  in  three  forms,  namely,  the  teleuto-, 
aecidio-,  and  uredo-spores.  The  spores,  in  the  host,  are  formed 
immediately  beneath  its  epidermis,  which  is  ruptured  on  the 
ripening  of  the  spores,  with  the  production  of  "rust,"  brown,  red, 
or  black  spots.  Those  chlamydospores  which  produce  basidia  are 
termed  teleutospores.  The  spore  on  germination  produces  a  trans- 
versely divided  basidium,  "  promycelium,"  on  which  basidio- 
spores,  "  sporidia,"  generally  four  in  number,  are  produced  on 
lateral  sterigmata.  This  basidio-fructification  is  gymnocarpic; 
the  basidia  neither  form  a  hymenium  nor  a  fruit-body  (only 
Cronartium  and  Gymnosporangium  have  a  slight  indication  of  a 
basidio-fructification). 

Many  Rust-Fungi,  in  addition  to  basidiospores,  have  small, 
unicellular  conidia,  "  spermatia,"  which  are  borne  in  conidio- 
carps,  "  spermofjnnia" 

The  TELEUTOSPORES  (Winter-spores)  maybe  either  unicellular  or 
multicellular ;  in  the  majority  of  cases  they  are  enclosed  in  a  hard 
outer  cell- wall,  the  exospore,  which  in  some  cases  is  very  strongly 
developed  ;  they  have  also  a  long  or  short  stalk,  the  remains  of 
the  spore-bearing  hypha.  Each  cell  of  the  teleutospore  has  one 
germ-pore  (a  thin  portion  of  the  wall,  for  the  protrusion  of  the 
germ-tube  ;  in  Phragmidium  and  Gymnosporangium  there  are,  how- 
ever, several  germ-pores).  The  colour  of  the  teleutospores  is 
generally  much  darker  than  that  of  the  uredospores,  and  it  is  by 
these  that  the  majority  of  the  Rust-Fungi  hibernate. 


BASIDIOMYCETES.  147 

In  Gymnosporangium,  two  kinds  of  teleutospores  are  found  (distinguished 
by  their  size  and  thickness  of  exospore).  In  many  species  of  Puccinia,  the 
form  of  the  teleutospores  varies  very  much,  so  that  in  the  same  layer  spores 
have  been  observed  with  the  characteristic  form  of  other,  allied  genera. — The 
teleutospores  of  Endophylltim  resemble  aecidiospores,  since  they  are  united 
in  chains,  whose  cells  are  easily  separated,  and  are  produced  in  the  interior  of 
a  "  peridium."  The  rnulticellular  teleutospores  of  Coleosporium  function  as 
basidia,  and  from  each  cell  immediately  produce  basidiospores. — The  teleuto- 
spores of  Coleosporium  and  Chrysomyxa,  differ  from  other  teleutospores  in  the 
absence  of  exospore  and  germ-pore. 

The  ^CIDOSPORES  (Spring -spores')  are  produced  in  chains  which 
are  generally  enclosed  in  an  envelope  of  hyphse,  the  peridium ;  the 
peridium  enclosing  the  spores  being  termed  the  cecidium.  The 
secidiospores  are  unicellular,  and  generally  of  an  orange  colour  ; 
they  are  often  separated  by  intermediate  cells  which  wither  and  so 
assist  in  the  distribution  of  the  spores.  The  exospore  is  made  up 
of  minute,  radially  arranged  rods.  Generally  germination  proceeds 
immediately,  the  eecidiospore  producing  a  germ-tube,  which  de- 
velopes  into  a  mycelium  bearing  either  uredo-  or  teleuto-spores. 

The  aecidia  of  many  Rust-Fungi  were  formerly  considered  as  distinct  genera. 
The  aecidia  of  Phragmidium,  Triphraymium,  and  Melampsora,  in  which  the 
peridium  is  wanting,  were  in  part  considered  as  Cteoma.  The  ascidia  with 
fimbriate  edge,  or  those  of  Gymnosporangium  with  longitudinal  lattice-like 
splits,  were  considered  as  "  Ecestelia  "  (Lattice-Bust) ;  large,  sac-shaped  ascidia 
on  the  Conifer®  were  known  as  Peridermium. 

The  UREDOSPOKES  (Summer- spores')  are  unicellular  and  arise 
singly,  seldom  in  chains  (Coleosporiuni).  Their  colourless,  warty 
exospore  bears,  in  the  equatorial  plane,  2-8  germ-pores.  In  the 
majority,  germination  proceeds  immediately,  and  a  mycelium  is 
produced  which  at  first  gives  rise  to  uredospores  and  afterwards 
to  teleutospores. 

The  uredospore-formations  of  Melampsorella  and  Cronartium  are  enclosed  in 
an  envelope,  and  hence  resemhle  a^cidia. — Between  the  uredospores  sterile, 
unicellular  hyphse  (paraphyses)  may  be  found. 


The  spermogonia  are  spherical  or  pear-shaped 
generally  embedded  in  the  substratum,  and  are  produced  before 
the  ascidia,  before  or  simultaneously  with  the  uredospores,  or  before 
the  teleutospores.  The  conidia,  as  far  as  observations  go,  do  not 
generally  germinate  under  ordinary  conditions. 

Among    the    Rust-Fungi    some  species    are  found   which  only 
form    basidiospores    and     teleutospores     (Puwinia    malvacearum, 


148  BASID10MYCETES. 

Chrysomyxa  abietis).  Other  species  have  in  addition  uredospores; 
others  spermogonia  and  uredospores ;  others  spermogonia  and 
secidia ;  others  spermogonia,  uredospores  and  secidia.  Those 
species  in  which  all  the  methods  of  reproduction  are  not  de- 
veloped must  not  be  considered  as  incomplete  forms. 

As  a  rule  the  mycelium,  which  is  produced  from  the  basidio- 
spores,  developes  secidia;  in  the  species,  however,  without  secidia, 
it  developes  the  uredo-form,~and  when  the  uredospores  are  also  ab- 
sent, the  teleutospore-form.  It  has  been  established  in  some  species 
of  Puccinia  and  Uromyces  that  the  formation  of  secidia  can  be  sup- 
pressed, and  it  is  not  a  necessary  part  of  the  cycle  of  develop- 
ment of  the  species. 

The  majority  of  Bust-Fungi  hibernate  in  the  teleutospore-form.  Many 
species  are  able  to  hibernate  in  the  uredospore-form  (Coleosporium  senecionis). 
Others  pass  the  winter  in  the  aecidio-form,  and  develope  aecidia  on  new  hosts 
(Uromyces  pisi,  on  Euphorbia  cyparissia* ;  Phragmidium  subcorticium,  on  Rosa; 
Meidium  elatinum,  on  Abies  alba).  In  Chrysomyxa  abietis,  the  mycelium,, 
developed  from  the  basidiospores,  survives  the  winter. 

Among  the  Rust-Fungi,  with  several  forms  of  reproduction,, 
there  are  about  sixty  whose  development  can  only  be  completed  by 
an  alternation  of  hosts,  that  is,  on  one  host  only  uredo-  and  teleuto- 
spores  are  produced,  while  the  further  development  of  the  ger- 
minating basidiospores,  and  the  formation  of  the  secidia  and 
spermogonia  from  its  mycelium,  can  only  take  place  on  a  second 
quite  distinct  and  definite  host  (hetero&cious  or  metoxenous  Fungi). 
Those  Fungi  which  have  all  their  forms  of  reproduction  on  the 
same  host  are  termed  autc&cious  or  autoxenous.  It  is  not,  however,, 
always  necessary  that  the  hetercecious  Rust-Fungi  should  regularly 
change  their  hosts ;  for  example,  Puccinia  graminis  can  hibernate 
in  the  uredo-form  on  the  wild  Grasses,  and  in  the  spring  can  dis- 
tribute itself  again  in  the  same  form. 

As  a  consequence  of  the  alternation  of  hosts  the  various  forms  of  develop- 
ment were  considered  as  independent  genera  (Uredo,  sEcidium,  Rastelia,  Cccoma,. 
Periderniium),  until  De  Bary  and  Oersted  established,  about  the  same  time 
(1865),  the  mutual  connection  of  some  forms,  and  paved  the  way  for  the  right 
conception  of  these  Fungi. 

As  an  example  of  one  of  the  most  highly  developed  species, 
Puccinia  graminis,  the  "  Rust  of  Wheat,"  holds  a  prominent  posi- 
tion. Its  uredospores  and  teleutospores  are  produced  (Fig.  146) 
on  Grasses  (on  cereals,  especially  Wheat,  Rye,  Oats,  and  many 
wild  Grasses),  while  the  secidia  and  spermogonia  are  confined  to- 


BAS1DIOMTCETES. 


149 


the  Berberidaceae.  The  teleutospores,  developed  on  the  Grasses, 
hibernate  on  the  dried  portions  of  their  host,  and  in  the  succeeding 
year  each  of  the  two  cells  of  the  teleutospore  may  develope  a 
basidium  with  four  basidiospores  (Fig.  146  D,  c).  The  basidiospores 
are  distributed  by  the 
wind,  germinate  quickly, 
and  only  proceed  to  fur- 
ther development  on  Ber- 
beris  or  Mahonia.  The 
germ-tube  bores  through 
the  epidermis  of  the  Bar- 
berry-leaf, and  forms  a 
mycelium  in  its  interior, 
its  presence  being  indi- 
cated by  reddish-yellow 
spots  on  the  leaf.  After 
6-10  days  the  flask-shaped 
apermogonia  appear  (Fig. 
147  B ;  C,  a;  conidia  in 
Fig.  147  D)  and  a  few 
days  later  the  cup-shaped 
cecidia  (Fig.  147  A  ;  C,  c, 
d,  e).  The  former  are 
generally  on  the  upper, 
and  the  latter  on  the  ' 
under  side  of  the  leaf.  The 
orange  -  coloured  aecidio- 
spores  scatter  like  dust, 
and  germinate  only  on 
Grasses  ;  the  germination 
takes  place  in  about  two 
days  when  placed  on  any 
green  part  of  a  Grass. 
The  germ-tube  enters  the 
Grass  -  leaf  through  a 
stoma ;  a  mycelium  is  de- 
veloped in  the  leaf,  giving 
rise  to  a  small,  oval,  rust-coloured  spot  (Fig.  146  A)  ;  in  about  6-9 
days  the  epidermis  is  ruptured  over  the  red  spot,  and  numerous 
reddish-yellow  uredospores,  formed  on  the  mycelium,  are  set  free. 
The  uredospores  (Fig.  146  B)  are  scattered  by  the  wind,  and  can 


FIG.  146.— Puccinia 


graminis. 


150 


BASIDIOMTCETES. 


germinate  should  they  fallen  the  green  portions  of  other  Grasses: 
they  then  emit  2-4  germ-tubes  through  the  equatorial] y-placed 
germ-pores.  The  germ-tubes  enter  a  leaf  through  a  stoma,  a 
new  mycelium  is  then  developed,  and  in  about  eight  days  a  fresh 
production  of  uredospores  takes  place,  which  germinate  as  before. 
The  uredospore-mycelium  very  soon  produces,  in  addition,  the 
brown  teleutospores,  which  give  a  brown  colour  to  the  rust-coloured 
spots,  the  familiar  uredospores  on  the  cereals  being  quite  sup- 
pressed towards  the  close  of  the  summer '(Fig.  146  (7,  D).  The 
"Rust  of  Wheat"  hibernates  on  some  wild  Grasses  in  the  uredo- 
spore-form. 


FIG.  147.— JEcidium  berleridis.  A  Portion  of  lower  surface  of  leaf  of  Barberry,  with 
cluster-cups  (secidia).  B  A  small  portion  of  leaf,  with  spermogonia,  from  above.  C  Trans- 
verse section  of  leaf  on  the  upper  side,  in  the  palisade  parenchyma  are  three  ppermogonia 
(a  b);  on  the  lower  side  an  unripe  aecidium  (c  d)  and  two  ripe  secidia  (d,  e,  /);  /  chain  of 
secidiospores.  D  Hypbae,  forming  conidia. 

GENERA.  Puccinia  (Fig.  146,  147)  has  bicellular  teleutospores,  each  having 
a  germ-pore,  and  the  apcidia  when  present  have  an  indented  peridium  ;  some 
species,  as  exceptions,  have  1-3-celled  teleutospores.  Many  species  are 
HETEKO3Cious,  for  example,  P.  graminis,  described  above  ;  P.  rubigo,  which  also 
infests  various  Grasses,  but  whose  ascidia  appear  on  Ancftusa;  the  masses  of 


BASIDIOMYCBTES. 


151 


teleutospores  are  small ;  they  contain  paraphyses,  and  are  for  a  long  time 
covered  by  the  epidermis.  P.  coronata,  on  Oats  and  Eye  Grass;  its  secidia  on 
Rhamnus  ;  the  teleutospores  are  surmounted  by  a  crown — "  coronate  processes." 
P.  phragmitis,  on  Reeds  ;  aecidia  on  species  of  Rumex  and  Rheum.  P.  molinice, 
on  Molinia  ccerulea  ;  the  aecidia  on 
Orchids.  P.  poanim,  on  Meadow- 
<3rass  ;  aecidia  on  Tussilago.  Various 
Puccinias  growing  on  species  of  Cares 
have  their  ascidia  on  Urtica,  Lysi- 
machia,  Cirsinm,  Pedicularis,  etc. — 
Of  those  AUTOZCIOUS  species,  which 
have  all  their  generations  on  the 
same  host,  may  be  noted  : — P.  galii, 
P.  menthce,  P.  violce,  P.  epilobii,  P. 
•axparagi,  which  grow  on  the  hosts 
from  which  they  have  taken  their 
spe-ific  names. — As  representative  ' 

of  a  group  which  have  spermogonia,  FIG.  148.—  Gymnospo rang ium  sabiua.  A 
uredo-andteleuto-sporesonthesame  small  portion  of  the  epidermis  of  a  Pear- 
host,  but  on  different  individuals,  P.  leaf  (fl)  pierced  afc  b  b^  the  germinating 

basidiospore  (ci. 
waveolen*,    on    the     Field-Thistle, 

may  be  mentioned.  The  spermogonia  have  a  strong  odour.— A  peculiar  group 
(Leptopuccinia)  has  only  teleutospores,  which  germinate  immediately,  and 
whilst  still  attached  to  their  living  host.  To  this  group  belong  P.  arenarice, 
on  a  number  of  Caryophyllaceae  ;  and  P.  malvacearum,  on  various  Malvaceae, 
introduced  in  1873  from  South  America  to  Europe,  where  it  soon  proved  very 
destructive  to  Hollyhocks. 

Uromyces  (Fig.  149)  differs  only  from  Puccinia  in  always  having  unicellular 
teleutospores.  Among  this  genus  both  heteroecious  and  autcecious  species  are 
found.  To  the  first  group  belong  U.  pisi,  whose  ascidia  are  found  on  Euphorbia 
cyparissias,  and  U.  dactylidis,  whose  ascidia  appear  on  Ranunculus;  to  the 
second  group  belong  U.  beta,  U.  pliaseoli,  U.  trifolii. 

Triphragtnium  has  teleutospores  with  three  cells  (one  below  and  two  above), 
on  Spiraea  ulinaria. 

Phragniidium  (Fig.  150)  has  teleutospores  consisting  of  a  row  of  cells  (3-10) 
arranged  in  a  straight  line  ;  the  upper  cell  has  one 
germ-pore  and  the  others  four  germ-pores  placed 
equatorially.  Both  this  and  the  preceding  genus 
have  large,  irregular  ascidia  without  peridia,  but  often 
with  bent,  club-like  paraphyses  (150  b  and  c)  ;  they 
-are  all  autcecious.  and  are  only  found  on  the  Eosacese. 

Endophyllinn   (see  above,  under  teleutospores,   p. 
147)  on  species  of  Sempervivum. 

Gymnosporangium  (Figs.  152,  154)  has  bicellular 
teleuto&pores  collected  in  large,  gelatinous  masses 
formed  by  the  swelling  of  the  long  spore-stalks  ;  in  each  cell  2-4  germ-pores  are 
found.  Uredospores  are  wanting.  All  the  species  are  heteroecious  ;  the  teleuto- 
•spores  appear  on  Juniperus,  the  aacidia  (Roestelia)  on  the  Pomaceae.  G.  sabin<z, 


FIG.  149.  —  Uromyces 
genistce ;  a  uredospore  ;  ?> 
teleutospore. 


152 


BAS1D10MYCETES. 


on  Juniperus  tabina,  J.  virginiana,  etc.,  has  the  secidia  ("  Rastelia  cancellata  ") 
on  Pyrus  comnmnis  (Figs.  152,  148)  ;  G.  juniperinum,  on  Juniperus  communis 
with  "  Roestelia  cornuta"  (Fig.  154  a)  on  tiorbui  aucvparia,  Aria  nivea  (S.aria) 
and  Mains  communis ;  G,  clavariceforme  on  Juniperus  comnmnis,  the  aacidium 
belonging  to  it  ("  Ewstelia  lacerata  ")  on  Crattfgusoxyacantha. 

Melampsora  has  prismatic  teleutospores  placed  parallel  to  each  other  and 
forming  a  crustaceous  layer ;  in  many  species  they  are  divided  longitudinally 
into  several  cells  (Fig.  151).  The  a3cidia,  without  peridium,  belonged  to  the  old 
genus  Cfsoma.  M.  caprearum,  on  Willows,  has  the  aseidia  (Cceoma  euonynd)  on 
Euonijinus.,  M.  Imrtitjii,  on  Osiers;  the  secidium  on  Eibes.  M.  mixta,  on  Salix 


FIG.  150.— Phragmidivm  gracile :  a  an  uredospore  ;  b  and  c  two  pnrapliyses;  da  TOUII^ 
teleutospore ;  e  a  teleutospore  with  a  basidium  and  two  basidiospores  (s) ;  /  two  series  of 
ascidiospores  (Ph.  rosce). 

rep  ens  and  Orchids.  M.  pinitorqua,  on  leaves  of  the  Aspen,  ascidia  on  Pine 
branches  (Pine  shoot  fungus) ;  J/.  populina  on  Popidus  moniUfera  and  ni<ir<i  ; 
M.  betulina  (Fig.  153),  on  Birch  leaves ;  M.  padi  (Fi<;.  151),  on  leaves  of  Prnnns 
padus,  developes  teleutospores  in  the  epidermal  cells  ;  M.  lini  is  the  cause  of 
injuiy  to  the  Flax  ;  M.  agrimonies. 

Cahjptospora  goeppertiana  ;  teleutospores  on  Vaccinium  vitis  idcca\  spermo- 
gonia  and  a?cidia  on  Abies  alba  (Firneedle-Bust). 

Coleosporium  (Fig.   155)  forms  its  uredospores  in  reddish  yellow  chains ;  for 


HASIDIOMYCETES. 


153 


the  teleutospores,  see  page  147.  C.  senecionis,  on  the  Groundsel ;  its  aecidium 
(Peridermium  woljfii)  on  Pine-leaves  (Fig.  155  a).  Other  species  on  Sonchus, 
Petasites,  Campanula,  Rhinanthacca. 

Clirysomyxa  (Fig.  156)  has  bright  red,  branched  teleutospore-chains  ;  each 
spore  developes  a  4-celled  basidium.  C.  ledi,  on  Ledum  palustre;'  its  secidia 
on  the  leaves  of  the  Fir.  C.  alietis  (Fig.  156),  without  uredo-  and  secidio- 
spores;  teleutospores  on  the  leaves  of  the  Fir.  In  the  first  summer,  yellow 


FIG.  151. — Melampsora  padi:  a  and.fr 
uredospores ;  c-f  teleutospores,  seen 
from  different  sides. 


Fi:j.  152. — Pear-leaf,  seen  from  the  under 
side,  with  "ttiesteliacancellata":  indifferent 
ages  (a  youngest,  d  oldest). 


.  FIG.  153.— ATel a  in  psora  uctulina:  a, 
uredospores  ;  b  three  contiguous  teleu- 
tospores, one  of  which  has  developed 
a  bnsidium  with  three  basidiospores. 
(x  400.) 


hands  are  formed  on  the  leaves,  and  in  the  following  spring  the  red  cushions  of 
spores. 

Cronartium  (Figs.  157,  159)  has  unicellular  teleutospores  united  in  numbers 
to  form  erect  threads  or  columns ;  the  uredospores  are  enclosed  in  a  "peridium  "  ; 
C.  ribicola  (Fig.  157),  on  leaves  of  Kibes  (especially  Black  Currants) ;  its  secidia 
(Peridermium  strobi,  or  P.  klebahni)  on  the  stems  and  branches  of  Finns  strobus 


154 


BASIDJOMYCETES. 


Fict.  154. — Gymnosporanginum  juniperinum  :  a  a  smal  leaf  with  three  clusters  of  secidia 
(Hat.  size);  b  three  conidia;  c  two  aecidiospores  on  one  of  which  are  seen  the  germ -pores; 
d  a  portion  of  the  wall  of  an  secidium ;  e,  /two  teleutospores. 


FIG.  155. — Coleosporium  senecionis  ;  a  Pine-leaves  with  secidia  (Peridermium  wolffii)  r<nt. 
size;  b  an  aecidiospore ;  c  a  germinating  eecidiospore  ;  d  a  chain  of  uredospores;  e  a  chain 
of  teleutospores  of  winch  the  terminal  one  has  germinated  and  produced  a  basidiospore  (s). 


BASIDIOMYCETES. 


155 


(Fig.  159),  on  which  it  causes  great  damage  ;  C.  asclepiadeum,  on  Vincetoxicum 
officinale ;  its  aecidia  (Peridennium  cornui)  on  the  stems  and  branches  of  Pinus 
silvestris. 


FIG.  156. — Clirysoinyxa  dbietis : 
a  leaf  of  the  Fir,  with  5  clusters 
of  basidiospores(  x  4);  b  branched 
rows  of  teleutospores  springing 
from  the  mycelium  (m). 


FIG.  157. — Cronartium  ribicola :  a  mass  of  uredo- 
spores  (  x  50) ;  b  an  uredospore  ;  c  a  column  of  teleu- 
tospores (x  60);  da  small  portion  of  the  same 
more  highly  magnified,  with  a  basidium  and  two 
basidiospores  (s). 


To  the  Fungi  of  which  the  seeidium  is  known,  whilst  the  remaining  forms  are 
still  undetermined,  but  which  are  without  doubt  heteroecious,  belong  JEcidium 
elatinum,  which  produces  the  enormous  "  witches'  brooms  "  and  barrel-shaped 
swellings   on  steins  and  branches  of  Abies    alba ;  and  JEcidium  strobilinum 
(Fig.  158),  which  attacks  Fir-cones, 
causing  all  the  scales  to  become 
covered    with    clusters   of    aecidia 
opening  by  a  lid.     Hemileia  vasta- 
trix   destroyed  the   coffee   planta- 
tions in  Asia. 


Order2.  Auriculariaceae. 

The  long,  transversely  divided 
basidia  bear  laterally  4  long 
sterigmata  with  basidiospores 
(Fig.  160  B}  and  are  united 
to  form  an  hymenium  on  the 
surface  of  the  fruit -body. 
Parasites  or  saprophytes. 


FIG.  158. — JEcidium  strolilinum  :  a  scale  of 
cone  of  Picea  excelsa,  with  numerous  secidia  ;  6. 
secidiospores  arranged  in;  a  series;  c  a  cell 
of  the  peridium. 


156 


BASID10MYCETKS. 


Auricularia  sambucina  (Auricula  judce),  Judas'-ear,  has  large  fruit-bodies, 
which  may  attain  the  size  of  several  inches,  resembling  an  ear  or  a  mussel  shell. 
In  the  moist  condition  they  are  flesh-coloured,  tough  and  gelatinous,  but  when 
dried,  become  hard,  grey  and  wrinkled  ;  the  exterior  is  covered  with  short  hairs ; 
while  the  internal  surface  bears  the  hymenium.  Habitat :  stems  and  branches 
of  old  Elder- trees  (Sambucus). 

Order  3.  Tremellaceae.  The  round, pear-shaped,  longitudinally 
divided  basidia  bear  4  elongated  sterigmata,  situated  apically,  and  4 
basidiospores  (Fig.  160  C,  D),  and  are  united 
into  the  hymenium  on  the  surface  of  the 
fruit-body.  The  fruit-bodies  are  frequently 
gelatinous  and  quivering ;  similar  fruit- 
bodies  are  also  found  in  the  Dacryomyce- 
taceas  and  HydnaceaB.  Simple  conidiophores, 
which  appear  not  unfrequently  in  the  basi- 
diocarps,  before  the  basidia,  are  known  in 
many  species.  Saprophytes. 

Exidia  has  kidney-shaped,  oblong  basidiospores, 
and  small,  hook-like  conidia ;  E.  glandulosa,  E.  albida, 
etc.,  on  wood. — Craterocolla  has  conidiocarps  ;  C.  cerasi 
on  Cherry-wood. — Sclacina  incrustans ;  the  yellow, 


&  D 


FIG.  159.— Peridermium 
«trobi  :  secidia  of  Cronar- 
tiuin  ribicola  (nat.  size). 


FIG.  160. — B  Auricularia  samlucina  :  a-d  basidia  in  vari- 
ous stages  of  development;  e  a  sterigma  bearing  a  spore.— 
C  Trcmella  hitescens  :  a-d  basidia  seen  from  various  sides 
(b  from  above)  and  in  various  stages  of  development; 
e  sterigma  with  basidiospore  (x  400).  D  Exidia  glandu- 
losa :  a-c  various  stages  in  the  development  of  a  basi- 
dium;  d  sterigma  with  basidiospore  (x  350). 


fleshy,  or  cartilaginous  fruit-bodies  are  found  in  autumn  covering  the  ground  in 
moist  woods. — Tremclla  has  round  basidiospores  ;  T.  mesenterica  has  irregularly- 


BASID10MTCETES. 


157 


folded,  quivering,  orange  fruit-bodies,  about  one  inch  in  breadth  ;  T.  lutescens 
(Fig.  161)  has  orange-yellow  conidial- and  yellow  basidial-layers ;  T.  frondosa 
has  fruit-bodies  upwards  of  a  foot  in  breadth. 

Order  4.  Pilacracese.  The  transversely  divided  basidia  have  no 
sterigmata,  but  sessile  basidiospores,  and  fill  up  the  cavity  of  a  closed 
(angiocarpic}  fruit-body  as  a  gleba  without  a  regular  arrangement 
(hymenium  wanting). 

Pilacre  fagl  on  the  old  stems  of  the  Copper-Beech  ;  P.  petersii,  on  dried 
branches  of  the  Hornbeam,  has  stalked,  capitate  fruit-bodies. 


FIG.  161.— TremWxi  lutescens:  I  and  II  fruit-bodies  (nat.  size);  III  vertical  section 
through  a  fruit-body  ;  b  basidia ;  c  conidia;  IV-VI  basidia;  VII  basidiospore  with  a 
second  spore  ;  VIII  a  basidiospore  with  yeast-like  budding  (cultivated) ;  IX  a  conidio- 
phore.  ( LII-IX  about  400. ) 


Series  2.    Autobasidiomycetes. 

This  second  and  larger  part  of  the  Basidiomycetes  is  character- 
ised by  its  more  highly  differentiated,  undivided,  club-shaped,  or 
cylindrical  basidia,  which  generally  bear  4  (seldom  2,  6,  8)  apically 
placed  sterigmata  and  basidiospores  (Fig.  145).  The  fruit-bodies 
are  partly  gymnocarpic  (in  the  first  3  orders  and  in  some  Agarica- 
ceoe).  partly  hemiangiocarpic  (in  orders  3-6  of  the  Hymenomycetes 


158 


BASIDIOMYCETES. 


and  in  the  Phalloidese,  the  fruit-bodies  in  these  orders  are  in  the 
young  conditions  more  or  less  angiocarpic,  but  later  on  generally 
open  below  and  bear  the  hymeniura  on  the  under  surface  of  the 
fruit-body),  partly  also  angiocarpic  (in  the  Gasteromycetes). 


FIG.  162. — Dao yomyces  deliquescens :  I  fruit-body  (nat.  size);  II  vertical  section  through 
the  hymenium;  III  germinating  basidiospore  ;  IV  a  portion  of  mycelium  with  conidia; 
V  a  germinating  conidium  ;  VI  and  VII  chains  of  oidia  more  or  less  strongly  magnified; 
VIII  basidiospoie  of  D.  longisporus  ;  IX  germinating  basidiospore  of  D.  ovisporus;  X 
and  XI  Calocera  viscosa;  X  fruit-body  (nat.  size);  XI  basidia  with  basidiospores  (highly 
magnified);  XII  Dacryomitra  glossoides  (nat.  size). 


BASID10MYCETES.  159 

Dacryomycetes. 

The  long,  club-shaped  basidia  bear  two  tapering  sterigmata,  which 
develope  remarkably  large  basidiospores  (Fig.  162  II,  XI)  and 
form  gymnocarpic  fruit- bodies  with  hymenium.  1  order  : 

Orpte?  1.  Dacryomycetacese.  This  order  comprises  4  genera 
of  Which  the  first  two  develope  the  hymenium  on  the  whole  surface 
of  the  fruit-body,  but  the  two  last  only  on  its  apex. 

Dacryomyces :  the  folded,  gelatinous,  Tremella-like  fruit-bodies  break  out  in 
winter  on  dried  wood  (hedges)  in  the  form  of  red  or  yellow  drops.  D.  deliques- 
cens  is  very  common  (Fig.  121).  The  following  genera  have  cartilaginous 
fruit-bodies. — Calocera  (Fig.  162),  with  club-like,  simple,  or  branched,  Clavaria- 
like,  fruit-bodies  ;  the  orange  coloured  fruit-bodies  of  C.  viscosa  grow  aggregated 
together  on  the  wood  of  Conifers. — Guepinia  resembles  a  Peziza,  and  has  the 
Lymenium  only  on  the  hollow  upper  surface. — Dacryomitra  resembles  a  Mit- 
rula  (Fig.  162). 

Family  2.    Hymenomycetes. 

This  family  is  very  rich  in  species  (more  than  8000  have  been 
described),  and  to  it  belong  all  the  "  Mushrooms  "  and  "  Toadstools." 
The  fruit-bodies  present  very  various  forms ;  they  are  generally 
fleshy,  very  perishable,  seldom  leathery  or  corky,  in  the  last  case 
often  perennial.  The  basidia  are  more  or  less  cylindrical  and  bear 
generally  4  (seldom  2,  6  or  8)  sterigmata  and  basidiospores.  The 
hymenium  in  the  fully-formed  fruit-bodies  lies  free  on  the  sur- 
face :  in  orders  1  and  2  and  a  portion  of  order  6  it  is  from  the 
commencement  exposed,  fruit-bodies  gymnocarpic  ;  orders  3-6  have 
hemiangiocarpic  fruit-bodies  (p.  157).  In  the  first  order  the 
basidia  (or  the  hymenium)  are  developed  immediately  from  the 
mycelium  (Fig.  163) ;  the  fruit-bodies  of  orders  2  and  3  present  a 
higher  grade  of  development,  and  have  between  the  mycelium  and 
hymenium  a  special  hyphal-tissue,  a  stroma,  which  is  crustaceous, 
club-like,  or  coralloid,  etc.,  and  in  general  bears  the  hymenium  on 
the  largest  part  of  the  free,  smooth  surface.  In  the  forms  most 
highly  developed  (orders  4-6)  a  new  tissue — the  JiymenopJiore — is 
introduced  between  the  stroma  and  hymenium,  which  appears  on 
the  under  side  of  the  fruit-body  in  the  form  of  warts,  projections, 
tubes,  folds  or  lamellae  (Figs.  166,  167,  174  be).  Paraphyses  are 
frequently  found  in  the  hymenium,  among  the  basidia.  In  the 
Hymenomycetes  few  examples  of  conidia  can  be  recognised  at 
first.  More  frequently  clilamydospores  are  found,  particularly  oidia. 
The  mycelium  is  richly  branched,  generally  colourless,  often  peren- 
nial ;  it  lives  in  humus  or  decaying  wood,  and  is  seldom  parasitic. 

W.  B.  Af 


160 


BASEDIOMYCETES 


The  hyphse  generally  have  clamp-connections  and  unite,  some- 
times, to  form  a  rhizomorpha  (Fig.  177)  or  sclerotia  with  coloured, 
pseudoparenchymatous  covering. 


Fio.  163. — Exolasidium  vaccinii.  I  Hypertrophied  stem  of  Vaccinium  mils  idcea  ;  II  leaf 
with  gall-like  swelling  ;  III  section  of  II ;  IV  transverse  section :  m  mycelium  between 
the  parencbymatous  cells  ;  p  hypodermal  cells ;  e  epidermis  with  basidia  in  various  stages 
of  development ;  V  epidermis  with  germinating  spores  ;  VI  and  VII  spores  germinating 
in  water  (IV-VII  X  620). 


BASIDIOMYCETES. 


161 


Order  1.  Tomentellaceae.  To  this  order  belong  the  simplest  of 
the  Hymenomycetes.  The  basidia  (Fig.  145)  arise  free  and  irregu- 
larly from  the  mycelium  ;  a  hymenium  is  entirely  absent  or  very 
slightly  formed  '(in  Corticium  it  attains  its  highest  development)  ; 
fruit-bodies  are  also  wanting. — In  general  they  form  flaky,  mem- 
branous or  leathery  coverings  on  bark  and  wood.  Some  are 
parasites. 

Hypochnus  without  conidia. — Tomentella  with  conidiophores ;  growing  on 
wood  or  earth. — Exobasidium  vaccinii  (Fig.  163),  a  parasite  on  Vaccinium, 
Andromeda,  Arctostaphylos,  and  Rhododendron,  forms  flaky-powdery,  white  or 
red  coverings  and  may  cause  hypertrophy  of  the  parts  attacked.  E.  warininyii 
is  parasitic  on  Saxifraga ;  E.  lauri  causes  outgrowths  on  the  stem  of  Laurus 
•canariensis  as  long  as  a  finger,  which  formerly  were  regarded  as  aerial  roots. — 
Corticium  forms  membranous  to  leathery  layers  or  crusts;  C.  quercinum  on 
•wood  and  bark,  particularly  Oak,  is  flesh-coloured ;  C.  cceru.leu.rn  has  a  blue 
hymenium  ;  C.  giganteum  on  the  bark  of  fallen  Pine-trees. 

Order  2.  Clavariacese.  The  hymenium  is  situated  on  a  stroma, 
and  either  completely  covers  the  smooth  surface  of  the  more  or  less 
fleshy  gymnocarpic  fruit-body, 
or  is  confined  to  a  tolerably 
well  defined  upper  portion 
of  it  (Typhula}.  Paraphyses 
absent.  The  vertical,  white, 
yellow,  or  red  fruit-bodies 
are  roundish  or  club -like, 
undivided  or  richly  branched 
(Fig.  125).  Generally  on  the 
ground  in  woods,  seldom  on 
tree-stems,  etc. 

GENERA  :  Clavaria,  generally 
large  Fungi  with  thick,  round 
branches.  C.  botrytis  has  a  very 
thick,  tubercular  stem  with  numer- 
ous short,  flesh-coloured  branches: 
it  has  an  agreeable  taste.  C.  coral- 

loides  has  a  brittle,  richly-branched  fruit-body  (Fig.  164)  ;  basidia  with  two 
large  spores.  C.  pistlllaris  consists  of  a  single,  undivided  club  of  a  yellowish- 
\vhite  colour. — Sparassis  has  compressed,  leaf-like,  curled  branches  ;  S.  crispa 
has  fruit-bodies  as  large  as  a  white  cabbage-head,  with  an  agreeable  taste. — 
Typhula  and  Pistillaria  are  small  Fungi  with  filamentous  stalks,  terminating 
in  a  small  club.  The  fruit-bodies  of  the  former  often  arise  from  a  small, 
spheroid  sclerotium  ;  the  latter  is  distinguished  by  the  basidia  bearing  only 
two  spores. 


FIG.  164. — Clavaria  coralloides  (nat.  size  . 


162  BASIDIOMYCETES. 

Order  3.  Thelephoracese.  The  hymenium  is  placed  on  a 
stroina  and  covers  the  smooth  surface  of  the  leathery  hemiangio- 
carpic  fruit-body,  generally  on  its  under  side.  The  edge  of  the 
stroma,  which  bounds  the  hymenium  is  sometimes  especially 
developed  (Sterewn).  Saprophytes. 

GENEKA:  Thelephora.  The  fruit-bodies  in  this  genus  are  brown,  very  ir- 
regularly shaped,  and  often  lobed.  The  spores  too  are  brown,  but  in  the  other 
genera  colourless.  The  species  are  found  growing  on  barren  soil.  T. 
laciniata  (Fig.  165)  has  imbricate,  semi-circular,  dark-brown  pileus,  which  i& 
jagged  at  the  edge  and  upper  surface.  The  fruit-bodies  are  very  often  raised 
above  the  ground,  and  although  this  species  is  not  a  parasite,  yet  it  destroys 
young  seedlings  by  growing  above  and  smothering  them.  —  Stereum  has  a 
stiffer  fruit-body,  with  a  distinct,  fibrous,  intermediate  layer.  It  grows  on 
bark  and  wood,  projecting  like  a  series  of  imbricate  brackets.  S.  hirsutum  is 
yellow  ;  its  free  edge  is  provided  with  a  number  of  stiff  hairs,  the  upper  surface 
being  divided  into  a  number  of  zones.  S.  purpnreum  has  a  red-  violet  hymenium 

which  distinguishes  it  from  the 
previous  species.  —  Cypliella  has  a 
membranous  cup,  or  bell-shaped 
fruit-body,  often  borne  on  a  stalk, 
the  concave  surface  being  covered 
with  the  hymenium.  They  are 
small,  white  Fungi,  growing  on 
Moss  and  dead  stems.  —  Solenia  is 
closely  related  to  Cyphella  ;  its 
fruit-bodies  are  smaller  and  hairy  ; 
they  are  found  clustered  together 
forming  a  crust-like  covering  on 
dead  ^oA.  -  Craterellus  has  a 


Pie.165.-Tfc.Up*oml«fl»ni«ta(nat.Bize). 

large,     funnel-shaped     fruit-body, 

the  hymenium  covering  the  external  surface.  C.  cornucopioidcs  is  shaped  like 
a  trumpet  or  a  "  horn  of  plenty."  It  is  dark-grey,  several  inches  in  height,  and 
grows  gregariously  on  the  ground  in  forests.  It  is  distinguished  by  the  basidia 
bearing  only  two  sterigmata. 

Order  4.  Hydnaceae.  The  fruit-body  is  most  frequently 
fleshy,  and  varies  considerably  in  shape,  the  simplest  forms  being 
resupinate,1  the  higher  ones  umbrella-like.  The  hymenophore  is 
found  on  the  free  or  downward-turned  surface,  and  always  takes 
the  form  of  soft  emergencies  hanging  vertically  downwards.  The 
emergencies  may  be  thorn-,  awl-,  or  wart-like.  The  species  are 
found  growing  on  the  soil  and  on  dead  wood. 

GENERA  :  Hydnum  has  subulate,  distinct  emergences.  H.  rcpandum  is 
yellow,  the  stalk  being  placed  in  the  centre  of  the  pileus.  It  is  an  edible 

1  In  the  resupinate  fruit-bodies  a  fertile  and  sterile  surface  cannot  be  dis- 
tinguished (cf.  Polyporaceae  and  some 


BASIDIOMYCETES 


163 


species,  and  often  forms  "  fairy  rings  "  in  woods.  H,  auriscalpium  (Fig.  166) 
is  dark-brown,  with  stalk  placed  at  the  edge  of  the  pileus.  It  grows  on  old  Fir- 
cones. H.  erinaceus  grows  on  old  tree-trunks.  The  fruit-body  is  yellow  and 
very  large — as  big  as  a  human  head — with  emergences  as  much  as  an  inch  in 
length. — Irpex  has  a  leathery  fruit-body,  partly  resupinata,  partly  with  free, 
projecting  edge ;  the  under  side  bears  tooth-like  emergences  which  are  arranged 
in  rows,  and  Irpex  thus  forms  a  transition  to  the  Agaricacese. — Phlebiais  entirely 
resupinate,  with  radially-arranged  folds  on  the  free  side,  and  pectinate  border. 


FIG.  166. — Hydiium auriscalpium,  upon  a  Fir-cone,  in  different  stages  of  development. 

Order  5.  Polyporacese  (Pore- Fungi).  An  order  very  rich 
in  species  (about  2000  species  are  described).  The  fruit-body  is 
of  very  different  forms — resupinate,  projecting  like  a  bracket, 
hoof-like,  or  umbrella-shaped.  In  some  it  is  fleshy  and  edible, 
in  others  leathery  or  corky,  persisting  for  several  years.  The 
hymenophore  is  situated  on  the  under  side  of  the  fruit-body,  and 
consists  of  wide  or  narrow  tubes  or  pores,  whose  inner  surface  is 
clothed  with  the  hymenium  (Fig.  167).  In  some  fruit-bodies  large 
cavities  are  to  be  found,  which  have  arisen  as  interstices  between 
the  labyrinthine  curved  and  reticulate  folds.  Chlamydospores  are 
known  in  some  species.  Conidia  occur  very  rarely.  Many  species 
work  considerable  damage  :  some  as  parasites  on  trees,  others  by 
destroying  timber. 

GENERA.  Polyporus  (Pore-Fungus).  The  tubes  are  narrow,  accurately  fitted 
together,  and  forming  a  thick  layer  on  the  under  side  of  the  fruit-body,  appearing 


164 


BASIDIOMYCETES. 


as  a  number  of  fine  holes.  The  fruit-body  most  frequently  resembles  a  bracket, 
or  is  hoof-shaped,  with  one  side  growing  from  a  tree-trunk  ;  it  is  very  often  per- 
ennial, and  a  new  layer  of  tubes  arises  in  each  succeeding  period  of  vegetation. 
Strata,  corresponding  to  the  periodically  interrupted  growth,  are  thus  formed  in 
stories  one  above  the  other,  and  are  visible  on  the  upper  surface  of  the  fruit- 
body,  as  well  as  in  the  interior,  as  a  series  of  concentric  belts,  sometimes  as 
many  as  half  a  score  or  more  in  number.  P.  fomentarius  (Touchwood)  attacks 
trees,  especially  the  Beech.  The  spores  germinate  on  wounds  from  broken 
branches,  and  the  hyphae,  following  the  course  of  the  medullary  rays,  find  their 
way  into  the  interior  of  the  tree,  from  whence  the  mycelium  spreads  upwards, 
downwards,  and  peripherally,  so  that  the  wood  becomes  rotten  ("  wh  te-rot") 
and  thick  felts  of  mycelium  are  formed  iu  radial  and  tangential  directions.  A 

dark  line,  caused  by  the 
youngest  parts  of  the  hy- 
phffi  containing  a  brown 
juice,  marks  the  boundary 
between  the  rotten  and 
the  unattacked  parts  of 
the  stem  (Fig.  168);  at 
places  where  the  mycelium 
extends  to  the  bark,  the 
cambium  becomes  de- 
stroyed and  further  growth 
is  arrested,  so  that  longi- 
tudinal furrows  arise  on 
the  stem.  It  is  at  these 
places,  too,  that  the  boof- 
shaped,  ash  -  coloured 
fruit-bodies  are  developed, 
which  may  attain  a  cir- 
cumference of  upwards  of 
7  feet.  The  interior  of 
the  fruit-body  consists  of 
a  dried-up,  loosely  felted, 
red-brown  mass  of  hyphas, 
which  has  been  used  for 
tinder  and  as  a  styptic 
("Fungus  chirurgorum"). 
P.  igniarius  has  a  harder, 
dark-brown,  more  rounded 
fruit- body  ;  it  grows  in  a 
similar  manner,  but 

especially  attacks  Oaks,  Poplars,  and  Plum-trees,  the  wood  of  which  becomes 
rotten,  and  is  called  touchwood.  P.  plni  (Trametes  pint),  (Fig.  170),  a  parasite 
on  the  stems  of  Pimts,  causes  a  kind  of  "  red-rot  "  in  the  stem.  P.  sulphureus 
has  a  soft,  cheesy,  yellow  fruit-body;  it  produces  "  rot"  in  Oaks  and  Apple- 
trees.  P.  ojllcinalis,  Larch-fungus  ("  Fungus  Laricis  "  in  Pharmocopoeia),  grows 
on  Larch- tr<.es  in  the  south-east  of  Europe.  P.  versicolor  has  thin,  semicircular 


FIG.  107.— Polyporus  igniarius.  Section  through  the 
under  side  of  the  Fungus  :  h-h  is  hyphal-tissue  between 
the  tubes,  formed  by  irregularly  felted  hyphse,  many  of 
which  are  seen  cut  across;  s  is  the  hymenium  which 
covers  the  walls  of  the  tabes,  and  from  which  the  basidia 
with  the  spores  protrude. 


i 


BASIDIOMYCETES. 


165 


fruit-bodies,  with  zones  of  various  colours  on  the  upper  side  ;  it  is  one  of  the 
most  frequent  species  on  tree-stems.  P.  frondosus  grows  on  soil  in  woods,  and 
consists  of  numerous  aggregated  fruit-bodies,  which  become  very  large  and 
fleshy.  This  species  is  edible.  P.  perennis  also  grows  on  the  soil  in  woods  ; 
it  is  very  leathery,  with  central  stalk,  and  has  concentric  zones  on  the  upper 
surface  of  the  fruit-body.  P.  vaporarius  destroys  the  wood  of  living  Pines 
(Pinus  silvestris)  and 
Firs  (Picea  excelsa), 
causing  it  to  become 
red-brown ;  in  timber 
this  Fungus  causes 
"red-strip"  followed 

by  a  "dry-rot."  P.  *JHn^g&%@jffi&®mi6^^^®&  t 
squamosus  destroys 
many  Walnut  -  trees, 
and  is  also  very  de- 
structive to  Limes  and 
Elms.  P.  fulvus  causes 
a  "white -rot"  in  Abies 
alba. 

Heterobasidion  an- 
nosum  (Polyporus  an- 
nosus,  Trametes  radici- 
perda,  Fig,  169)  is 
characterized  by  its 
Aspergillus-like  coni- 
diophores.  It  is  a 
parasite  on  the  Pine, 
Fir,  Birch,  Beech,  etc., 


FIG.  168. — Section  of  stem  of  a  Beech  attacked  by  P.  fomen- 
tarius  :  anon-attacked  pares  of  the  stem;  b  the  furrows  where 
the  mycelium  has  reached  the  bark,  and  where  the  thick 
mycelium-strands  reach  the  exterior  ( Jth  of  the  nat.  size). 


FIG.  169.— Base  of  a  Fir-tree,  with  a  number 
of  fruit-bodies  of  Heterubnsidion  annosum  just 
beneath  the  surface  of  the  soil,  indicated  by 
the  dotted  line  (ith  nat.  size). 


FIG.  170.— A  fully  developed  fruit- 
body  of  Polyporus  pini  (Trametes 
pini),  lateral  view  (nat.  size). 


166 


BASIDIOMYCETES 


and  is  the  chief  cause  of  a  root-disease  (red-rot)  in  Pines  and  Firs ;  the  fruit- 
bodies  develope  a  large  number  of  basidiospores ;  they  may  be  very  large  and 
are  found  just  beneath  the  surface  of  the  soil  (on  living  or  dead  roots),  and 
exposed  to  the  air  (on  felled  stems  and  roots,  in  Scandinavia). 

PtychogastcY  has  cushion-like  fruit-bodies,  which  consist  chiefly  of  chlamy- 
dospore-chains,  formed  of  ellipsoidal  spores,  which  alternate  with  short  hyphae 
having  transverse  septa  and  clamp-connections.  The  hymenial  portion  is 
limited  to  a  small  group  of  tubes.  Pt.  albus  (Oligoras  ustilayinoides)  grows  on 
stumps  of  Conifers  and  forms  irregular  cushions,  at  first  white  and  later  on 
"brown,  which  consist  almost  entirely  of  chlamydospores. 

Boletus  (Fig.  171)  has  a  fleshy  fruit-body  resembling  a  common  Mushroom, 
with  central  stalk.     The  layer  of  tubes  is  easily  detached  from  the  pileus,  and 
the  tubes  are  easily  separable  from  one  another.     They  grow  on  the  ground  in 
woods.     Edible  species  are:  B.  edulis,  with  thick,  reticulate  stalk;  B.  scaber, 
with  thin  stalk  and  rough  pileus  ;  B.  luteus,  with 
a  ring  on  the  stalk.     B.  luridus   is   poisonous, 
its  tubes  have  red  openings,  and  the  flesh  turns 
quickly  blue  when  broken  and  exposed  to  the  air. 
Fistulina  liepatica  (Beef-steak  Fungus),  has  a 
red,  fleshy,    edible   fruit-body,   with  red  juice. 
The  tubes  are  individually  distinct ;   conidia  are 
also  developed.     Grows  on  old  Oaks. 

Merullus  lacrymans  ("  Dry-rot")  has  a  resu- 
pinate  fruit-body  with  white,  cotton-like  border, 
and  the  remaining  portions  covered  by  reticulate, 
ramified  veins  of  a  rust-brown  colour.  In  favour- 
able vegetative  conditions  it  is  fleshy  and  exudes 
large  drops  of  water — hence  its  specific  name 
and  also  the  name  "  Tear  Fungus."  The  my- 
celium is  at  first  colourless,  and  then  yellow- 
brown  ;  when  dry  it  is  tough  and  leathery.  It 
destroys  tbe  timber  in  damp  houses,  extends  far 
and  wide  over  boards  and  beams  and  even  over 
the  masonry,  giving  rise  to  a  disagreeable  smell 
in  the  rooms  in  which  it  lodges.  In  woods  the 
Fungus  lives  on  Pine-stems.  It  is  brought  from  the  forest  on  the  logs  of  timber, 
and  is  distributed  from  log  to  log  by  the  mycelium  and  the  basidiospores. 
The  living  mycelium  can  be  recognised  by  the  clamp-connections  shooting  out 
branches.  The  basidiospores  are  often  ejected  a  distance  of  a  metre  ;  they  are 
elliptical  (10-11  /j,  long  and  5-6  //,  broad),  and  germinate  easily  on  damp  wood, 
or  in  fruit-juice  which  has  been  neutralized  with  urine  or  alkaline  carbonates. 

Dccdalea  (Labyrinth  Fungus),  has  bracket-like,  corky  fruit-bodies  with  irregu- 
larly-folded plates  or  discs  on  the  under  side.  It  forms  a  transition  to  the 
Agaricacea3.  D.  quercina  is  frequent  on  Oak-stumps. 


FIG.  171.— Boletus  edulis  (about 
Jth)  :  "b,  longitudinal  section  of 
a  portion  of  the  pileus. 


Order  6.    Agaricaceae    (Mushrooms,    Toadstools).      The 

Jiymenophore  consists  of  knife-like  plates  (lamellce,  gills),  which  are 
situated  on  the  under  side  of  the  umbrella-like  pileus  of  the  fruit- 


BASIDIOMYCETES. 


167 


body,  and  radiate  from  the  central  stalk.  Those  which  are  first 
formed  extend  from  the  edge  of  the  pileus  to  the  stalk;  those 
formed  later  reach  only  a  longer  or  shorter  portion  of  this  distance, 
according  to  their  age.  In  structure  the  lamellae  (Fig.  174)  con- 
sist of  a  central  mass  of  hyphae,  the  trama,  continuous  with  the 
hyphge  of  the  pileus ;  these  terminate  in  a  layer  of  shorter  cells, 
the  sulky  menial  layer,  immediately  beneath  the  hymenium  which 
is  composed  of  basidia  and  paraphyses.  In  a  few  species,  but  not 
in  the  majority,  the  lamellae  are  branched,  and  in  some  they  are 
dec uiTent.  A  few  have  the  stalk  placed  excentrically,  or  it  may 
be  entirely  absent. 


FIG.  172. — Development  of  Psalliota  campestris :  a,  b,  c,  d  show  the  various  stages  of 
the  development  of  the  fruit- bodies  and.  the  mycelium  (m)  (nat.  size);  e  the  fruit-body 
in  a  somewhat  later  stage,  slightly  magnified  ;  /  longitudinal  section  of  e  ;  n  first  forma- 
tion of  the  hymenium ;  g  longitudinal  section  of  a  more  advanced  fruit-bsdy  (nat.  size) ; 
n  the  hymenium  ;  o  velum  partiale  (see  Fig.  133.) 

In  the  early  stages  of  its  development  the  fruit-body  is  more  or 
less  enclosed  in  a  hyphal  tissue — the  "  veil  "  (velum  unicersale,  or 
volva).  The  veil  at  first  completely  encloses  the  young  fruit-body, 
but  is  afterwards  ruptured  as  the  latter  grows,  part  remaining 
at  the  base  of  the  stalk  as  the  "  sheath  "  (annulus  inferus)',  and 
part  on  the  pileus  as  scales  or  warts.  In  the  "  Fly  Mushroom  " 
(Amanita  muscaria)  the  remains  of  the  veil  are  especially  con- 


168 


BASIDIOMYCETES. 


spicuous  as  white  patches  on  the  bright  red  ground  of  the  upper 
surface  of  the  pileus,  and  as  a  sheath  at  the  base  of  the  stalk  (Fig. 
178  v.).  Another  veil — the  velum  partiale — a  hyphal  tissue  (Figs. 
178  a;  173)  stretches  from  the  edge  of  the  pileus  to  the  stalk,  and 
encloses  the  lamellae.  This  veil  is  ruptured  as  the  pileus  expands, 
a  portion  attached  to  the  stalk  remaining  as  the  "  upper  ring  " 
(annulus  snperus]  (Figs.  173,  178  a),  or  a  part  attached  to  the 
pileus  hanging  down  as  a  fringe  round  its  edge. — Some  genera 
have  no  veil,  the  under  side  of  the  pileus  being  exposed  from  the 
first  (gymnorarpic  Agaricaceae) .  Those  which  have  a  veil  (hemi- 
angiocarpicA.)  afford  a  transition  to  the  angiocarpic  Grasteromycetes. 


FIG.  173.— The  cultivated  Mushroom  (Psalliota  campestris). 

The  mycelium  mostly  grows  in  soils  rich  in  humus  or  dung,  on 
decaying  trees  and  similar  objects.  Many  species,  e.g.  Tricholoma 
personatum  and  Marasmius  oreadcs,  form  the  so-called  "  fairy  rings." 
The  fruit-bodies  in.  these  species  are  confined  to  a  larger  or  smaller 
surface  on  which  they  are  very  regularly  arranged  in  a  ring.  The 
reason  for  this  is  found  in  the  radial  growth  of  the  mycelium,  so 
that  the  oldest  portion,  or  the  starting  point,  is  found  at  the  centre 
of  the  ving,  and  the  younger  ones,  on  which  the  fruit-bodies  are 
formed,  at  the  circumference.  The  older  hyphae  gradually  die,  arid 
at  the  same  time,  the  radial  growth  continuing,  the  ring  of  fruit- 
bodies  becomes  larger  and  larger.  The  "  fairy-rings  "  are  marked 


BASIDIOMYCETES. 


169 


not  only  by  the  fruit-bodies,  but  also  by  the  more  vigorous  growth 
and  darker  colour  of  the  grass  upon  these  spots. 

Some  species  are  parasites.  An  example  is  presented  by  Armil- 
laria  mellea,  a  remarkable  and  very  destructive  Fungus  in  woods  and 
forests  (Figs.  176,  177).  In  addition  to  the  filamentous,  white  mycelium, it 
has  also  black,  or  black-brown,  horny,  root-like  mycelium-strands  (rhizoinorpba) 
which  were  "formerly 
considered  to  belong  to 
a  special  genus  of  Fungi 
described  under  the 
name  "  Rhizomorpha." 
The  mycelium  lives 
parasitically  on  the  Con- 
ifers and  other  trees, 
forcing  its  hyphse  into 
the  bark  and  between 
the  bark  and  wood,  and 
thence  penetrating  into 
the  wood  so  that  the 
tree  is  very  severely  at- 
tacked. It  may  also 
live  saprophytically,  and 
clusters  of  fruit-bodies 
are  often  found  on  old 
stumps  and  stems,  on 
old  timber,  and  in  the 
rich  soil  of  woods.  The 
rhizomorpha,  living  un- 
derground, can  extend 
for  considerable  dis- 
tances and  infect  the 
roots  of  neighbouring 
trees,  and  spreads  in 
tbis  way  the  diseases 
known  as  "Harzsticken" 
and  "Bark -Canker," 
which  are  very  destruc- 
tive to  young  trees. 


FIG.  174. — Psalliota  campestris.  A  Tangential  section  of 
pileus  showing  lamellae  (1).  B  Portion  of  gill  more  highly 
magnified;  ttrama;  hy  hymenium \vithbasidia and  basidio- 
spores;  sTi,  sub-hymenial  layer.  C  A  portion  of  the  same 
more  highly  magnified ;  s'  s"  s'"  s""  various  stages  in  the- 
development  of  basidiospores ;  q  paraphyses. 


The  chief  charac- 
teristics by  which 
the  numerous  genera 
are  separated  are 
the  presence  or  the  absence  of  the  two  kinds  of  veils,  the  nature 
of  the  fruit-body,  the  form,  branching  of  the  lamellee,  and  their 
position  and  relation  with  respect  to  the  stem,  the  shape  of  the- 


170 


BASIDIOMTCETES. 


pileus,  the  colour  of  the  spores,  etc.,  etc. 
colour  may  be  obtained  by  plac- 
ing the  pileus  with  the  lamellte 
turned  downwards  on  a  piece  of 
white  or  coloured  paper,  so  that 
the  spores,  as  they  fall  off,  are 
collected  on  the  paper,  and  the 
arrangement  of  the  lamellae  can 
then  be  clearly  seen. 


A  knowledge  of  the 


FIG  175. — Cantharellus  cibarius  (reduce^). 


FIG.  176.— Armillaria  mellea  (£  nat.  size) : 
o  root  of  a  fir;  b  rhizomorpha-strands  ; 
c-f  fruit-bodies  in  four  different  stages  of 
development. 


•jrrG-  177  —The  mycelium  of  Armillar>'a  mellea  ("E/u'zomorpTia")  (nat.  size). 


BASIDIOMYCETES. 


171 


About  4,600  species  belonging  to  this  order  have  been  described. 

On  account  of  the  large  number  of  species  the  order  is  divided  into  several 
sections  : 

1.  Agaricinei ;  fruit-body  fleshy ;  lamellae    membranous,    knife-like,    with 
sharp  edge  ;  basidia  crowded  together.     The  FOLLOWING  HAVE  WHITE  SPORES  : — 
Amanita  (Fly  Mushroom),  with  volva,  and  generally  also  the  upper  ring  on  the 
stalk;  many  are  poisonous,  such  as  A.  muscaria  (Fig.  178)  which  has  bright 
red  pileus  with  white  spots,  A.  pantherina  and  A.plialloides;  A.  ccesarea  is  edible. 
— Lepiota  procera  (Parasol  Fungus)  is  one  of  the  largest  Mushrooms  ;  it  has  a 
scaly  pileus  and  moveable  ring  (edible). — Armillaria  mellea  has  been  mentioned 
above  (Figs.  176,  177). — Tricholoma,  lamellae  indented  near  the  stalk;  T.  gam- 
bosum  (Pomona  Fungus)  belongs  to  the  best  of  edible  Fungi ;    T.  personatum 
often   forms  fairy  rings 

(see  above).  —  Clitocybe, 
lamella  decurrent  ;  G. 
nebularis  is  edible. — 
Pleurotus,  stalk  eccen- 
tric ;  P.  ostreatus  (Oyster 
Mushroom)  grows  in 
clusters  on  tree-stems 
(edible). —  Collybia  and 
Mycena,  species  numer- 
ous, small.  —  SPORES 
ROSE-RED  :  Volvaria  and 
Hyporhodius.  —  SPORES 
BROWN  :  Cortinarius, 
with  cob  web -like  veil ; 
Pholiota,  membranous 
veil  and  ring  ;  P.  squar- 
rosa  in  clusters  on  tree- 
stems  ;  P.  mutabilis,  on 
tree-stumps  (edible). — 
SPORES  VIOLET-PUBPLE  : 
Hyphcloma,Psalliota;  to 
this  section  the  common 
edible  Mushroom  (Fig. 
172-174)  belongs,  with  annulus  and  chocolate-coloured  lamellas ;  it  is  cultivated" 
for  the  sake  of  the  fine  flavour.— SPORES  BLACK:  Coprinarius. 

2.  Marasmiei.    Fruit-body  tough,  almost  leathery,  and  persistent ;  spores 
white.     Marasmius  oreades  forms  large,   regular  fairy-rings  on  pastures  and 
commons  ;  it  is  used  as  seasoning  in  food. — Panus  stipticus  with  eccentrically- 
placed  stalk,  in  clusters  on  tree-stumps. — Schizophyllum  has  the  edge  of  the 
lamellas    divided    longitudinally,    and  the   split   portions   revolute. — Lentinus 
affords  a  transition  to  Dadalea  among  the  Polyporaceee. 

3.  Russulei.    Fruit-body  fleshy  and  fragile,  in  which  two  different  systems 
of  hyphae  may  be  distinguished  ;  spores  thorny,  white,  or  pale-yellow.   Many  are 
poisonous. — Russula  has   generally   fragile  and   thick   lamellae  reaching  from 
stalk  to  edge  of  pileus  ;  pileus  frequently   red. — Lactarius  has  white  or  yellow 


FIG.  178. — Fly  Mushroom  (Amanita  muscaria). 


172  BASIDIOMYCETES. 

milky  juice,  which  often  is  very  acid.    L.  deliciosus  has  red-yellow  milky  juice, 
and  is  of  a  pleasant  flavour.     L.  torniinosus  is  poisonous. 

4.  Hygrophorei.     Lamellae    thick   and   waxy,   widely  separated  ;     spores 
white.     Many  species  of  Hygrophorus  have  brightly- coloured  pileus  and  grow 
among  the  grass  on  moors  and  commons. — Nyctalis  is  parasitic  on  larger  Toad- 
stools.    It  is  remarkable  for  its  abundant  formation  of  chlamydospores,  whilst 
the  basidiospores  are  little  developed. 

5.  Coprinei.     Fruit-bodies  very  soft,  quickly  perishable;  lamellae  membran- 
ous and  deliquescent.      The  basidia  are  separated  from  each  other  by  para- 
physes.     Coprinus  has  coal-black   spores,  grows   on  manure,  and  sometimes 
developes  sclerotia. 

6.  Paxillei.    Fruit-body  fleshy;  lamellae  easily  detached  from  the  pileus  and 
reticulately -joined  near  the  stalk.     They  form  a  connecting  link  between  the 
Agaricaceae  and  Boletus. 

7.  Cantharellei.     Lamellae  reduced  to  dichotomously-divided  folds,  decur- 
rent  on  the  stalk.    Cantharellus  cilarius  (Fig.  175)  is  yolk-yellow,  and  grows  on 
the  ground  in  woods  (edible).     It  is  allied  to  C rater ellus. 

Family  3.     Phalloidese. 

The  fruit-bodies  before  they  are  ripe  are  spherical  or  ovoid,  and 
enclosed  by  a  flesliy  covering,  the  peridium,  which  is  perforated  at 
'maturity  and  remains  as  a  sheath  (Fig.  179)  ;  the  fruit-bodies 
are  hemiangiocarpic. 

Order  1.  Phallacese  (Stink-horns).  The  peridinm  has  a 
•complicated  structure  and  is  composed  of  three  layers,  the  in- 
termediate one  being  thick  and  gelatinous.  The  gleba  (the  tissue 
which  bears  the  hymenium)  is  situated  upon  a  peculiar  receptacle 
which  expands  into  a  porous  stalk  and  by  its  sudden  distension, 
rupturing  the  peridium,  elevates  the  gleba  and  hymenium  above 
the  peridium,  which  remains  as  a  sheath.  The  gleba  becomes 
gelatinous  and  dissolves  away  as  drops.  To  this  order  belong  many 
peculiar  and  often  brightly  coloured  forms,  which  are  natives  of 
the  Southern  Hemisphere. 

Phallus  impudicus  (Stink-horn)  (Fig.  179),  has  a  fruit-body  which  at  first  is 
-white,  heavy,  and  soft,  and  resembles  a  hen's  egg  in  shape  and  size.  The  peri- 
dium is  divided  into  three  layers  (Fig.  179  e,  g,f)  of  which  the  external  and 
internal  are  membranous,  and  the  middle  one  very  thick  and  gelatinous;  each 
of  these  has  again  a  laminated  structure.  The  peridium  when  ruptured  re- 
mains as  a  sheath  (/c)  at  the  base  of  the  stalk.  The  receptacle  at  first  is 
strongly  compressed  (ft)  but  afterwards  expands  into  a  long  stalk  (I)  which 
bears  the  conical  gleba  (m).  Prior  to  the  rupture  of  the  peridium  the  gleba 
consists  of  a  greenish  mass  (i)  which,  when  exposed,  emits  a  carrion-like  stench 
serving  to  attract  flies,  by  whose  agency  the  spores  are  distributed.  It  is  found 
commonly  in  hedgerows  and  in  woods,  growing  on  the  ground.  The  much 


BAS1DIOMYCETES. 


173 


smaller  arid  less  common  P.  caninus  is  found  on  rotten  tree-stumps. — In 
Clathrus  cancellatus  the  receptacle  expands  into  a  bright  red,  reticulate  struc- 
ture. A  native  of  the  South  of  Europe.  Colus,  Aseroe,  Mitromyces. 

Order  2.  Sphaerobolaceae.  An  intermediate  layer  of  the  peridium  swells 
when  ripe,  becomes  convex,  and  ejects  the  remaining  spherical  portion  of  the 
fruit-body  which  contains  the  spores.  Sphcerobolm  carpobolw  has  small, 
spherical  fruit-bodies  which  open  in  the  form  of  a  star. 


FIG.  179.— Phallus  impudicus  (Stink-horn),  somewhat  diminished.  Fruit-bodies  in  all 
stages  of  development  (b,  c,  d  and  fc-m)  are  seen  arising  from  a  root-like  mycelium  (a) ;  d 
longitudinal  sections  through  a  fruit-body  b3fore  the  covering  has  ruptured. 

Family  4.     Gasteromycetes. 

The  fruit-body  is  angiocarpic,  fleshy  at  first,  and  later  generally 
more  or  less  hard  and  continues  closed  after  the  spores  are  ripe. 
The  tissue  lying  immediately  inside  the  peridium  is  termed  the 
gleba;  it  is  porous,  containing  a  larger  or  smaller  number  of 
chambers  lined  with  the  hymenium,  which  is  either  a  continuous 


174 


BASIDIOMYCETES. 


layer  of  basidia  or  else  it  fills  up  the  entire  cavity.  The  basidia 
as  a  rule  bear  four  spores,  sometimes  eight  (Geaster),  or  two- 
(Hymenogaster).  The  tissue  of  the  walls  (trama)  consists  often 
(Lycoperdacece}  of  two  kinds  of  hyphse,  some  thin  and  rich  in 
protoplasm,  divided  by  transverse  septa  and  bearing  the  basidia  ; 
others  thicker  and  thick- walled  which  do  not  dissolve  like  the 
former  on  the  ripening  of  the  spores,  but  continue  to  grow  and 
form  a  woolly,  elastic  mass,  the  capillitium,  which  may  be  regarded 
as  highly  developed  paraphyses.  The  peridium  may  be  either 
single  or  double,  and  presents  many  variations  in  its  structure 
and  dehiscence.  The  mycelium  is  generally  a  number  of  string- 
like  strands,  living  in  soils  rich  in  humus. 

Order  1.  Tylostomaceae.  Capillitium  present.  After  the  rupture  of  the 
peridium  the  remaining  part  of  the  fruit-body  is  elevated  on  a  long  stalk. 
Tylostoma  wainmosum,  on  heaths. 

Order  2.  Lycoperdaceae.  The  fruit-body  has  a  double 
peridium ;  the  external  one  at  length  breaks  into  fragments 
(Lycoperdon,  Bovista),  or  it  has  a  compound  structure  of  several 
layers  (Geaster)  and  detaches  itself  as  a  continuous  envelope  from 
the  inner  layer,  which  is  membranous  and  opens  at  its  apex. 
The  interior  of  the  fruit-body  consists  either  solely  of  the  fertile 
gleba  (Bovista,  Geaster),  or,  in  addition,  of  a  sterile  tissue  at  the 
base  (Ly  coper  don).  A  capillitium  is  also  present. 

Lycoperdon  (Puff-ball)  has  a  sterile  part  at  the  base  of  the  fruit-body  which 
often  forins  a  thick  stalk.  The  surface  of  the  peridium  is  generally  covered 

with  warts  or  projections.  When 
young  this  Fungus  is  edible,  but 
when  ripe  it  is  dry,  and  used  for  stop- 
ping the  flow  of  blood.  L.  giyanteum, 
which  is  often  found  growing  in 
meadows,  attains  a  considerable  size, 
its  diameter  reaching  as  much  as 
eighteen  inches.  L.  gemmatum  (Fig. 
180)  is  covered  with  pyramidal  warts ; 
in  woods.  —  Bovista  has  no  sterile 
basal  part ;  the  external  peridium  is 
smooth,  and  falls  away  in  irregular 
patches.  J5.  plumbea,  on  links  near 
the  sea. — Geaster  (Earth-star)  has  an 
external  peridium  composed  of  several 
layers,  which  when  the  fruit-body  opens,  split  into  several  stellate  segments. 
These  segments  are  very  hygroscopic,  and  in  dry  weather  bend  backwards  and  so 
raise  the  inner  peridium  into  the  air.  The  inner  peridium  contains  the  spores 


FIG.  180. — Lycopirdon  gemmatum 
(inat.  size). 


BASIDIOMYCETES. 


175 


and  capillitia.  G.  coliformis  has  several  apertures  in  the  inner  peridium.  The 
other  species  have  only  one  regular  aperture  at  the  apex.  G.  striatus  has  a 
pedicellate  inner  peridium,  with  conical,  striped  peristome.  G.  fomicatus  has 
an  external  peridium  split  into  four  segments.  This  last  and  several  other 
species  produce  "  mycorhiza  "  on  the  roots  of  Conifers. 


Fis.181. — I  Hymenogaster  citrinus  (nat.  size) ;  II  longitudinal  section  through  H.  tener 
(x  5) ;  III  portion  of  a  section  of  H.  calosporus  ;  g  a  chamber  ;  ft  hymenium;  sp.  spores; 
t  trama  (x  178) ;  IV  Rhizopogon  luteolus  (nat.  size);  V  Scleroderma  vulgare,  VI  section  of  V; 
VII  basidia  with  spores  belonging  to  the  same  Fungus. 

Order  3.  Sclerodermataceae.  Capillitium  wanting.  The 
peridium  is  simple  and  thick,  gleba  with  round,  closed  cham- 
bers, which  are  filled  with  basidia. 

Scleroderma  has  a  corky  peridium.   The  fruit-bodies  commence  their  develop- 
ment under  ground.    S.  vulgare  (Fig.  181  V-VII),  has  a  hard,  slaty-black  gleba. 
W.  B.  N 


176  BASIDIOMYCETES. 

Order  4.  Nidulariacese  (Nest-Fungi).  Small  Fungi  of 
which  the  fruit-body  at  first  is  spherical  or  cylindrical  but  upon 
maturity  it  becomes  cupular  or  vase-like,  and  contains  several 
lenticular  "  peridiola "  lying  like  eggs  in  a  nest.  The  peridiola 
are  the  chambers  which  contain  the  hymenium,  covered  by  a  thin 
layer  of  the  gleba,  all  the  remaining  portion  of  the  gleba  becoming 
dissolved.  On  decaying  wood. 

Nidularia  has  spherical  fruit-bodies  containing  a  large  number  of  lenticular 
peridiola,  embedded  in  a  slimy  mass. — Crucibulum  has  fruit-bodies  resembling 
crucibles  with  discoid  peridiola,  each  with  a  spirally-twisted  stalk. — Cyathns 
has  a  fruit-body,  which  when  open  is  campanulate,  with  stratified  peridium, 
and  long-stalked,  lense- shaped  peridiola. 

Order  5.  Hymenogastraceae.  Fruit-bodies  tubercular, 
globose  and  subterranean,  resembling  very  closely  the  Truffles, 
from  which  they  can  only  be  distinguished  with  certainty  by 
microscopic  means.  The  peridium  is  simple,  capillitium  wanting, 
and  the  gleba  encloses  a  system  of  labyrinthine  passages  covered 
with  a  continuous  hymenium.  The  fruit-bodies  persist  for  some 
time,  and  form  a  fleshy  mass,  the  spores  being  only  set  free  by 
the  decay  of  the  fruit-body,  or  when  it  is  eaten  by  animals.  The 
majority  are  South  European.  Hymenogaster,  Melanogaster, 
RUzopogon  (Fig.  181  I-IV). 

APPENDIX  TO  THE  BASIDIOMYCETES  : 
Basidiolichenes  (Lichen-forming  Basidiomycetes). 

Several  Fungi  belonging  to  the  Basidiomycetes  have  a  symbiotic 
relationship  with  Algae  exactly  similar  to  that  enjoyed  by  certain 
Ascomycetes,  and  these  are  therefore  included  under  the  term 
Lichens  (p.  136).  They  are  chiefly  tropical. 

Order  1.  Hymenolichenes.  To  this  order  belong  some 
gymiioca.rpic  forms  :  Oora,  Dictyonema,  Laudatea. l 

Order  2.  Gasterolichenes.  To  this  belong  some  angiocarpic 
forms  :  Emericella,  Trichocoma. 

APPENDIX  TO  THE  FUNGI. 
Fungi  imperfect!  (Incompletely  known  Fungi). 

1.  The  Saccharomyces-forms  are  Fungi  which  are  only 
known  in  their  yeast-conidial  form.  They  are  conidia  of  higher 

1  The  two  last  genera  are  identical,  the  Algal  part  being  a  Scytonema,  that  of 
Cora  a  Chrorwecus ;  while  the  same  Fungus— a  Thelephora — takes  part  in  t  e 
formation  of  all  three  (A.  Moller,  Flora,  1893). 


JTUNGI    IMPERFECTI 


177 


Fungi  which  can  multiply  to  an  unlimited  extent  by  budding  in 
nutritive  solutions,  and  in  this  way  maintain  their  definite  size  and 
shape.  The  budding  takes  place  only  at  the  ends  of  the  conidia. 
The  wall  of  the  conidium  forms  at  one  or  at  both  ends  a  small  wart- 
like  outgrowth,  which  gradually  becomes  larger,  and  is  finally 
separated  from  its  mother-cell  as  an  independent  cell,  surrounded 
by  a  closed  cell-wall  (Fig.  182  a,  6). 


FIG.  182.— Beer-yeast  (SaccTiaromt/ces  cerevisice)  :  a-b  (x  400);  c-/(x  750)  ;  c  a  cell  in  the 
process  of  forming  spores ;  d  a  cell  vyith  four  ripe  spores  ;  e  the  spores  liberated  by  the 
•dissolution  of  the  cell-wall:  /  three  germinating  spores;  g  mycelium-like  cell-chains. 
<x  1000:  after  Em.  Chr.  Hansen.) 

Under  very  favourable  conditions  multiplication  occurs  so 
rapidly  that  the  daughter-cells  themselves  commence  to  form 
buds,  before  they  have  separated  from  their  mother-cell,  with  the 
result  that  pearl-like  chains  of  cells  are  produced.  When  the 
yeast-cells  have  only  limited  nutriment,  with  an  abundant  supply 


178 


FUNGI   IMPERFECT1 


of  air,  at  a  suitable  'temperature,  an  endogenous  formation  of 
spores  takes  place.  The  protoplasm  of  the  cells  divides  into  1-4 
(rarely  a  greater  number)  masses  (Fig.  182  c,  d,  e)  which  surround 
themselves  with  a  thick  cell- wall,  and  in  this  state  can  with- 
stand adverse  conditions  and  periods  of  dryness  lasting  for  several 
months. 

The  sporangia  are  not  asci  since  they  have  no  definite  form,  and 
a  definite  number,  form  and  size  of  spores  is  not  found.  The 
spores  in  the  different  species  and  kinds  occupy  varying  periods 
for  their  development,  although  exposed  to  the  same  temperature, 
a  fact  of  importance  in  determining  one  from  another.  On 
germination  the  wall  of  the  mother-cell  is  destroyed,  and  each 
spore  gives  rise  to  a  new  cell,  multiplication  taking  place  by 

budding  (Fig.  182 /).  The  majority 
of  Yeast-Fungi  are  able  to  produce 
alcoholic  fermentation  in  saccharine 
fluids. 

The  most  important  of  these 
Fungi  is  the  Beer-yeast  (Saccharo- 
mycescerevisicB)  with  ovate,  ellipsoidal 
or  spherical  cells  (Fig.  182).  It  is 
a  plant  which  has  been  cultivated 
from  time  immemorial,  on  account 
of  its  property  of  producing  alco- 
holic fermentation  in  sugar-contain- 
ing extracts  (wort),  derived  from 
germinating  barley  (malt).  Car- 
bonic acid  is  also  set  free  during  this  process.  The  4>  surface- 
yeast  "  (Fig.  182  a),  which  produces  ordinary  beer  when  the 
brewing  takes  place  at  higher  temperatures,  has  cell-chains ;. 
"  sedimentary  yeast  "  (Fig.  182  6),  used  in  the  brewing  of  Bavarian 
beer,  has  spherical  cells,  solitary,  or  united  in  pairs.  Both  these 
and  the  following  Yeast-Fungi  include,  according  to  Hansen, 
several  species  and  kinds. 

The  "  Ferment  of  Wine"  (Saccharomyces  ellipsoideus)  produces 
wine  in  the  juice  of  grapes.  Uncultivated  yeast-cells  are  always 
present  on  grapes  ;  an  addition  of  this  species  to  the  "  must  "  is 
not  necessary  to  secure  fermentation.  A  large  number  of  other 
"  uncultivated  "  yeast-cells  appear  in  breweries  mixed  with 
the  cultivated  ones,  and  cause  different  tastes  to  the  beer  (S. 
pastorianus,  etc.).  8.  ludwigii,  found,  for  instance,  on,  the  slimy 


FIG.  183. — Saccharomi/ces  mycoderma. 


FUNGI    IMPERFECTI  179 

discharge  from  Oaks,  produces  abundant  cell-chains  on  cultiva- 
tion. S.  apiculatus  is  very  frequently  met  with  on  all  kinds  of 
sweet  fruits,  it  has  orange-like  cells.  S.  mycoderma  has  cylindri- 
cal cells,  often  united  together  in  chains  (Fig.  183)  :  it  forms  a 
whitish-gray  mass  ("  fleur  de  vin  ")  on  wine,  beer,  fruit-juice,  etc., 
standing  in  bottles  uncorked  or  not  entirely  filled.  It  is  thought 
that  this  Fungus  causes  decomposition  and  oxydises  the  fluid  in 
which  it  is  found,  but  it  cannot  produce  alcoholic  fermentation  in 
saccharine  liquids,  and  it  does  not  form  endospores  ;  hence  it  is 
uncertain  whether  it  is  true  Saccharomyces. 

The    "Dry-yeast"    used   in   baking  white    bread  is  "surface- 


FIG.  181.— Oidium  lactis :  a  branched  hypha  commonly  met  with  ;  b  a  hypha  lying  in 
milk  and  producing  aerial  hyphse  which  give  rise  to  oidia ;  c  a  branch  giving  rise  to 
oidia,  the  oldest  (outermost)  oidia  are  becoming  detached  from  one  another ;  d  a  chain 
of  divided  cells;  e  germinating  oidia  in  different  stages  (slightly  more  magnified  than  the 
other  figures)/ 

yeast."  In  leaven,  a  kneaded  mixture  of  meal,  barm  and  water, 
which  is  used  for  the  manufacture  of  black  bread,  Saccharomyces 
minor  is  present,  and  a  species  allied  to  this  produces  alcoholic 
fermentation  in  dough  with -the  evolution  of  carbonic  acid,  which 
causes  the  dough  to  "rise." 

2.  Oidium-forms.  Of  many  Fungi  only  the  Oidium-forms 
are  known,  which  multiply  in  endless  series  without  employing 
any  higher  form  of  reproduction.  Oidium  lactis  (Fig.  184)  is  an 
imperfectly  developed  form  which  frequently  appears  on  sour 


180  FUNGI    IMPERFECTI 

milk  and  cheese.  It  can  produce  a  feeble  alcoholic  fermentation 
in  saccharine  liquids.  Thrush  or  aphthae  (0.  albicans)  appears  as 
white  spots  in  the  mouths  of  children.  Several  similar  Oidium- 
forms  are  parasites  on  the  skin  and  hair  of  human  beings,  and 
produce  skin  diseases,  such  as  scurvy  (0.  schoenleinii)  and  ring- 
worm (0.  tonsurans). 

3.  Mycorrhiza.  Certain  Fungi,  which  have  been  found  on 
the  roots  of  man}'  trees  and  heath-plants,  particularly  Cupuliferse 
and  Ericaceae,  consist  of  septate  hyphse,  and  belong  partly  to  the 
Hymenomycetes,  partly  to  the  Gasteromycetes.  It  has  been  shown 
that  they  enter  into  a  symbiotic  relationship  with  the  roots  of 
higher  plants,  producing  a  condition  known  as  Mycorrhiza. 


DIVISION  II. 

MUSCINE^  (MOSSES). 

In  this  Division  a  well-marked  alternation  of  generations  is  to  be 
found.  The  development  of  the  first  or  sexual  generation  (gameto- 
phyte\l  which  bears  the  sexual  organs,  antheridia  and  archegonia, 
commences  with  the  germination  of  the  spore,  and  consists,  in  the 
Liverworts,  of  a  thallus,  but  in  the  true  Mosses  of  a  filamentous 
protonema,  from  which  the  Moss-plant  arises  as  a  lateral  bud. 
The  second  or  asexual  generation  (sporopliyte) ,  developed  from  the 
fertilised  oosphere,  consists  of  a  sporangium  and  stalk. 

The  sexual  generation,  the  gametophyte.  The  protonema 
in  the  Liverworts  is  very  insignificant,  and  not  always  very  sharply 
demarcated  from  the  more  highly  developed  parts  of  the  nutritive 
system.  In  the  true  Mosses  the  protonema  is  well-developed,  and 
consists  of  a  branched,  alga-like  filament  of  cells,  the  dividing 
cell-walls  being  always  placed  obliquely.  In  the  parts  exposed  to 
the  light  it  is  green,  but  colourless  or  brownish  in  those  parts 
which  are  underground  (Fig.  186).  The  protonema  is  considered 
to  be  a  lower  form  of  the  stem,  and  grows  in  the  same  manner 
by  means  of  an  apical  cell ;  at  its  apex  it  may  directly  develope 
into  a  leaf-bearing  stem,  or  these  arise  from  it  as  lateral  branches 
(Fig.  186  &). 

The  more  highly  differentiated  part  of  the  vegetative  system, 
the  "  Moss-plant,"  which  is  thus  developed  from  the  protonema, 
is  in  the  "  thalloid "  Liverworts  generally  a  dichotomously- 
branched  thallus  without  any  trace  of  leaf-structures  (Fig. 
194)  ;  in  Marchantia  (Fig.  197)  and  others,  scale-like  leaves 
(amphigastria)  are  found  on  the  under  surface.  The  higher 
Liverworts  and  the  Leafy-Mosses  are  differentiated  into  a  fila- 
mentous, ramified  stem  with  distinct  leaves  arranged  in  a  definite 
manner,  resembling  the  stem  and  leaves  of  the  higher  plants  (Figs. 
186,  195,  200). 


1  Formerly  termed  oophyte. 
181 


182 


MUSCINE^:. 


True  roots  are  wanting,  but  are  biologically  replaced  by  rhizoids. 
These  are  developed  on  the  stems  or  thallus :  in  the  Liverworts 
they  are  unicellular,  but  in  the  Leafy-Mosses  generally  multicel- 
lular  and  branched.  In  the  latter  group  they  are  considered 
identical  with  the  protonema,  and  may  become  true  protonema, 
and  new  plants  may  be  developed  from  them  (Fig.  186  6). 

The  internal  structure  of  the  sexual  generation  is  very  simple. 
The  leaves  in  nearly  all  cases  are  formed  of  a  single-layered  plate 
of  cells;  in  the  Leafy-Mosses,  however,  a  midrib  is  very  often 


FIG.  166.— -.4  Lower  portion  of  a  Moss-plant  with  rhizoids  (r),  one  of  which  bears  a 
reproductive  bud  (1).  The  dotted  line  indicates  the  surface  of  the  ground;  tlie  portions 
projecting  above  this  become  preen  protonema  (p);  fc  is  a  young  Moss-plant  formed  or. 
one  of  these.  B  Germatining  spore  of  Funaria  hygrometrica,  with  exospore  still 
attached.  C,  D  Older  stages  of  the  protonema. 

formed,  and  sometimes,  also,  marginal  veins ;  and  along  these  lines 
the  leaves  are  several  layers  of  cells  in  thickness.  The  stem  is 
constructed  of  cells  longitudinally  elongated,  the  external  ones  of 
which  ai-e  narrower  and  sometimes  have  thicker  walls  than  the 
more  central  ones.  Vessels  are  not  found,  but  in  several  Mosses 
there  is  in  the  centre  of.  the  stem  a  conducting  strand  of  narrow, 
longitudinal  cells,  which  represents  the  vascular  bundle  in  its  first 


MUSCINEJ;. 


183 


stage  of  development.  This  strand  contains  elements  for  convey- 
ing water  as  well  as  sieve-tubes.  Stomata  are  entirely  wanting  in 
the  sexual  generation  of  the  Leafy-Mosses  ;  they  are  found  in  a 
few  Liverworts  (Marchantia) ,  but  their  structure  is  not  the  same 
as  in  the  higher  plants. 

VEGETATIVE  REPRODUCTION  takes  place  by  gemmee  or  buds  which 
arise  on  the  protenema,  the  rhizoids,  the  thallus,  or  the  shoots,  and 
become  detached  from  the  mother-plant ;  or  else  the  protonema 
and  the  older  parts  of  the  plant  simply  die  off,  and  their  branches 
thus  become  independent  plants.  This  well-developed  vegetative 
reproduction  explains  why  so  many  Mosses  grow  gregariously. 
In  certain  Marchantiacese  special  cupules,  in  which  gemmae  are 
developed,  are  found  on  the  surface  of  the  thallus  (Fig.  197  A,s-s). 


FIG.  187. — Marchantia  poli,morphat 
a  mature  antheridiutn. 


FIG.  188.  — Sperinatozoids. 


Again,  protonema  may  also  arise  from  the  leaves,  and  thus  the 
leaves  may  act  as  reproductive  bodies.  Certain  Mosses  nearly 
always  reproduce  vegetatively,  and  in  these  species  the  oosphores 
are  seldom  fertilised. 

The  first  generation  bears  the  SEXUAL  ORGANS  ;  both  kinds  are 
found  either  on  the  same  plant  (monoecious),  or  on  separate  plants 
(direcious).  In  the  thalloid  Liverworts  they  are  often  situated  on 
the  apex  of  small  stems  (gametophores^,  springing  from  the  surface 
of  the  thallus.  In  the  Leafy- Liverworts  and  true  Mosses  the  leaves 
which  enclose  the  sexual  organs  often  assume  a  peculiar  shape,  and 
are  arranged  more  closely  than  the  other  leaves  to  form  the  so-called 
"  Moss-flower."  The  male  sexual  organs  are  called  anfheridia. 
They  are  stalked,  spheroid,  club-  or  egg-shaped  bodies  whose 
walls  are  formed  of  one  layer  of  cells  (Fig.  187),  enclosing  a  mass 
of  minute  cubical  cells,  each  one  of  which  is  a  mother-cell  of 


184  MUSCINEJ;. 

n  spermatozoid.  The  spermatozoids  are  self-motile ;  they  are 
slightly  twisted,  with  two  cilia  placed  anteriorly  (Fig.  188),  while 
posteriorly  they  are  generally  a  trifle  club-shaped,  and  often  bear 
at  that  part  the  remains  of  the  cytoplasm,  the  spermatozoid  itself* 
being  formed  from  the  nucleus.  In  the  presence  of  water  the  ripe 
antheridium  bursts,  and  its  contents  are  ejected  ;  the  spermato- 
zoids, being  liberated  from  their  mother-cells,  swarm  about  in  the 
water  in  order  to  effect  fertilisation. 

The  female  sexual  organs  are  termed  archegonia.    They  are  flask- 
shaped  bodies  (Fig.  189),  the  lower,  swollen  portion  (renter)  having 


FiQ.  189. — MarcTiantto  polymer  pha.  A  A  young,  and  B  a  ripe  arclifgonium  with  open 
neck.  C  An  unripe  sporangium  enclosed  by  the  archegoniutn  a  :  st  the  s^talk  ;  /  the  wall 
of  the  sporangium.  Elaters  are  seen  between  the  rows  of  spores. 

a  wall,  in  most  cases  from  1-2  cells  thick,  enclosing  the  oosphere 
(Fig.  189  B,  k)  :  the  long  neck  is  formed  of  tiers  of  4-6  cells, 
enclosing  a  central  row  of  cells — the  neck-canal-cells  (Fig.  189  A). 
When  the  archegonium  is  fully  developed,  the  walls  of  the  neck- 
canal-cells  become  mucilaginous  and  force  open  the  neck  of  the 
archegonium.  The  mucilage  thus  escapes,  and,  remaining  at  the 
mouth  of  the  archegonium,  acts  in  a  somewhat  similar  manner  to 
the  stigma  and  conducting  tissue  of  a  carpel,  by  catching  and  con- 
ducting the  spermatozoids  to  the  oosphere  (Fig.  189  B,  m),  with 


185 


whose  cell-nucleus  they  coalesce.  With  regard  to  the  formation 
of  the  oosphere,  it  may  further  be  remarked  that  the  lower  part  of 
the  archegonium  originally  encloses  the  so-called  "  central  cell  "; 
but  shortly  before  the  archegonium  is  ripe,  this  cuts  off  a  small 

portion,  the  ventral-canal-cell,  which 
lies  immediately  beneath  the  neck, 
and  the  larger,  lower  portion  be- 
comes the  oosphere. 

The  organs  mentioned  here,  antheridia 
and  archegonia,  are  present  in  the  Crypto- 
gams (Pteridophyta)  and  the  Gymno- 
sperms.  They  have  always  the  same 
fundamental  structure,  but  with  slight 
modifications  of  detail.  These  plants  are 
therefore  known  as  the  ARCHEGONIATA. 

The  fertilisation  of  the  Mosses 
cannot  be  effected  without  water. 
Rain  and  dew  therefore  play  a 
very  important  part  in  this  pro- 


Fm.  190.— Avdrecea  rupestri*.  Longi- 
tudinal section  through  a  sporangium 
at  the  time  when  the  mother-cells  of 
the  spores  are  dividing :  p  pseudo- 
podium  ;  /  foot ;  v  vaginula ;  7i  neck ; 
c  columella  ;  w  wall  of  the  sporan- 
gium ;  e  external  row  of  cells;  s  the 
spore-sac ;  t  the  spore-mother-cells ; 
r  the  calyptra  with  the  neck  of  arche- 
gonium (Zj. 


FIG.  191.  —  Andrecea  rupestris.  Transverse 
section  through  a  ripe  sporangium.  In  the 
middle  is  seen  the  four-sided  columella,  sur- 
rounded by  the  numerous  spores,  drawn  dia- 
grammatically.  Surrounding  them  is  seen  the 
wall  of  the  sporangium,  whose  outer  layer  *>£ 
cells  is  thickened  and  coloured.  The  layer  of 
cells  is  unthickened  in  four  places  (a-)»  indi- 
cating the  position  of  the  clefts  (see  Fig. 
193). 


cess,  and  for  this  end  various  modifications  of  structure  are  found. 


186  MUSCINE^E. 

Among  the  sexual  organs,  paraphyses — filamentous  or  club- 
shaped  bodies — are  to  be  found. 

The  asexual  generation,  the  sporophyte  (Moss-fruit  or 
sporogonium).  As  the  result  of  fertilisation  the  oosphere  sur- 
rounds itself  with  a  cell-wall,  and  then  commences  to  divide  in 
accordance  with  definite  laws.1  The  embryo  (Fig.  189  C)  pro- 
duced by  these  divisions  remains  inside  the  wall  a-a  of  the  arche- 
gonium  (Figs.  190,  199  D,  E),  and  developes  into  the  sporogonium, 
which  remains  attached  to  the  mother-plant,  often  nourished  by 
it,  as  if  the  two  were  one  organism.  The  lower  extremity  of  the 
sporogonium,  the  foot  (Figs.  190  /;  199  D),  very  often  forces  its 
way  deep  down  into  the  tissue  of  the  mother- pi  ant,  but  without  an 
actual  union  taking  place.  The  central  portion  of  the  sporogonium 
becomes  a  shorter  or  longer  stalk  (seta),  while  the  sporangium 
itself  is  developed  at  the  summit.  .  At  a  later  stage,  during  the 
formation  of  the  spores,  the  sporangium  very  often  assumes  the 
form  of  a  capsule,  and  dehisces  in  several  ways  characteristic  of  the 
various  genera  (Figs.  192,  193,  194,  195,  200).  The  basal  portion 
of  the  archegonium  grows  for  a  longer  or  shorter  period,  forming 
a  sheath,  the  calyptra,  in  which  the  capsule  is  developed,  but 
eventually  it .  ceases  to  enlarge,  and  is  then  ruptured  in  different 
ways,  but  quite  characteristically,  in  each  group.  Anatomically, 
the  asexual  generation  is  often  more  highly  differentiated  than  the 
sexual ;  thus,  for  instance,  stomata  are  present  on  the  sporangia 
of  the  true  Mosses,  but  are  absent  in  the  sexual  generation. 

As  the  capsule  developes,  an  external  layer  of  cells — the  amphi- 
thecium  —  and  an  internal  mass — the  endothecium — are  differ- 
entiated. As  a  rule  the  former  becomes  the  wall  of  the  capsule 
while  the  latter  gives  rise  to  the  spores.  In  this  Division,  as"  in 
the  Pteridophyta,  the  name  archesporium  (Fig.  ]90  £)  is  given  to 
the  group  of  cells  inside  the  sporangium  which  gives  rise  to  the 
mother-cells  of  the  spores.  The  archesporium  is  in  general  a 
unicellular  layer ;  in  Sphagnum  and  Anthoceros  it  is  derived  from 
the  most  internal  layer  of  the  amphithecium,  but  with  these  ex- 
ceptions it  arises  from  the  endothecium,  usually  from  its  most 
external  layer.  In  the  true  Mosses  and  in  Eiccia  only  spore- 
mother-cells  are  produced  from  the  archesporium,  but  in  the 

1  The  oospore  divides  by  a  wall  transverse  or  oblique  to  the  longer  axis  of 
the  archegonium.  From  the  upper  (epibasal)  cell,  tbe  capsule  (and  seta)  is 
derived,  while  the  lower  (hypobasal)  gives  rise  to  the  foot.  In  Iticcia  the  hypo- 
basal  half  takes  part  in  the  formation  of  the  sporangium. 


187 


majority  of  the  Liverworts  some  of  these  cells  are  sterile  and 
become  elaters  (cells  with  spirally  thickened  walls,  Figs.  196,  189), 
or  serve  as  "  nurse-cells  "  for  the  spore-mother-cells,  which  gradu- 
ally absorb  the  nutriment  which  has  been  accumulated  in  them. 
In  Anthoceros,  and  almost  all  the  Leafy-Mosses,  a  certain  mass  of 
cells  in  the  centre  of  the  sporangium  (derived  from  the  endo- 
thecium)  does  not  take 
part  in  the  formation 
of  the  archesporium, 
but  forms  the  so  called 
"column"  or  "  colum- 
ella"  (Figs.  190,  191). 
The  spores  arise  in 
tetrads,  i.e.  four  in  each 
mother-cell,  and  are 
arranged  at  the  corners 
of  a  tetrahedron,  each 
tetrahedron  assuming 
the  form  of  a  sphere  or 
a  triangular  pyramid. 
The  mature  spore  is  a 
nucleated  mass  of  pro- 
toplasm, with  starch  or 
oil  as  reserve  material. 
The  wall  is  divided  into 
two  layers:  the  exter- 
nal coat  (exospore) 
which  is  cuticularized 
and  in  most  cases 
coloured  (brown,  yel- 
lowish), and  the  internal 
coat  (endospore) ,  which 
is  colourless  and  not 
cuticularized.  On  ger- 
mination the  exospore 
is  thrown  off,  the  endo- 
spore protrudes,  and 
cell-division  commences 
and  continues  with  the  growth  of  the  protonema  (Fig.  186,  B-D).. 

The  morphological  explanation  which  Celakovsky  has  given  of  the  sporo- 
gonium,  and  which  is  not  at  all  improbable,  is,  that  it  is  homologous  with* 


FIG.  193. — Andrecea  petro- 
pMla.  An  empty  capsule  ; 
the  calyptrahas  fallen  off. 

(Mag.  25  times.) 
/ 

FIG.  192.— Andrecea  petrophila.  A  ripe  sporogonium  : 
a  an  archegonium  which  has  been  raised  with  the 
pseudopodium ;  p  the  foot ;  6  the  neck  ;  d-e  the  dark- 
coloured  portion  of  the  sporangium,  whose  outer  cell- 
walls  are  considerably  thickened ;  c-c  the  thin-walled 
portions  where  the  dehiscence  occurs  ;  o  the  lower  ex- 
tremity of  the  spore-sac;  /calyptra;  g  the  apex  of 
the  sporangium.  (Mag.  25  times.) 


188  MUSCINEJl. 

an  embryo  consisting  of  a  very  small  stem-portion  and  a  terminal  spore-pro- 
ducing leaf.  This  will  be  further  explained  in  the  introduction  to  the  Flower- 
ing-plants (p.  236). 

In  the  Liverworts  the  young  sporogonium  lives  like  a  parasite, 
being  nourished  bj  the  sexual  generation  (only  in  Anthoceros  has  it 
a  slight  power  of  assimilation).  In  the  Leafy-Mosses,  on  the  other 
hand,  with  regard  to  the  power  of  assimilation,  all  transitions 
are  found  from  abundant  assimilation  (Funaria,  Physcomitrium) 
to  almost  complete  "  parasitism  "  (Sphagnum,  Andrecea).  In  the 
majority  of  the  operculate  Mosses  the  sporogonium  has  a  more 
or  less  perfect  system  of  assimilation,  and  is  able  itself  to  form  a 
large  portion  of  the  material  necessary  for  the  development  of  the 
spores,  so  that  it  chiefly  receives  from  the  sexual  generation  the 
inorganic  substances  which  must  be  obtained  from  the  soil.  The 
more  highly  developed  the  assimilative  system  of  the  sporogonium, 
the  more  stomata  are  present. 

APOSPORY.  In  some  operculate  Mosses  it  has  been  possible  to  obtain  a  pro- 
tonema  with  small  Moss-plants  from  the  seta,  when  severed  from  its  Moss- 
plant,  and  grown  on  damp  sand. 

The  Mosses  are  the  lowest  plants  which  are  provided  with  stem 
and  leaf.  They  are  assigned  a  lower  place  when  compared  with 
the  higher  Cryptogams,  partly  because  there  are  still  found  within 
the  Division  so  many  forms  with  a  mere  thallus,  partly  because 
typical  roots  are  wanting  and  the  anatomical  structure  is  so  ex- 
tremely simple,  and  partly  also  because  of  the  relation  between 
the  two  generations.  The  highest  Mosses  terminate  the  Division, 
the  Muscinege  and  Pteridophyta  having  had  a  common  origin  in 
the  Algae-like  Thallophyta. 

They  are  divided  into  two  classes  : — 

HEPATICJ:,  or  Liverworts. 

MUSCT  FRONDOSI.     True  Mosses  or  Leafy-Mosses. 

Class  \.    Hepaticae  (Liverworts). 

The  protonema  is  only  slightly  developed.  The  remaining  part 
of  the  vegetative  body  is  either  a  prostrate,  often  dichotomously- 
branched  thallus,  pressed  to  the  substratum  (thalloid  Liverworts), 
with  or  without  scales  on  the  under  side  (Figs.  194,  197)  ;  or  a 
thin,  prostrate,  creeping  stem,  with  distinctly-developed  leaves, 
which  are  borne  in  two  or  three  rows  (Figs.  195,  198),  viz.,  two  on 
the  upper  and,  in  most  cases,  one  on  the  under  side.  The  leaves 
situated  on  the  ventral  side  (amphigastria)  are  differently  shaped 
from  the  others  (Fig.  198  a),  and  are  sometimes  entirely  absent. 


MUSOINE^K. 


189 


In  contradistinction  to  the  Leafy-Mosses,  stress  must  be  laid  on 
the  well-marked  dorsiventrality  of  the  vegetative  organs  ;  i.e.  the 
very  distinct  contrast  between  the  dorsal  side  exposed  to  the  light 
and  the  ventral  side  turned  to  the  ground.  Veins  are  never  found 
in  the  leaves. 

The  ventral  part  of  the  archegonium  (calyptra)  continues  to  grow 
for  some  time,  and  encloses  the  growing  ambryo,  but  when  the 
spores  are  ripe  it  is  finally  ruptured  by  the  sporangium,  and  re- 
mains situated  like  a  sheath  (vagimtla)  around  its  base.  The 
sporangium  opens,  longitudinally,  by  valves  or  teeth  (Fig.  194,  195, 
197  6),  very  rarely  by  a  lid,  or  sometimes  not  at  all.  A  columella 
is  wanting  (except  in  Anthoceros,  Fig.  194)  ;  but  on  the  other  hand, 
a  few  of  the  cells  lying  between  the  spores  are  developed  into 


FIG.  194.— Anthtceros   Imvit 
(nat.  size) :  K-K  capsules. 


FIG.  196.  — An 
elater  with  two 
spores. 


FIG.  195.— Plagiochila  asplenioides :  a  unripe,  atid  b  an  open  capsule;  p  involucre.  The 
ventral  edge  of  each  leaf  is  higher  than  its  dorsal  edge,  and  covered  by  the  dorsal  edge  of 
the  next  one. 

elaters  (Fig.  196),  i.e.  spindle-shaped  cells  with  spirally-twisted 
thickenings,  which  are  hygroscopic,  and  thus  serve  to  distribute 
the  spores.  (They  are  seen  in  Fig.  189  (7,  not  yet  fully  developed, 
as  long  cells  radiating  from  the  base  of  the  sporangium.  They 
are  wanting  in  Riccia). 

Round  the  entire  archegonium,  (or  group  of  archegonia,  when  several  are 
developed  on  the  same  receptacle)  a  sheath — the  involucre — is  often  formed, 
which  persists,  and  encloses  the  base  of  the  stalk  of  the  sporangium,  together 
with  the  sheath  of  the  archegonium  (Fig.  195^).  In  the  MarchantiaceaB  each 
archegonium  is  enclosed  in  a  loose  investment,  the  perigynium,  which  is 
developed  as  an  outgrowth  from  the  cells  of  its  stalk. 


190 


The  majority  of  the  Liverworts  are  found  in  damp  and  shady 
places,  pressed  to  the  substratum  ;  a  few  are  found  floating  in 
fresh  water. 

Family  1.     Marchantiese. 

This  embraces  only  forms  with  a  thallus,  which  is  more  or  less 
distinctly  dichotomously  branched,  in  some,  one  or  two  rows  of 
thin  leaves  are  situated  on  its  under  surface.  On  the  upper 
surface  of  the  thallus  are  found  large  air-chambers. 

Order  1.  Ricciaceae.  The  sporogonia  are,  with  the  excep- 
tion of  a  few  genera,  situated  singly  on  the  surface  of  the  thallus, 
and  consist  only  of  a  capsule  without  foot  or  stalk.  They  always 
remain  enclosed  by  the  wall  of  the  archegonium  (calyptra),  and  open 


FIG.  197. — Marcliantia  polymorplia.  A  Female  plant  (nat.  size)  :  a  and  b  are  arche- 
goniophores  in  various  stages  of  development;  s  cupules  with  gemrnas  (see  page  183). 
B  An  archegoniophore  seen  from  below,  the  short-stalked  sporangia  are  seen  placed  in 
8-10  double  rows.  C  Male  plant,  with  a  young  and  an  older  autheridiophore.  D 
Antheridiophore  halved  vertically  to  show  the  antheridia  (7i) ;  m  the  aperture  of  the 
pits  in  which  they  are  sunk— the  older  ones  to  the  left,  the  younger  to  the  right. 

only  by  its  dissolution.  Elaters  are  not  developed.  Some  genera 
are  found  floating  like  Duckweed. — Riccia  glauca  grows  on  damp 
clay  soil.  R,.  fluitans  and  JR.  natans  float  in  stagnant  waters. 

Order  2.  Corsiniaceae.  (Not  native).  Intermediate  forms  between  the 
preceding  and  the  following  order.  In  internal  and  external  structure  mainly 
resembling  the  Marchantiaceae.  Corsinia  ;  Boschia. 

Order   3.      Marchantiaceae,    are    large,    fleshy    forms.      The 


MUS01NBJB. 


191 


surface  of  the  thallus  is  divided  into  small  rhombic  areas,  in  the 
centre  of  each  of  which  is  found  a  large,  peculiarly  constructed 
stoma  (Fig.  197  A)  ;  beneath  each  of  these  a  large  air-cavity  is  to 
be  found.  From  the  floor  of  the  air-cavity  a  number  of  alga- 
like  cells  project  into  it;  these  contain  chlorophyll  and  are 
therefore  the  assimilating  cells.  The  antheridia  and  archegonia 
are  each  found  aggregated  on  specially  formed  branches  (some- 
what resembling  Mushrooms)  projecting  from  the  surface  of  the 
thallus.  The  antheridia  are  developed  on  the  upper  surface  (Fig. 
197  C,  D)  and  the  archegonia  on  the  lower  (Fig.  197  A,  B),  near 
the  centrally-placed  stalk. 

Marchanticu  polymorpha  is  dioecious  (Fig.  197),  and  very  common 
on  damp  places.  Lunularia  (South  Europe),  frequently  found 
on  flower-pots  in  conservatories ;  Preissia,  Fegatella,  Rebotilia, 
Targionia. 

Family  2.     Anthoceroteae. 

These  have  an  entirely  leafless,  fleshy,  flat,  and  irregularly-shaped  thallus. 
In  its  intercellular  chambers  Nostoc-colonies  are  often  found,  which  have  forced 
their  way  through  the  stomata  situated  on  the  under  side.  The  antheridia  and 
archegdhia  arise  from  the  cells  lying  inside  the  thallus.  The  capsule  resembles 
a  long,  thin  pod;  it  has  two  valves  and  a  columella.  Anthoceros  (A.  lcevist 
Fig.  194,  and  punctatus) . 

Family  3.    Jungermannieae. 

Some  forms  in  this  family  have  a  thallus  in  which  leaf -like 
structures  are  found  (Blasia),  while  in 
others  (e.g.  Metzgeria,  Pellia,  Aneura)  they 
are  entirely  absent.  The  majority,  how- 
ever, have  round,  thick  stems,  bearing 
dorsally  two  rows  of  leaves,  and  one  row 
ventrally.  Some  of  these  have  the  leaves 
"  underlying "  (Fig.  195),  while  in  others 
(Fig.  198)  they  are  "  overlying."  (See  Figs. 
195,  198,  with  explanation). 

The  sporangia  are  spherical,  stalked,  and 
situated  singly  on  the  apex  of  the  branches, 
and  open  by  four  valves  (in  Sphcerocarpus 
they  are  indehiscent). 

All  the  species  in  this  family  were 
formerly  reckoned  as  belonging  to  one 
genus,  Jungermannia,  but  now  they  are 
divided  into  several,  arranged  as  follows  : — 


\v.  B. 


FIG.  198.  —  Frullania  dila. 
tata.  Portion  of  a  branch 
seen  from  the  under  side : 
r  and  b  are  the  anterior  and 
posterior  edges  of  the  same 
dorsal  leaf;  a  ventral  leaves 
(amphigastria).  The  dorsal 
leaves  are  "  overlying,"  i.e. 
the  anterior  edge  of  the 
leaf  overlaps  the  posterior 
edge  of  the  preceding  one. 
O 


192 

I.  ANACROGYN^;.      The  archegonia  are    situated  on   the   upper 
side  of  the  thallus  or  stem,  placed  laterally.,  and  covered  by  an 
"  involucre,"  formed  by  the  calyptra  together  with  the  tissue  of 
the  stem  or  thallus. 

a.  ANELATEREJ:.     Without  any  elaters  :    Sphcerocarpus,  Hiella. 

b.  ELATEREJ:.     a.  Thalloid  :  Aneura  pinguis,  in  damp  situations ; 
Metzgeria  furcata,  on  trees ;  Pellia  epiphylla,  in  damp  situations ; 
Blasia  pusilla,  on  damp  clay  soil,  in  the  shade  (scales  are  present 
on  the  thallus) .     ft.  Foliose  and     ot  dorsiventral :  Haplomitrium 
hookeri. 

II.  ACROGYNJ:.     The  apex  of  the  stem  or  of  certain  branches  is 
adapted  for  the  formation  of  female  shoots.     The  archegonia  are 
most  frequently  aggregated  on  the  apex  of  the  shoots,  and  are 
encircled  by  their  leaves  (perichaetium).     Between  these  and  the 
archegonia,  enclosing  the  latter,  a  peculiar  cup-shaped  organ  (the 
involucre)    is    formed.      This   group    only   includes    leaf-bearing 
genera  :    Frullania,    Radula,    Madotheca,    Ptilidium,    Calypogeia, 
Lepidozia,     Mastigobryum,    Lophocolea,    Jungermannia,     Scapania, 
Plagiochila. 

Class  2.     Musci  frondosi  or  .veri  (True  Mosses). 

In  this  class  the  protonema  is  well  developed,  and  resembles  a 
branched  filamentous  Alga,  from  which  it  can  be  easily  distin- 
guished by  its  oblique  septa  (in  Sphagnum  it  is  a  cellular  expan- 
sion). The  Moss-plant,  which  is  developed  directly  from  the 
protonema,  generally  has  an  erect,  thick,  cylindrical  stem  similarly 
constructed  on  all  sides.  The  leaves  are  arranged  spirally,  the 
most  frequent  divergence  being  |-  or  ^  (Fig.  200  -4).  A  midrib  is 
often  present  and  also  marginal  veins  formed  by  longitudinally 
elongated  cells ;  at  these  veins  the  leaf  is  more  than  one  layer  in 
thickness.  In  Leucobryum  the  leaves  are  generally  constructed 
of  more  than  one  layer. 

The  stem  grows  by  means  of  a  three-sided,  pyramidal,  apical  cell 
which  gives  rise  to  three  rows  of  segments,  each  segment  forming 
a  leaf.  The  lateral  branches  arise  from  the  lower  portions  of  the 
segments,  the  upper  portion  of  which  does  not  take  any  part  in  the 
construction  of  the  leaf.  From  their  mode  of  origin  the  branches 
are  not  axillary,  and  differ  in  this  respect  from  the  Flowering- 
plants. 

The  ventral  portion  of  the  archegonium  is  very  early  ruptured 


MUSCINEJJ.  193 

at  its  base  by  the  growing  sporogonium,  upon  which  it  remains,  and 
it  is  thus  raised  into  the  air,  forming  a  "  hood,"  the  calyptra 
'(Figs.  192  ;  200  J5).  In  the  Sphagnaceae  the  hood  is  not  present ; 
in  this  order,  as  in  the  Liverworts,  the  archegonium  remains  at  the 
base  of  the  sporogonium.  The  sporangium  opens  by  cireumsessile 
dehiscence  ;  the  upper  portion  (operculum)  being  separated  along 
a  specially  constructed  ring  of  cells,  and  falls  off  like  a  u  lid  " 
(Fig.  200).  Only  in  a  few  forms  (families  2  and  3)  does  any  vari- 
ation of  this  take  place.  Elaters  are  never  found,  but  (with  the 
•exception  of  Archidium)  there  is  always  present  in  the  sporangium 
a,  central  mass  of  cells,  the  columella,  which  take  no  part  in  the 
formation  of  the  spores.  The  columella,  in  some,  does  not  reach 
quite  to  the  operculum  and  in  these  cases  the  spore-sac  is  bell- 
-shaped  and  covers  the  columella  (Andrecea,  Fig.  190 ;  Sphagnum, 
iFig.  199  D)  ;  but  in  the  majority  of  Mosses  the  columella  extends 
io  the  lid,  so  that  the  space  containing  the  spores  becomes  a 
ihollow  cylinder. 

The  sporangium  is  generally  raised  on  a  long  stalk ;  in  the  great 
•majority  this  stalk  is  formed  from  the  lower  half  of  the  oospore 
and  belongs  to  the  asexual  generation — it  is  then  known  as  the 
>seta.  In  Andrecea  and  Sphagnum  the  seta  is  very  short,  and  the 
sporangia  are  raised  upon  a  long  stalk  (pseudopodium)  developed 
from  -the  summit  of  the  sexual  generation  (Figs.  190,  192).  In 
the  latter  figure  an  archegouium  (a)  is  seen  attached  to  the 
pseudopodium,  having  been  carried  up  with  this  during  the  course 
of  its  development.  The  summit  of  the  pseudopodium  is  enlarged 
'to  embrace  the  foot  of  the  sporogonium  (Figs.  192,  199  D) 

A.  The  sporangium  is  supported  on  a  pseudopodium  ;  the  columella  does 
,not  extend  to  the  operculum. 

Family  1.     Sphagneae  (Bog- Mosses). 

'The  protonema  has  been  already  described.  The  stem  is 
•regularly  branched  owing  to  the  fact  that  a  branch,  or  collection  of 
branches,  arises  at  every  fourth  leaf.  These  branches  are  closely 
covered  with  leaves,  some  are  erect,  while  others  hang  down  and 
surround  the  stem.  No  rhizoids  are  developed.  These  Mosses  are 
of  a  whitish-green  colour,  and  when  water  is  present  are  always 
saturated  with  it  like  a  sponge,  the  reason  for  this  being  found  in 
the  construction  of  the  stem  and  leaves.  The  stems  are  covered 
by  an  external  layer  of  large  clear  cells,  without  chlorophyll,  but 
<with  annular  or  spiral  thickenings  on  the  walls,  which  are  also 


194 


MUSCINE^E. 


perforated  by  large  holes.  By  means  of  capillary  attraction,  water 
is  thus  raised  to  the  summit  of  the  stem.  Similarly  constructed 
cells  are  also  found  in  the  leaves,  but  they  are  surrounded  by  a  net 
of  very  narrow,  chlorophyll-containing  cells  (Fig.  199  F),  whose 
colour  is  thus  to  a  great  extent  lost  amongst  those  which  are 


FIG.  199.— Sphagnum  acutifolium.—A  The  upper  portion  of  a  plant :  a  branches  with 
antheridia ;  cli  branches  -with  terminal  archegonia  and  perichaetia;  b  the  upper  stem- 
leaves.  B  A  male  branch  whose  leaves  are  partly  taken  off  in  order  to  show  the 
antheridia.  C  Group  of  three  archegonia :  the  central  one  (a)  is  formed  from  the  apical 
cell.  D  Sporogonium  in  longitudinal  section :  the  broad  foot  (sg')  is  sunk  in  the  vaginula, 
v  ;  c  calyptra;  ar  neck  of  the  archegonium  ;  ps  pseudopodium.  E  ripe  sporangium  with 
operculum,  and  the  remains  of  the  archegonium  situated  on  the  pseudopodium  which  is 
still  surrounded  by  the  perichsetium ;  to  the  left  is  a  barren  branch.  F  Portion  of  a 
f oliage-lpaf  seen  from  above :  I  perforations ;  b  chlorophyll-containing  cells ;  s  spiral 
thickenings. 

colourless.     This  anatomical  structure  is  an  essential  condition  for 
the  formation  of  peat.     The  Bog-Mosses  grow  by  preference  on 


MUSCINEJ3.  195 

moors,  which  they  cover  with  a  thick  carpet  saturated  with  water. 
The  lower  extremities  of  the  plants  perish  very  rapidly,  and 
gradually  become  converted  into  peat,  and  the  branches  thus 
separated  from  each  other  become  independent  plants.  The 
sporangia  (Fig.  199  D,  E)  are  spherical,  but  with  a  very  short 
stalk.  They  open  by  a  lid,  but  have  110  annulus.  The  archegonium 
(Fig.  199  (7)  persists  at  the  base  of  the  sporogonium  as  in  the 
Liverworts.  Only  one  genus,  Sphagnum. 

Family  2.     Schizocarpeae. 

The  Mosses  which  constitute  this  family  are  of  a  brownish -black  colour  and 
are  found  living  on  rocks.  The  sporangium  resembles  that  of  the  Liverworts 
inasmuch  as  it  opens  by  four  valves,  but  these  continue  attached  to  each  other  at 
the  apex  as  well  as  at  the  base  (Fig.  193).— There  is  only  one  genus :  Andreaa. 

B.  The  stalk  is  formed  from  the  lower  portion  of  the  sporogonium.  The 
columella  is  continued  to  the  summit  of  the  sporangium  and  united  with  it 
(Arclddium  has  no  columella.) 

Family  3.     Cleistocarpeae. 

The  fruit  does  not  dehisce  in  the  regular  way,  but  the  spores  are  liberated  by 
decay.  They  are  small  Mosses  which  remain  in  connection  with  their  protonema 
until  the  sporangium  is  mature.  The  archegonium  remains  sessile  at  the  base 
of  the  short  capsule-stalk,  and  is  not  raised  into  the  air  (compare  Hepaticae). — 
Phascum,  Ephemerum,  Archidium,  Pleuridium. 

Family  4.     Stegocarpeae. 

To  this  belong  the  majority  of  the  Mosses,  about  3,000  species. 

The  capsule  opens  as  in  Sphagnum  by  means  of  a  lid  (operculum) , 
which  is  often  prolonged  into  a  beak.  Round  the  mouth  of  the 
opened  capsule,  a  number  of  peculiar  yellow  or  red  teeth  are  to  be 
found.  These  constitute  the  peristome  ;  their  number  is  four,  or  a 
multiple  of  four  (8,  16,  32  or  64).  The  form  and  thickenings 
of  these  teeth  are  widely  different,  and  on  this  account  are  used  by 
Systematists  for  the  purposes  of  classification.  In  some  Mosses 
(Fig.  200  0,  D)  there  is  a  double  row  of  teeth.  Except  in 
Tetraphis  they  are  not  formed  from  entire  cells,  but  from  the 
strongly  thickened  portions  of  the  wall  of  certain  layers  of  cells 
belonging  to  the  lid,  and  persist  when  this  falls  off.  They  are 
strongly  hygroscopic,  and  assist  greatly  in  the  ejection  of  the  lid, 
in  which  operation  they  are  considerably  aided  by  a  ring  of  elastic 
•cells  with  thickened  walls,  situated  in  the  wall  of  the  lid  near  the 
base  of  the  teeth.  This  ring  is  known  as  the  annulus.  The  arche- 
gonium. is  raised  into  the  air  like  a  hood,  the  calyptra,  which 


196 


MUSCINE2E. 


either  covers  the  sporangium  on  all  sides  (having  the  shape  of  a 
bell),  or  is  split  on  one  side  (Fig.  200  J3,  Ji). 

Among  peculiar  forms  may  be  mentioned :  Splachnum,  which  is  especially 
remarkable  for  the  collar-like  expansion  at  the  base  of  the  capsule.     Fissidens 

deviates  in  having  a  flat  stem  and  leaves 
arranged  in  two  rows.  The  leaves  are 
boat-shaped  and  half  embrace  the  stem. 
— Schistostega  has  two  kinds  of  stems. 
The  barren  ones  resemble  Fern-leaves  ; 
they  have  two  rows  of  leaves,  which  are 
attached  together  vertically,  are  decur- 
rent,  and  coalesce  at  their  bases.  The 
fertile  ones  have  an  ordinary  appearance. 
— Tetraphis  :  the  peristome  is  composed 
of  four  teeth,  which  are  formed  from  entire 
cells.  T.  pellucida  has  peculiar  gemmae. 

The  family  is  divided  into  two 
groups  :  the  Musci  acrocarpi,  the 
growth  of  whose  main  axis  is 
limited  and  terminated  by  the 
formation  of  the  sexual  organs ; 
and  the  Musci  pleurocarpi,  whose 
sporogonia  are  situated  on  special 
lateral  shoots,  while  the  growth  of 
the  main  axis  is  unlimited. 


FIG.  200.  —  A  Hypnum  populeum.  S 
and  C  Sporangia,  with  hood  (7i),  and 
operculum  (I'),  and  without  these  (C), 
showing  the  peristome  (p).  D  The 
mouth  of  the  capsule  of  Fontinalis  anti- 
pyretic (. 


A.   Acrocarpi. 

Order  1.  Weisiaceae.  Peristome, 
with  16  teeth  arranged  in  one  series,  rarely 
wanting.  Leaf  with  midrib.  Campylopus, 
Dicranella,  Cynodontium. — 


Dicranum   (D.  scoparium,   common    in   forests), 
Weisia,  Gymnostomum  (no  peristome),  Systegium. 

Order  2.  Leucobryaceae.  Peristome  with  16  teeth.  Leaves  with  three  or 
more  layers  of  cells,  of  which  the  external  ones  are  air-conducting  and  per- 
forated (as  in  the  Sphagneaa),  the  middle  one  containing  chlorophyll.  Lenco- 
bryum, 

Order  3.  Fissidentaceae.  Peristome  as  in  the  preceding  ones.  The 
leaves  are  arranged  in  two  rows  on  the  plagiotropic  shoots ;  in  Fissidens  the: 
midrib  of  the  leaf  bears  wing-shaped  outgrowths.  Conomitrium,  Fissidens. 

Order  4.  Seligeriaceae.  Peristome  with  16  undivided  teeth.  Very  smalll 
Eock-mosses.  Seligcria. — Blindia. 

Order  5.  Pottiaceaa.  Peristome  with  16  teeth,  which  are  divided  almost 
to  the  base,  or  with  32  teeth.  Calyptra  hood-like. — Barbula  (B.  imtralis,  B. 
ruralis}.  Trichostomum,  leptotrichum.—  Ceratodon  purpureus. — Distichium. — 
Pottia. 


MUSCINEJi.  197 

Order  6.  Grimmiaceae.  The  leaf-cells  are  often  papillose ;  in  the  upper 
portion  of  the  leaf,  small,  and  of  roundish  shape.  The  calyptra  is  most 
frequently  hood-like  or  conical.  Eucalypta. — Orthotrichum,  often  with  short- 
stalked  capsule,  is  found  on  trees. — Coscinodon.—Hedwigia. — Grimmia,  Racond- 
trium. — Cinclidotus. 

Order  7.  Schistostegacese.  The  stems  are  of  two  kinds  (see  above)  ; 
Schistostega  osmundacea,  in  caves,  has  a  bright  emerald  protonema. 

Order  8.  Splachnaceae.  The  capsule  has  a  large,  collar-like  neck  (see 
above).  Splachnum  (especially  on  manure). 

Order  9.  Funariaceae.  Capsule  pear-shaped.  Funaria  (F.  hygrometrica  has 
a  very  hygroscopic  seta,  becoming  twisted  when  dry,  and  straightening  with 
moisture) ;  Physcomiirium  ;  Discelium. 

Order  10.  Bryaceae.  The  capsule  is  thicker  towards  the  apex ;  most 
frequently  pendulous.  Philonotis,  Bartramia. — Aulacomnium. — Paludella, 
Mecsea. — Mnium. — Bnjum,  Webera,  Leptobryum. 

Order  11.  Polytrichaceae.  Single  peristome,  formed  by  16,  32,  or  64  teeth. 
Leaves  with  longitudinal  lamellae  on  upper  surface. — Polytriclium  has  long, 
hairy  calyptra.  Catharinea  (C.  undulata,  in  forests). 

Order  12.  Georgiaceae.  Peristome  with  4  teeth  (see  above).  Tetraphis 
(T.  pellucida  has  gemmae  1. 

Order  13.  Buxbaumiaceae.  Capsule  asymmetrical ;  double  peristome  : 
the  interior  one  conical,  with  16  or  32  longitudinal  folds. — Buxbaumia  (B. 
aphylla) ;  Diphyscium. 

B.    Pleurocarpi. 

Order  14.  Fontinalaceae.  Long,  floating  Water-Mosses.  Fontinalis  (F. 
antipyretica  is  found  in  streams).  Dichelyma. 

Order  15.     Hookeriaceae.     Pterygophyllum. 

Order  16.  Leskeaceae.  Dull-looking  Mosses,  with  papillose  or  warted 
leaves. — Thuidium,  Thuja-like  with  regularly  arranged  1-3  doubly  pinnate 
stems ;  Anomodon,  Leskea. 

Order  17.     Pterogoniaceae.     Pterigynandrum  filiforme,  etc. 

Order  18.     Fabroniaceae.     Anacamptodon. 

Order  19.  Neckeraceae.  Steins  most  frequently  with  flat,  leafy  branches. 
The  leaves  are  smooth,  never  with  longitudinal  folds. — Neckera. 

Order  20.  Hypnaceae.  The  leaves  are  smooth  with  square,  often  bladder- 
like,  cells  at  the  edge.  Hylocomium  (H.  splendens,  H.  triquetrum) ;  Hypnum  ; 
Brachythecium  ;  Plagiothecium. — Eurhynchium. — Homalothecium,  Isothecium, 
Orthothecium,  Homalia. — Climacium,  Lescurcea,  Leucodon. 

The  Mosses  occur  all  over  the  globe.  Many  are  found  in  great  numbers,  and 
growing  thickly  massed  together,  they  form  an  important  feature  in  landscapes 
(for  example  Sphagnum  and  Polytrichum  in  the  Arctic  Tundra).  In  the  Northern 
and  Arctic  regions  the  Mosses  are  very  plentiful,  and  often  form  a  considerable 
part  of  the  vegetation,  while  in  the  Tropics  they  are  insignificant. 

Species  of  Hypnum  and  Polytrichum,  like  Sphagnum,  play  an  important  part 
in  the  formation  of  peat. 


DIVISION  III. 

PTERIDOPHYTA   (VASCULAR 
CRYPTOGAMS). 

The  alternation  of  generations  is  as  distinct  in  this  Division  as 
in  the  Mosses,  but  the  sexual  generation  consists  of  only  a  small 
thallus,  the  prothallium,  which  bears  directly  the  sexual  organs, 
antJieridia  and  archegonia -,  and  the  asexual  generation,  which 
arises  from  the  fertilisation  of  the  oosphere,  is  no  longer  a  single 
short-lived  sporangium,  but  a  highly  developed,  generally  per- 
ennial, plant  provided  with  stem,  leaves  and  true  roots  (Ferns, 
Horsetails,  etc.),  the  sporangia  being  borne  on  the  leaves.  In  this 
latter  generation  the  tissues  are  differentiated  into  epidermis, 
ground  tissue  and  vascular  tissue;  in  the  last  named  the  bundles 
are  closed,  and  in  the  majority  of  cases  concentric. 

The  sexual  generation,  gametophyte,  or  prothallium,  is 
always  a  thallus,  although  not  always  green  and  leaf -like  (Figs. 
'205,  215,  222,  229,  235,  etc.)  It  is  very  small,  even  in  cases  where 
it  attains  the  greatest  development,  and  consists  only  of  paren- 
chymatous  cells.  The  prothallium  is  nourished  by  hair-like  roots 
(rhizoids)  and  has  only  a  transitory  existence,  dying  soon  after  the 
fertilisation  of  its  oosphere. 

The  ANTHERIDIA  exhibit  great  variations  in  structure  which, 
however,  must  be  considered  as  modifications  of  the  fundamental 
type  which  is  found  in  the  Mosses.  These  modifications  will  be 
mentioned  under  the  various  families.  The  spermatozoids  are 
always  spirally-coiled,  self-motile,  protoplasmic  bodies,  with  most 
frequently  a  large  number  of  fine  cilia  on  the  anterior  end  (Figs. 
206, 223,  234).  They  are  formed  principally  from  the  nucleus  of  the 
mother-cell,  and  portions  of  the  cytoplasm  often  remain  for  a 
time  attached  to  their  posterior  end. 

The  ARCHEGONIA  are  more  uniform  throughout  the  entire 
Division,  and  more  closely  resemble  those  of  the  Mosses.  They 
are,  as  in  the  previous  Division,  principally  flask-shaped ;  but  the 


PTERIDOPHYTA.  199 

central  portion,  which  encloses  the  oosphere,  is  always  embedded 
in  the  tissue  of  the  prothallium,  so  that  the  neck,  which  is  formed 
of  4  rows  of  cells,  projects  above  the  surface  (Figs.  201  3,  222  h). 
The  development  of  the  archegonium  in  a  Fern  is  seen  in  the 
accompanying  figure  (Fig.  201).  The  archegoniumis  developed  from 
a  surface  cell,  which  divides  into  three  cells  by  two  walls  in 
a  direction  parallel  to  the  surface  of  the  prothallium  (Fig.  201). 
The  most  internal  cell  becomes  the  ventral  portion  of  the  arche- 
gonium. The  external  one  (6)  divides  perpendicularly  to  the  sur- 
face of  the  prothallium  into  four  cells,  which  again  divide  parallel 
to  the  surface  and  form  the  neck  (6,  in  2  and  3).  The  interme- 
diate cell  projects  upwards  into  the  neck  and  divides  into  two, 


FIG.  201.— Pleris  serrulata.     Development  of  archegonia. 


the  lower  one,  after  the  separation  of  the  ventral  canal-cell,  becom- 
ing the  oosphere,  arid  the  upper  one  the  neck-canal-cell  (c,  in  2  and  3). 
.  As  in  the  Mosses,  the  divisional  walls  of  the  neck-canal-cells 
become  mucilaginous,  causing  the  rupture  of  the  neck  of  the 
archegonium.  Fertilisation  takes  place  as  in  the  Mosses,  and  the 
passage  of  the  spermatozoids,  along  the  neck,  to  the  oosphere, 
has  been  observed.  Water  (rain  or  dew)  is  similarly  necessary  for 
the  movements  of  the  spermatozoids,  and  hence  for  fertilisation. 
The  other  classes  of  the  Division  chiefly  deviate  from  the  Ferns 
in  having  the  archegonium  sunk  deeper  into  the  prothallium,  and 
the  neck  reduced  in  length  (compare  Fig.  201  with  Figs.  216,  222, 
235,  236). 


200  PTERIDOPHYTA. 

According  to  the  nature  of  the  spores,  the  three  classes  of 
the  Vascular  Cryptogams  are  each  divided  into  isosporous  and 
heterosporous  groups. 

I.  The  isosporous  Vascular  Cryptogams  have  only   one   kind 
of  spore.     The  prothallium  developed  from  this  is  in  some  cases 
monoecious,  bearing  both  antheridia  and  archegonia ;  but  in  others 
there  is  a  distinct  tendency  for  each   prothallium  to  bear  only 
antheridia  or  archegonia  (dioecious) — true  Ferns  and  Lycopodium. 

In  Equisetum  there  is  only  one  kind  of  spore,  but  two  kinds  of 
p.rothallia  are  developed,  one  of  which  bears  only  antheridia. 
(male),  the  other  only  archegonia  (female)  ;  but  the  one  that 
bears  antheridia  may  be  transformed  into  the  one  that  bears 
archegonia  and  vice  versa. 

II.  In  the  higher  group,  heterosporous  Vascular  Cryptogams 
(Selaginella   and    Isoetes,    etc.),    there    are   two  distinct  kinds  of 
spores,   the  small,  microspores,  and  the  large,  macrospores.     The 
microspores    are  male,    and    produce    prothallia  which  bear  only 
antheridia.     The  macrospores  are  female,   and  produce   prothallia 
which  bear  only  archegonia. 

Corresponding  to  this  difference  in  the  spores,  there  is  also 
found  a  difference  in  the  development  of  the  prothallium.  In  the 
Isosporeae  the  prothallium  is  large,  and  either  green,  leaf-like,  and 
provided  with  rhizoids  (most  of  the  Ferns,  Horsetails,  etc.),  or 
subterranean,  pale-coloured,  and  globular  (Ophioglossum,  Lycopo- 
dium). It  lives  vegetatively  for  a  fairly  long  time,  and  generally 
produces  a  large  and  varying  number  of  archegonia  and  an- 
theridia. The  prothallium  in  the  Heterosporeas  is  gradually  more 
and  more  reduced,  its  independent  and  vegetative  life  becomes  of 
less  and  less  importance,  it  becomes  more  dependent  on  the  mother- 
plant,  and  projects  from  the  spore  very  slightly,  or  not  at  all. 
The  antheridia  and  archegonia  become  reduced  in  number  to  one, 
and  also  degenerate  in  point  of  development. 

It  may  here  be  remarked  that  the  gradual  development  of  the- 
asexual  generation,  the  development  of  the  two  kinds  of  spores, 
and  the  progressive  reduction  of  the  prothallium  and  sexual 
organs  which  is  found  in  this  Division,  is  continued  to  the  Gymno- 
sperms  and  Angiosperms.  The  microspores  are  in  these  called 
pollen-grains,  and  the  male  prothallium  is  very  rudimentary. 
The  macrospores  arc  termed  embryo-sacs,  and  the  female  prothal- 
lium, the  endosperm. 

The  asexual  generation,  sporophyte.     When  the  oospherer 


PTERIDOPHYTA. 


201 


which  in  this  cas"e  as  in  all  others  is  a  primordial  cell,  is  fer- 
tilised, it  surrounds  itself  with  a  cell-wall  and  commences  to  divide 
into  a  number  of  cells,  to  form  the  embryo. 

The  first  dividing  wall  (basal  wall)  is  nearly  horizontal,  and  in  the  direction  of 
the  longitudinal  axis  of  the  archegonium.  The  next  wall  is  vertical,  and  the 
next  perpendicular  to  the  other  two.  The  oosphere,  therefore,  is  now  divided 
into  eight  octants  by  these  three  walls.  The  basal  wall  divides  the  embryo  into 
a  hypobasal  and  an  epibasal  half.  From  the  first  one,  by  continued  divisions, 
the  first  root  is  developed  ;  from  the  latter,  the  stem  and  leaves.  After  the  for- 
mation of  the  octants  the  development  proceeds  in  somewhat  different  ways  in 
the  various  classes.  In  addition  to  the  stem,  leaf,  and  root,  a  "  foot "  is  deve- 
loped from  the  hypobasal  half  which  remains  enclosed  in  the  prothallium,  and 
conveys  nourishment  from  the  prothallium  to  the  young  plant  until  it  is  able 
to  sustain  itself  (Fig.  202).  The  formation  of  these  members  in  the  embryo 
depends  on  the  position  of  the  oosphere  in  the  archegonium  and  prothallium, 
and  is  independent  of  gravity. 


FIG.  202.—  Adiantum  capillus  veneris.  Vertical  section  through  a  prothallium  (//), 
with  a  young  plant  attached  on  its  under  side  (mag.  about  10  time>) ;  r  the  first  root,  and 
b  the  first  leaf  of  the  young  Fern-plant ;  m  the  foot.  In  the  angle  between  m  and  b  lies  tLe 
apex  of  the  stem  :  h  the  rhizoids  of  the  prothallium ;  ce  ce  unfertilised  archegonia. 

In  the  Mosses  the  asexual  generation  is  the  sporogonium,  which 
is  limited  in  its  development  and  in  a  great  measure  dependent 
upon  the  sexual  generation,  upon  which  it  is  situated  ;  but  in  the- 
Pteridophyta  this  generation  is  an  independent  and  highly  developed 
plant,  provided  with  stem,  leaf,  and  true  roots,  and  has  in  many 
instances  an  unlimited  development.  The  Pteridophyta  are  the 
lowest  Division  with  true  roots.  The  root  which  is  first  formed  ia 
very  similar  in  nature  to  the  primary  root  of  the  Monocotyledons ; 
it  very  soon  dies  and  is  replaced  by  others  which  are  more  per- 
manent, and  developed  upon  the  stem  (adventitious  roots)  ;  roots 
are  wanting  in  Salvinia,  Psilotum,  and  some  Hymenophyllacea3. 
The  differentiation  is,  however,  not  so  complete  as  in  the  Flower- 
ing-plants, and  so  many  leafy  forms  are  not  found.  The  various 
members  of  these  plants  are  anatomically  much  higher  than  in  the- 


202 


PTERIDOPHYTA. 


Mosses,  having  an  epidermis,  a  ground  tissue  with  variously  dif- 
ferentiated cells,  and  a  highly  developed  vascular  system.  The 
vascular  bundles,  like  those  in  the  Monocotyledons,  are  without 
cambium,  and  closed ;  they  are  therefore  incapable  of  any  increase 
in  thickness.  In  general  the  bundles  are.  concentric,  with  the 
bast  round  the  wood  (Fig.  203).  The  wood  is  almost  entirely 
made  up  of  scalariform  tracheides. 

In  Isoetes  &  secondary  thickening  takes  place  by  a  cambium,  which  is  formed 
inside  the  cortex,  constructing  secondary  cortex  to  the  exterior,  and  secondary 
wood  towards  the  interior. — Botrychium  bas  also  a  thickening  growtb.  Collateral 
vascular  bundles  occur  in  Osmundacecz,  Equisetacecs,  and  the  leaves  of  many 
Pulypodiacete,  etc. 

It  is  a  point  of  special  interest,  that  the  gigantic  forms  of  Ferns, 

Equisetums,  and  Club- 
Mosses  (which  flourished 
in  earlier  geological 
periods,  when  these  classes 
attained  their  highest  de- 
velopment) possessed 
some  means  of  increasing 
in  thickness. 

The  sporangia  are  in 
all  cases  capsule-like,  and 
burst  open  when  ripe  to 
eject  the  spores.  They 
are  nearly  always  situated 
on  the  leaves  (in  Lycopo- 
diacece,  in  the  axils  of  the 
leaves,  or  above  these,  on 

FIG.  203.— Portion  of  the  stem  of  a  Fern.    Above  ,  i         \      T 

is  see-,  the  transverse  section,  with  vascular  bundles  the  stems  themselves).     In 

of  different  form  and  size.    The  rhombic  figures  on  some      f Orms      (LiEPTOSPOR- 

the  side  of  the  stem  are  leaf- scars.  ANGIAT*),     the      Sporangia 

are  developed  from  a  single  epidermal  cell ;  in  others  (EusPOR- 
ANGIAT^:),  from  a  group  of  epidermal  cells,  or  from  cells  which 
lie  beneath  the  epidermis.  In  the  first  group  a  primitive  mother- 
cell  (archesporium)  is  formed,  which  divides  commonly  into  sixteen 
special  mother-cells.  In  the  latter  group,  on  the  other  hand,  a 
number  of  primitive  spore-mother-cells  are  developed.  In  each 
sporangium  three  different  tissues  are  generally  developed  ;  an 
innermost  sporogenous  one  (s  in  Fig.  204  A),  which  arises  from 
the  archesporangium  ;  an  outermost  one,  which  forms  the  wall  (a), 


PTEEIDOPHYTA. 


203 


and  may  be  one  or,  more  rarely,  several  layers  in  thickness ;  and 
an  intermediate  one,  the  tapetum  (Fig.  204  A,  B,  6  £),  which  is  rich 
in  protoplasm,  and  whose  cells  are  dissolved  so  that  the  spores 
float  freely  in  the  fluid  thus  provided.  The  spores  arise  as  in  the 
Mosses  (in  tetrads),  by  the  cross-division  of  the  special  mother- 
cells,  and  according  to  the  manner  in  which  they  are  arranged 
in  the  mother-cell  have  either  a  tetrahedral  form,  with  a  large 
base  resembling  a  segment  of  a  ball,  or  are  oblong  (bilateral 
spores).  Their  construction  is  the  same  as  in  the  Mosses  (p. 
187). 

The  spore-formation  in  its  earliest  commencement  takes  place  in 
the  same  way  in  the  Isosporous  and  the  Heterosporou's  Vascular 


FIG.  204.— SelaglneUa  inaqualifolia.  A  A  young  sporangium,  which  may  develope  either 
into  a  macro-,  or  a  micro-sporaugium.  B  A  microsporangium. 

Cryptogams;  but  from  a  certain  point,  after  the  tetrahedral  divi- 
sion, a  difference  occurs  with  regard  to  the  macrosporangia.  All 
the  spores  formed  in  the  microsporang'ium  may  complete  their 
development ;  but  those  which  are  formed  in  the  macrosporangium 
are  generally  aborted,  with  the  exception  of  one  or  four,,  and  these 
consequently  attain  a  much  larger  size  (see  Fig.  239. — The  series  to- 
the  left  are  microsporangia ;  those  to  the  right,  macrosporangia) . 

APOGAMY.  In  some  Fern.s  (Pteris  cretica ;  Aspidium  filix  mas,  var.  cristatum; 
A.falcatum;  Todea  africana)  the  young  plant  is  not  developed  as  a  consequence 
of  fertilisation,  but  as  a  bud  from  the  protballium.  This  is  known  as  apogamy, 
or  loss  of  the  power  of  sexual  reproduction.  The  antheridia  are  generally 
more  or  less  developed ;  archegonia  are  entirely  wanting  in  Asp.  filix  mas,  var. 


204  PTERIDOPHYTA. 

cristatum.  This  variety  has  probably  only  become  apogamous  through  cultiva- 
tion. Many  specimens  of  Isoetes  lacustris,  in  a  lake  in  the  Vosges  mountains, 
produce  in  the  place  where  the  sporangia  are  usually  found,  a  vegetative  shoot 
which  grows  into  a  new  plant,  so  that  the  sexual  generation  is  wanting  in 
this  case.  Some  specimens  have  sporangia  on  some  leaves,  and  shoots  on 
others. 

Apospory,  or  the  formation  of  prothallia  instead  of  sporangia  and  spores  on 
the  leaves,  is  found  in  Athyrium  fdix  femina,  var.  clarissimum.  In  this  case 
the  development  of  the  sporangia  proceeds  only  to  a  certain  point,  and  from 
these  arrested  sporangia  the  prothallia  are  produced.  Normal  sporangia  are 
entirely  wanting  in  this  variety,  and  in  Aspidium  angulare,  var.  pulcherrimum, 
sporangia  are  completely  wanting.  Compare  the  Mosses  (page  188). 

The  Vascular  Cryptogams  are  divided  into  three  large  classes, 
in  each  of  which  a  progressive  development  can  be  traced  from  the 
isosporous  to  the  heterosporous  forms,  but  some  of  these  are  now 
only  known  as  fossils. 

Class  1.  Filicinae  (Ferns).— The  stem  is  small  in  comparison 
with  the  leaves,  and  branches  only  seldom,  and  then  by  lateral 
shoots.  The  leaves  are  scattered,  large,  often  deeply  divided,  and 
•of  various  highly  developed  forms.  The  undeveloped  leaves  are 
j-olled  up  in  the  bud,  having  what  is  termed  circinate  venation. 
The  sporangia  are  situated  on  the  edge  or  on  the  lower  side  of 
the  leaves,  those  on  which  the  sporangia  are  borne  (sporophylls) 
being  often  the  ordinary  foliage-leaves  ;  but  in  a  few  cases  the 
fertile  differ  from  the  barren  ones  (a  higher  stage  in  development). 
The  fertile  leaves  are  not  confined  to  definite  parts  of  the  shoot, 
and  do  not  limit  its  growth.  The  archesporium  is  most  frequently 
-unicellular. 

•A.  Isosporous:  Sub-Class  1.     Filices  (True  Ferns). 

B.  Heterosporous  :  Sub-Class  2.  Hydropteridese  (Water 
Ferns). 

Class  2.  Equisetinae  (Horsetails),  in  its  widest  meaning. — 
The  leaves  in  this  class  are  small  in  comparison  with  the  stem. 
They  are  arranged  in  whorls,  and  unite  to  form  a  sheath.  The 
sporangia  are  situated  on  specially  modified,  shield-like  leaves, 
which  are  closely  packed  together  and  form  a  "  cone."  The  cone 
is  borne  terminally,  and  limits  the  growth  of  the  shoot.  The 
sporangia  are  developed  from  a  large  group  of  epidermal  cells,  the 
archesporium  being  unicellular.  The  branches  are  arranged  in 
whorls,  and  develope  acropetally. 

A.  Isosporous:  Sub-Class  1.     Equisetaceae.     Existing  forms. 

B.  Heterosporous  :  Sub-Class  2,     Extinct  forms. 


PTERIDOPHYTA.  205 

Class  3.  Lycopodinae  (Club-Mosses).  —  Roots  generally 
branching  dichotomously.  The  leaves  are  scattered  or  opposite, 
and  in  proportion  to  the  stem  very  small,  undivided,  and  simple. 
They  are  scale-like  and  triangular,  tapering  from  a  broad  base  to 
a  point.  The  sporangia  are  situated  singly  (except  in  Psilotacece), 
and  almost  in  every  case  on  the  upper  side  of  the  leaf  or  in  the 
axil  of  a  leaf ;  but  in  some  cases  they  are  borne  on  the  stem,  just 
above  the  leaf-axil.  The  sporangia  arise  from  groups  of  epidermal 
cells.  The  sporophylls  are  often  modified,  and  differ  from  the 
foliage-leaves ;  they  are  then  arranged  in  cones  placed  terminally 
on  branches,  thus  limiting  their  growth. 

A.  Isosporous  :   Sub-Class  1.     Lycopodieae. 

B.  Heterosporous  :  Sub-Class  2.     Selaginelleas. 

Class  i.    Filicinae  (Ferns). 

The  characteristics  of  this  class  have  already  been  given  on  page 
-204. 

The  class  is  divided  into  two  sub-classes : — 

1.  The   TRUE   FERNS,  FILICES,  have  one   kind  of  spore  which 
generally  developes    monoecious   prothallia,  relatively   large   and 
green.     The  sporangia    are   most    frequently  situated   in   groups 
(sort),  which  are  often  covered  but  not  enclosed  by  an  indusium. 

2.  WATER    FERNS,    HYDROPTERIDE^E,    have   microsporangia  with 
many  (4  x  16)  microspores,  and  macrosporanyia,  each  with  one  mac- 
rospore.     The  prothallium  is  small,  and  projects  but  slightly  from 
the  germinating  spore.     The    sporangia   are    situated   in    groups 
(soH),  which  are  either  enclosed  by  an  indusium,  or  enveloped  in 
a  portion  of  a  leaf,  to  form  "  fruits  "  termed  sporocarps. 

The  old  name  for  the  Hydropteridese,  "Khizocarpeae,"  i.e.  the  "  root-fruited," 
originated  from  the  erroneous  supposition  that  the  sporocarps  were  borne  on 
the  roots. 

Sub-Class  1.     Filices  (the  True  Ferns). 

Of  the  eight  orders  (with  about  4,000  species)  comprised  in  this 
sub-class,  the  Polypodiaceee  is  the  largest  (having  about  2,800 
species)  and  the  most  familiar ;  for  this  reason  it  will  be  taken  as 
typical. 

The  sexual  generation.  When  the  spore  germinates,  the 
external  covering  (exospore)  is  ruptured,  as  in  the  Mosses.  The 
internal  cell-wall  (endospore)  grows  out  as  a  filament,  which  soon 
divides  and  gives  rise  to  the  prothallium,  a  flat,  cellular  expansion 
resembling  the  thallus  of  a  Liverwort.  In  its  fully  developed  state 


206 


PTER1DOPHYTA. 


the  prothallium  is  generally  heart-shaped,  dark  green,  and  pro- 
vided with  root-hairs,  and  it  attains  a 
diameter  of  about  one  centimetre  (Fig. 
205).  It  is  formed  of  one  layer  of  cells, 
except  along  the  central  line  near  the 
anterior  depression,  where  it  becomes 
several  layers  of  cells  in  thickness, 
forming  the  "  cushion,"  on  the  lower  side 
of  which  the  archegonia  are  developed. 
The  antheridia  are  first  formed ;  they 
are  thus  found  on  the  oldest  parts  of  the 
prothallium,  on  its  edge,  or  among  the 
root-hairs.  The  archegonia  are  developed 
later,  and  are  therefore  found  near  the 
apex.  Several  tropical  Ferns  have  pro- 
thallia1 deviating  from  this  typical  form; 
Trichomanes  (Order  HymenopJiyllacece) 
has  filamentous,  branched  prothallia, 
which  resemble  the  protonema  of  a  Moss. 

Others,  again,  have  strap-shaped  prothallia,  which  resemble  the 

thallus  of  certain  Liverworts. 


w' 


FIG.  205.— Prothallium  (pp)  of 
Maiden  hair  (Adiantum  capillus 
veneris)  with  a  young  plant 
attached :  b  first  leaf ;  w'  primary 
root ;  w"  adventitious  roots ;  h  h 
root-hairs  of  the  prothallium 
( x  abt.  30). 


FIG.  203.—  Antheridia  of  Maiden-hair  (x  550).  A  Unripe;  B  ripe,  but  unopened  ;  Copen 
and  ejecting  the  spermatozoids  (s).  Those  which  have  been  last  ejected  are  still  lying  en- 
closed in  their  mother-cells,  the  others  are  coiled  up  and  drag  with  them  the  cytoplasmic 
remains  (b) ;  /cells  of  the  prothallium. 

1  In  the  Polypodiaceas  unisexual  prothallia  as  distinct  as  those  of  Equisetum 
are  of  common  occurrence. 


PTERIDOPHYTA.  207 

The  ARCHEGONIA  have  been  already  mentioned  (p.  199,  Fig.  201). 
The  ANTHERIDIA  are  hemispherical  or  slightly  conical  bodies  (Fig. 
206).  They  consist,  as  in  the  Mosses,  of  a  wall  formed  by  one 
layer  of  cells,  which  encloses  a  number  of  spermatozoid-mother- 
cells  (A  and  B).  The  antheridia  when  ripe  absorb  water,  and 
are  ruptured,  and  the  spirally-coiled  spermatozoids  liberated  (Fig. 
206  S).  The  spermatozoids  have  been  observed  to  pass  down  the 
neck  of  the  archegonium,  and  to  fuse  with  the  oosphere. 

The  asexual  generation.  The  first  leaf,  the  "cotyledon," 
of  the  embryo  developed  from  the  oospore  (Figs.  202,  205)  is 
always  small,  and  has  a  very  simple  shape.  The  leaves  which 
occur  later  become  more  perfect,  stage  by  stage,  until  the  perma- 
nent form  of  leaf  has  been  attained. — The  STEM  is  most  frequently 
a  subterranean  or  a  semi-aerial  rhizome ;  it  is  only  in  the  tropical, 
palm-like  Tree-Ferns,  that  the  stem  raises  itself  high  in  the  air  and 
resembles  that  of  a  tree,  with  leaf -scars  or  with  the  remains  of 
leaves  attached  (Figs.  207,  203) ;  in  certain  species  the  stem  is  en- 
cased in  a  thick  mat  of  aerial  roots  (Dicksonia  antarctica).  When 
the  rhizome  is  horizontal  the  internodes  are  frequently  elongated, 
and  the  leaves  are  arranged  in  two  rows,  as  in  Polypodium  vulgare 
and  in  the  Bracken-Fern  (Pteridium  aquilinum),  etc. ;  it  is  also 
generally  dorsiventral,  having  a  dorsal  side  on  which  the  leaves  are 
situated,  and  a  ventral  side,  different  from  the  former,  on  which 
the  roots  are  borne.  When  the  stem  ascends  in  an  oblique  direc- 
tion, or  is  nearly  vertical,  its  internodes  are  extremely  short,  and 
the  leaves  are  arranged  in  a  spiral  line  with  a  complicated  phyllo- 
taxis,  e.g.  in  Athyrium  filix-fcemina,  Aspidium  filix-mas,  etc.  The 
BRANCHING  upon  the  whole  is  extremely  slight,  and  is  generally 
confined  to  the  petiole  (e.g.  Aspid.  filix-mas) ,  or  to  the  stem  near 
the  insertion  of  the  leaves.  Several  species  normally  form  buds  on 
different  parts  of  the  lamina.  The  buds  which  are  formed  on  the 
stem  are  not  confined  to  the  leaf-axil  as  in  the  higher  plants. 
The  Tree-Ferris,  generally,  do  not  branch  at  all. 

The  VASCULAR  BUNDLES  are  concentric,  with  the  wood  surrounded 
by  the  soft  bast.  In  tranverse  section  they  are  seen  as  circles  or 
irregularly-shaped  figures  (Fig.  203),  the  name  of  "  King  Charles 
and  the  Oak"  (Bracken-Fern)  having  originated  from  the  appear- 
ance which  the  bundles  present  in  oblique  section.  In  Osmunda 
they  are  collateral  and  resemble  those  of  the  Flowering-plants. 
Round  each  individual  bundle  is  often  a  sheath  of  thick- walled, 
hard,  brown,  sclerenchymatous  cells,  which  act  as  a  mechanical 

w.  B.  P 


208 


PTERIDOPHYTA. 


tissue ;    similar    strands    are    also    found    in    other    parts    of   the 
stem. 

The  LEAVES  in  nearly  all  species  are  only  foliage-leaves,  borne 


FIG.  207.— Various  Ferns  (1,  2,  3,  4). 

in  a  spiral.  They  have  an  apical  growth  which  continues  for  a 
long  time,  and  some  require  several  years  for  their  complete  de- 
velopment. In  the  buds  they  are  rolled  up  (circinate) ;  not  only 
the  midrib,  but  also  all  the  lateral  veins,  and  even  the  terminal 


PTEEIDOPHYTA.  209 

portions  of  a  leaf  are  sometimes  rolled  up  together,  the  tissues  of 
the  leaf  being  already  fully  developed  and  only  waiting  to  expand. 
The  leaves  are  often  excessively  divided  and  compound,-  with 
pinnate  branches,  and  have  an  epidermis  with  stomata  and  a  well- 
developed  system  of  venation.  Stipules  are  only  found  in  Marat- 
tiacece  and  Opliioglossacece. 

Very  often  peculiar  hairs  or  scales  (palece,  ramenta),  dry,  brown, 
flat  and  broad,  are  found  on  stem  and  leaf. 

The  SPORANGIA  are  small,  round  capsules,  which,  in  a  very  large 
number  of  Ferns,  are  formed  on  the  back,  but  more  rarely  on  the 
edge  of  the  ordinary  foliage-leaves.  It  is  very  seldom  that  there 
is  any  difference  in  form  between  the  barren  foliage-leaves  and  the 
fertile  leaves,  as  is  found  for  example  in  Blechnum  spicant  or 
Struthiopteris ;  or  that  the  fertile  part  of  the  leaf  is  differently 
constructed  from  the  barren  portion  of  the  same  leaf,  as  in  the 
Royal-Fern  (Osmunda).  In  such  instances  the  mesophyll  of  the 
fertile  parts  is  poorly  developed. 

The  sporangia  in  the  Polypodiacece  are  lens-shaped,  with  long 
stalk  (Fig.  211  D)  :  their  wall  consists  of  one  cell-layer  on  which 
a  single  row  of  cells,  passing  vertically  over  the  top  (that  is  along 
the  edge  of  the  sporangium),  is  developed  into  the  "ring"  (annu- 
lus).  The  cells  of  the  annulus  are  very  much  thickened  on  the 
inner  and  side  walls,  and  are  yellowish-brown.  The  thickened 
cells,  however,  do  not  entirely  encircle  the  sporangium,  and  on 
one  side,  near  the  stalk,  they  pass  over  into  large,  flat,  thin-walled 
cells.  These  form  a  weak  point  in  the  wall,  and  it  is  here  that 
the  sporangium  is  opened  diagonally  by  the  elongation  of  the 
annulus.  The  sporangium  of  the  Polypodiacese  opens  as  it  dries. 
The  cells  of  the  annulus  are  very  hygroscopic,  and  in  straighten- 
ing, the  annulus  bends  back  with  a  jerk,  thus  ejecting  the  spores 
to  considerable  distances.  The  cells  of  the  annulus  absorb  water 
with  great  readiness.  [The  sporangium  arises  as  a  single  epi- 
dermal cell,  from  which  a  basal  stalk-cell  is  cut  off.  Three  oblique 
cell-walls,  intersecting  near  the  base,  are  next  formed  in  the  upper 
cell,  and  a  fourth  between  these  and  parallel  to  the  free  surface ; 
an  inner  tetrahedral  cell  enclosed  by  four  others  is  thus  formed, 
the  outer  cells  become  the  wall  of  the  sporangium,  while  the  inner 
cell,  by  a  series  of  walls,  parallel  to  its  sides,  cuts  off  a  layer  of 
cells  which  eventually  form  the  tapetum,  the  remaining  central 
•cell  constituting  the  archesporium.] 

The  SPORES  are  either  oblong  and  bilateral,  or  they  are  tetra- 


210  PTERIDOPHYTA. 

hedric  with  curved  sides,  depending   upon  the  way  in  which  the 
tetrad  division  has  taken  place. 

The  sporangia  are  almost  always  situated  on  the  nerves  and 
gathered  into  groups,  sort,  which  differ  in  form  in  the  various- 
genera.  The  sori,  in  many  genera,  may  be  covered  by  a  scale-like- 
structure,  the  indusium  (Figs.  211  B,  212). 

In  the  majority  of  cases,  each  sorus  is  situated  on  a  small 
papilla  (placenta,  or  receptacle),  which  is  supplied  by  a  small  vascu- 
lar bundle.  Between  the  sporangia,  hairs  (paraphyses)  are  often 
situated,  which  spring  either  from  the  placenta  or  from  the  stalks 
of  the  sporangia. 

Systematic  Division.  The  Ferns  may  be  divided  into  -two- 
groups,  characterized  by  the  structure  and  development  of  the- 
sporangia.  The  sporangia  in  the  EUSPORANGIAT^E  take  their  origin 
from  a  group  of  epidermal  cells,  and  their  walls  are  formed  by 
several  layers  of  cells.  The  archesporium  is  the  (not  tetrahedric) 
hypodermal  terminal  cell  of  the  axial  row  of  cells  which  give  rise- 
to  the  sporangium.  In  the  LEPTOSPORANGIAT.E  the  sporangia  are 
developed  from  single  epidermal  cells,  and  their  walls  are  uni- 
layered.  The  archesporium  is  a  central,  often  tetrahedric  cell,, 
from  which  sixteen  spore-mother-cells  are  developed.1  It  is  diffi- 
cult to  say  which  form  is  the  oldest  (according  to  Prantl,  those- 
which  have  the  sori  on  the  nerve-endings)  ;  however,  the  Eusporan- 
giatise  would  seem  to  have  made  their  appearance  long  before  the 
others,  and  also  well  denned  Marattiaceae  and  Ophioglos&acese- 
occur  in  the  Kulm  and  Coal  period,  before  the  true  Polypodiaceae. 

About  4,000  species   of   Ferns  are  now  existing,   and  they  are-- 
found especially  in  tropical  and  sub-tropical  forests, 
Family  1.     Eusporangiatae. 

Order  1.  Ophioglossaceae.  The  prothallium  differs  from 
that  of  all  other  Ferns  in  being  subterranean,  free  from  chlorophyll,, 
pale  and  tuberous.  The  stem  is  extremely  short,  with  short 
internodes,  most  frequently  unbranched,  vertical,  and  entirely 
buried  in  the  ground  (Fig.  208  st}.  In  several  species  (among 
which  are  the  native  ones)  one  leaf  is  produced  every  year,  which 
has  taken  three  to  four  years  for  its  development.  In  Botry- 
chium  a  closed,  sheath-like  basal  part  of  each  leaf  cover's  the  sub- 
sequent leaves  during  their  development.  In  Ophioglossum  and 

1  The  position  of  the  annulus  varies  in  the  different  orders ;  longitudinal 
in  Polypodiaceae,  Hymenophyllaceaa,  and  Cyatheacese ;  transverse  in  Schizaa- 
acese,  Gleicheniacese :  indistinct  or  apical  in  Osniundacese,  Ophiaglossaceae,, 
Marattiaceae,  Salviniaceae,  Marsiliaceae. 


PTERIDOPHYTA. 


211 


others  each  leaf  has  at  its  base  an  intrapetiolar,  cap-like  sheath, 
which  protects  the  succeeding  leaf.  The  leaves  are  of  two  kinds  : 
(a)  foliage,  which  in  Ophioglossum  vulgatum  are  lanceolate  and 
entire,  but  in  Botrychium  however,  are  pinnate  (6  in  Fig.  208 
J,  J5);  and  (6)  fertile,  which  are  found  facing  the  upper  side  of 


3. — .4  Oi  Jiioglossum    vulgatufn,   (Adder's-tongue) :   B  Botrycliium    lunaria  (Moon- 
wort),  both  natural  size ;  r-n-oots;  bs  leaf-stalk ;  ststem;  l>  foliage-leaf ;  /fertile  leaf. 

the  foliage-leaves.  These  latter  in  Ophioglossum  are  undivided 
and  spike-like  (Fig.  209  A),  but  pinnate  in  Botrychium  (Fig.  208  B). 
Each  foliage  and  fertile  leaf  are  branches  from  the  same  petiole. 
The  large  sporangia  are  placed  laterally,  and  open  by  two  valves. 
No  annulus  is  formed  (Fig.  209). — Ophioglossum  reproduces  vege- 
tatively  by  adventitious  buds  on  the  roots. 


212 


PTERIDOPHYTA. 


Three  genera  with  about  twelve  species. 

Order  2.  Marattiacese  are  tropical  Ferns,  whose  gigantic 
leaves  resemble  those  of  the  Polypodiacese,  but  have  stipules  in 
addition.  The  sporangia  are  grouped  in  sori,  situated  on  the  lower 
side  of  the  leaves,  the  sporangia  in  each  sorus  being  arranged 
either  in  two  rows  or  in  a  ring.  In  Angiopteris  they  are  isolated 
(Fig.  210  A),  but  in  the  other  species  (Kaulfussia,  Dancea,  Marattia), 
they  are  united,  and  form  "synangia"  divided  into  a  number  of 
chambers  corresponding  to  the  sporangia.  These  open  by  clefts 
or  pores.  Marattia  presents  the  highest  development,  as  its 
sporangia  are  completely  united  in  a  capsule-like  synangium, 
which  is  closed  until  maturity,  and  then  opens  by  two  valves. 
In  each  valve  there  is  a  row  of  three  to  eleven  sporangia,  each 
opening  by  a  slit  towards  the  inside  (Fig.  210  B,  C).  An  indusium 
encloses  the  sorus,  except  in  Kanlfussia ;  it  is  formed  of  flat  and 
lobed  hairs,  which  resemble  the  hairs  of  the  other  portions  of  the 

leaves.  In  An giopteris  and  Mar- 
attia the  indusium  is  very  rudi- 
mentary ;  in  Dancea  it  forms  a 
kind  of  cupule. 

The  numerous  fossil  Marattiaceae- 
(15  genera,  with  98  species)  present 
similar  differences  to  those  now  living, 
but  more  various  forms  are  found,  for 
example,  with  solitary  free  sporangia. 
Those  now  living  are  the  last  small 
remnant  (4  genera  with  only  2& 
species)  of  a  once  dominant  family, 
which  existed  from  very  early  times, 
and  whose  culminating  point  was 
reached  in  the  Kulm  and  Coal  periods. 
The  Ophioglossaceas  appear  also  in 
the  Kulm  and  Coal  periods,  and  were 
about  as  numerous  as  at  the  present  time  (presumably  2  genera,  with  19 
species).  Leptosporangiate  Ferns  appear  however  to  have  occurred  first  of  all 
in  the  Trias-formation. 

Family  2.     Leptosporangiatse. 

Order  1.  Polypodiaceae.  Sporangia  on  the  lower  side  of  the 
leaves,  talked  and  provided  with  a  vertical,  incomplete  annulus  \ 
dehiscing  by  a  transverse  cleft  (Fig.  211  D). — The  genera  are 
distinguished  by  the  form  of  the  indusium  and  the  position  of 
the  sori,  etc. 


FIG  .210.— Sporangia  of  the  Marattiaceae  : 
A  Angiopteris;  B  and  C  Marattia;  C  is  a 
half  sorus  with  nine  sporangia,  each  of 
which  Las  opened  by  a  longitudinal  cleft. 


PTER1POPHYTA. 


213 


1.  The  sporangia  cover  the  entire   lower  surface   of   the   leaf 
(Tropical  America  and  Asia).     Acrostichum,  Platycerium. 

2.  Sori  without  indusia,  circular  or  oval.    Polypodium  (Fig.  211 
A}.     The  leaves  are  most  frequently  situated  in  two  rows  on  the 
dorsal  side  of  the  creeping  rhizome,  and  fall  off  leaving  a  smooth 
scar  behind. — P.  vulgare,  common  in   woods,  on  stones.     (Phegu- 
pteris  also  has  no  iridusium  ;  see  page  214). 

3.  The  sporangia  are  situated  in  continuous  lines  just  inside  the 
margin  of  the  leaf. — Pteris1 :  the  sporangia  form  a  continuous  line 
along  the  entire  margin  of  the  leaf  (Fig.  211  C),  which  bends  over 
and  covers  the  sporangia,  forming  a  "  false-indusium."     Pteridium 
has  linear  sori  situated  on  a  marginal  vascular  bundle,  covered 
by  two  linear  basal  indusia,  of  which  the  outer  is  bent  over  like 
the  edge  of  a  leaf. — P.  aquilinum  (Bracken;  has  a  wide-spreading 


FIG.  211.— Portions  of  leaves  with  sori.  A  Polypodium.  B  Aspidium.  C  Pteridium.  I> 
A  sporangium  of  one  of  the  Polypodiaceae  :  r  the  annulus ;  s  spores. 

rhizome  with  large  alternate  leaves,  placed  on  opposite  sides,  at 
some  distance  apart.  Only  one  leaf  is  developed  from  each  branch 
every  year. 

Adiantum  (Maiden-hair)  :  sori  on  the  underside  of  small  portions  of  the  edge 
of  the  leaf,  which  are  bent  over  (false  indusium).  Cryptogramme  (Allosorus), 
Cheilanthes. 

4.  The  sori  are  oval  or  linear,  situated  on  one  side  of  the 
vascular  bundle. — Asplenium  (Fig.  212  J.)  ;  sori  linear;  indusium 
with  one  of  its  edges^  attached  at  the  external  side!  A.  ruta  mura- 
ria  (Wall-Rue)  ;  A.  septentrionale  ;  A.  trichomanes. — Athyrium  :  sori 
linear  cr  curved;  A.  filix-fuemina  (Lady-Fern). —  Scolopendrium 
1  The  former  genus  Pteris  is  divided  into  Pteris  and  Pteridium. 


214 


PTEEIDOPHYTA. 


(Fig.  212  B}  :  sori  as  in  Asplenium,  but  situated  in  pairs  across 
the  lanceolate,  entire  leaves.  Each  sorus  is  covered  on  the  ex- 
ternal side  by  an  indusium,  whose  free  edges  are  parallel  and  ap- 
proach each'othe'r.  S.  vidgare  (Hart's-tongue). — Blechnum  (B.  spicant, 
Hard  Fern  ;  the  fertile  leaves  differ  from  the  barren,  the  pinnae  being  narrower, 
while  the  underside  is  almost  entire'y  covered  with  sori,  and  hence  they  are  of 
a  much  darker  brownish  hue  than  the  barren  ones) — Ceterach  :  indusium 
rudimentary  or  absent. 

5.  Sori    circular    and   covered    by    a    shield-like,    or   reniform 
indusium. — Aspidium  (Fig.  211  B) ;  the  leaves  wither  away  and 
leave  no  scar  upon  the  root-stock.     A.  filix-mas  (Male-Fern)  ;  A. 
spinulosum. — Phegopteris  has  no  indusium,  the  withered  bases  of 
the  leaf-stalks  are  persistent ;  P.  dryopteris  and  P.  polypodioides. 

6.  The  indusium  is  situated  below  the  sori,  and  has  the  shape  of 
a  one-sided  scale  (Cystopteris,  Struthiopteris},  or  of  a  cup  or  cupule, 
which  in  Woodsia  is  sometimes  fimbriate  (Fig.  212  0,  D). 


FIG.  212.—^!  Asplenium.  B  Scolopendrium.  C  Woodsia ;  D  single  sorus  of  the  same. 
E  Cyathea  :  the  sporangia  have  fallen  off  in  the  upper  sori.  (All  magnified.) 

7.  The  sori  are  situated  on  the  margin  of  the  leaf,  and  at  the  end  of  a 
vascular  bundle.  Indusium,  semi-cupnlar.  Davallia.  Principally  tropical 
species.  1  in  S.  Europe. 

This  order  is  the  greatest,  comprising  about  2,800  species,  the 
majority  being  perennial  plants.  A  few  are  large,  and  known  as 
Tree-Ferns. 

As  plants  in  conservatories  and  rooms  the  following  are  cultivated :  species  of 
Gymnogrartime  (tropical  America),  Lomaria,  Nephrolepis,  Pteris  (P.  serrulata, 
cretica). 

Officinal.  Aspidium  filix -mas,  rhizome  and  the  withered  petioles.— Species  of 
Alsophila  and  Cibotium  give  Penghawar  Djambi.  The  rhizome  of  Pteridium 
aquilinum,  var.  esculentum,  contains  so  much  starch  that  it  is  used  as  food. 

The  other  orders  of  true  Ferns  deviate  from  the   Polypodiacese,  especially  in 


PTERIDOPHYTA. 


215 


the  formation  of  the  ammlus,  the  bursting  of  the  sporangium  and  its  mode  of 
attachment  and  development,  and  in  the  differences  in  the  formation  of  the  pro- 
thallium,  etc.  The  principal  are  : — 

Order  2.  Hymenophyllaceae.  To  this  order  belong  the  lowest  and  most 
Moss-like  Ferns ;  the  leaves,  with  the  exception  of  the  veins,  are  most  frequently 
formed  of  only  one  layer  of  cells,  and  consequently  stomata  are  wanting  ;  the 
formation  of  the  prothallium  also  somewhat  resembles  the  Mosses.  Sori 
marginal,  on  the  extremities  of  the  vascular  bundles,  and  surrounded  by  a 
cupular  indiisium.  The  sporangia  are  sessile,  with  equatorial  annulus.  Hy- 
menopliyUum  (H.  tunbridgense,  European).  Trichomanes  (T.  speciosum,  Euro- 
pean). Species  about  200,  which  live  especially  on  rocks  and  trees  in  damp 
and  shady  tropical  forests.  Some  have  no  roots. 

Order  3.  Cyatheaceae.  Annulus  complete  and  oblique.  To  this  order  be- 
long, principally,  the  tree-like  Ferns  with  palm-like  habit.  The  number  of 
ppecies  is  about  200,  .they  are  all  tropical  and  form  forests  in  some  regions  of 
Australia.  Cibotium  and  Dicksonia  have  marginal  sori,  with  cupular,  basal 
indusium.  (The  stem  of  D.  antarctica  is  covered  with  aerial  roots.)  Alsophila 
(without  indusium) ;  Cyathea  with  cupular,  inferior  indusium  (Fig.  212  E). 


FIG.  213.— Gleiclienia :  A  part  of  a  leaf  with  sori ;  B  a  single  sorus. 

Order  4.  Gleicheniaceae.  Sporangia  with  equatorial  annulus,  and  longi- 
tudinal dehiscence,  most  frequently  groups  of  3-4  in  sori  without  indusium  (Fig. 
213).  Gleiclienia:  the  apical  growth  of  the  leaves  continues  for  a  long  time. 

Order  5.  Schizaeaceae.  Annulus  apical.  To  this  order  belongs  Aneimia, 
which  is  so  commonly  cultivated  in  conservatories.  The  two  lowest  pinnae  are 
metamorphosed,  having  no  leaf -parenchyma  and  being  covered  with  sporangia. 
Schizcea.  Mohria.  Lygodium,  a  climber,  whose  leaves  have  unlimited  growth 
and  attain  a  length  of  several  metres.  About  70  species.  Tropical. 

Order  6.  Osmundaceae.  The  sporangia  have  at  the  apex  a  lateral  group 
of  strongly  thickened  cells,  which  gradually  pass  over  into  the  ordinary  cells. 
The  sporangia  open  by  a  longitudinal  cleft.  Indusium  wanting.  Osmunda 
bears  the  sporangia  upon  peculiar,  branched  pinnae,  without  parenchyma  (the 
uppermost  in  the  leaf).  0.  regalis  (Royal-Fern) :  European. 


Sub-Class  2.     Hydropterideee  (formerly  Rhizocarpae), 
Water  Ferns. 

The  following  further  characteristics  must  be  added  to  those 
given  on  page  205  : — 


216 


PTERIDOPHYTA. 


Sexual  generation.      The  MICROSPORES  produce  an  extremely 
rudimentary  prothallium,  formed  of  only  two  cells,    and   having 


D 


FIG.  214.— Sulrinta  natans  :  A  miorosporanginm  with  germinating  microspores  and  pro- 
trading  prothallia  (s) ;  B  a  prothallium  with  the  bicellular  antheridium  (s)  growing  out  of 
the  microsporangium ;  C  the  two  cells  of  the  antheridium  have  opened  by  transverse 
clefts  ;  beneath  is  seen  the  microspores  enclosed  by  the  hardened  mucilage  ;  D  spermato- 
zoids  still  enclosed  in  the  mother-cells. 

also  a  very  much  reduced  bicellular  antheridium  with  a  small 
number  of  spermatozoid  mother-cells  in  each  cell  (in  Salvinia  4,  in 
Marsilia  and  Pilularia  16).  In  Salvinia  the  microspores  remain 
embedded  in  a  hard  mucilaginous  mass  (at  first  frothy)  which  fills 
up  the  cavity  of  the  sporangium.  The  prothallinm  must  therefore 


FIG.  215.—  Sahinia  natans.  A,  B  Female  prothnllin,/-/,  protruding  from  the  macrospore 
which  is  still  enclosed  in  the  macrosporangium;  ce  archegonia.  C  An  embryo  (x  16) 
still  in  connection  with  the  spore  (s) :  a  the  scutiform  leaf ;  b-e  the  subsequent  foliage- 
leaves,  of  which  b  and  c  stand  singly,  d-e-v  in  a  whorl ;  v  the  submerged-leaf ;  /-/  wing, 
like  lobes  of  the  prothallium  ;  m  the  foot. 


PTER1DOPHYTA.  217 

grow  out  through  this  slime  and  also  through  the  wall  of  the  spor- 
angium (Fig.  214),  and  it  thus  terminates  in  a  relatively  long 
cell. 

In  Marsilia  the  microspores  are  set  free  from  the  microsporan- 
gium,  and  the  prothallia,  with  the  antheridia,  remain  in  them 
until  the  spermatozoids  are  liberated.  The  latter  are  spirally- 
twisted  threads. 

The  MACROSPORES,  on  germination,  give  rise  to  a  very  reduced  pro- 
thallium,  which  in  Salvinia  bears  3  archegonia  ;  but,  if  these  are 
not  fertilised,  the  prothallium  may  continue  to  grow  and  become  a 
fairly  large,  green  body  with  several  archegonia  (Fig.  215  A,  B). 
In  Marsilia  the  prothallium  is  still  more  reduced,  it  is  enclosed  in 
the  macrospore,  and  only  bears  one  archegonium.  The  archegonia 


FiG.  21G.— Salvinia  natan?.  A  An  archegonium,  unripe,  teen  in  longitudinal  section: 
h  the  neck-cells;  k  the  neck-canal-cclls  ;  c  the  central  cell.  B  An  open  atchegonium  of 
which  the  neck-cells  have  separated  off.  C  An  open,  old  archegoniuin  seen  from  the  top. 

are  similar  in  structure  to  those  of  the  Ferns,  but  are  smaller,  and 
sunk  more  deeply  in  the  tissue  of  the  prothallium. 

The  asexual  generation  is  developed  from  the  fertilised 
egg-cell.  It  is  a  dorsiventral,  horizontal  shoot.  In  Salvinia  it  bears 
at  first  a  shield-like  leaf,  the  scutiform  leaf  (Fig.  215  (7,  a),  which 
is  succeeded  by  the  ordinary  foliage-leaves.  The  young  plants 
of  Marsilia,  likewise,  have  less  perfect  leaves  in  the  very  early 
stage. 

The  formation  of  the  sporangium  is  the  same  as  in  the  Lepto- 
sporangiate  Ferns.  (The  16  spore-mother-cells  originate  from  one 
central,  tetrahedric  archesporium.) 

The  Hydropteridese  are  divided  into  2  orders,  the  chief  differ- 
ences between  them  being  found  in  the  asexual  generation. 


218 


PTERIDOPHYTA, 


Order  1.     Salviniacese.     This  order  more  nearly  approaches 
the  true  Ferns,  especially  so  on  account  of  the  form  of  the  indusium. 


A  B 

FIG.  2l7.—Salvinia  -naians   (natural  size):  A  seen  from    above,  floating  on  the  water; 
B  a  portion  seen  from  the  side  in  its  natural  position  in  the  water. 

Only  one  species  is  found  in  Europe,  Salvinia  natans  (Fig.  217). 
This  is  a  small,  floating,  annual,  aquatic  plant,  entirely  destitute  of 
roots.  The  dorsiventral,  horizontal  stem  bears  two  kinds  of  leaves, 
which  are  arranged  in  whorls  of  three.  Two  of  these  which  turn 
upwards  are  oval,  entire,  "  aerial  foliage-leaves  "  (Fig.  217  J5,  62-63)  ; 

the  third;  the  "water-leaf"  (61)  is 
submerged  and  divided  into  a 
number  of  hair-like  segments, 
similar  to  the  submerged  leaves 
in  many  aquatic  plants,  for  in- 
stance, Water-buttercup  (see  also 
Fig.  215  C).  The  whorls  of  leaves 
alternate  with  each  other ;  there 
are  thus  4  rows  of  dorsally-placed 
aerial  leaves,  and  two  rows  of 
ventrally-placed  submerged  leaves. 
The  sporangia  are  situated  in  sori, 
each  sorus  being  borne  on  a  small 
column  (receptacle  or  placenta)  and  enveloped  by  a  cupular, 
but  entirely  closed  indusium  (Fig.  218).  The  sori  are  situated  on 


FIG.  21S. — Sori  of  Salvinia  in  longi- 
tudinal section  :  h  microsporangia ;  m 
macrosporangia.  (x  10.) 


PTERIDOPHYTA. 


219 


the  submerged  leaves  (Fig.  217  B,  s-s)  and  are  unisexual,  i.e.  each 
sorus  contains  micro-sporangia  only,  or  macrosporangia. 

Azolla  belongs  to  this  order.  It  is  a  very  small,  floating,  tropical  water-plant 
(America  and  East  India),  with  horizontal,  root-bearing  stem.  The  stem  branches 
profusely  by  lateral  buds,  and  bears  the  two  rows  of  leaves  on  its  dorsal  side, 
the  roots  on  the  ventral  side.  Each  leaf  is  bifid,  and  divided  into  an  upper  dorsal, 
and  a  lower  ventral  portion.  The  upper  segments  float  on  the  surface  of  the 
water  and  are  arranged  like  tiles  on  a  roof,  each  one  overlapping  its  neighbour. 
In  each  floating  segment  a  large  cavity  is  found,  in  which  Anabcena  is  always 
present.  The  lower  segments  are  submerged. 

Order  2.  Marsiliacese.  The  characteristic  feature  of  this 
order,  and  one  not  possessed  by  other  Fern-like  plants,  is  that  the 
sori  (2-many)  are  enveloped  in  leaf- 
segments  which  close  round  them  and 
form  a  "  sporocarp,"  just  in  the  same 
manner  as  the  carpels,  in  the  Angio- 
spermous  Flowering-plants,  close  round 
the  ovules  and  form  ovaries.  The  sori 
contain  both  micro-  and  macro-sporangia. 
When  the  spores  are  ripe,  the  sporocarp 
opens  in  order  to  disperse  the  spores 
(Fig.  220). 

The  two  genera  (with  57  species, 
Temperate,  Tropics)  are  land-  and  marsh- 
plants,  whose  dorsiventral,  creeping  stem 
bears  roots  on  the  under  surface,  and  the 
leaves  in  two  rows  on  the  upper  side 
(Figs.  219,  221).  The  leaves  *of  Mar- 
silia  are  compound,  and  divided  into  four 
small  leaflets  springing  from  the  apex 
of  the  petiole  (Fig.  219),  and  resemble 
the  leaves  of  Oxalis.  In  the  bud  the 
leaves  are  circinate  (Fig.  219  6),  and  at 
night  they  exhibit  the  well-known  sleep- 
movements.  The  sporocarps  are  borne 
on  the  petioles  of  the  fertile  leaves,  near 
their  bases  (Fig.  219 /)  ;  they  are  oblong 
and  resemble  small  beans,  the  outer  cells 
being  hard  and  sclerenchymatous,  while  FIG.  219.  —  Manill 

,  i  n  •    •  i     i   •  i  (natural  size) :   K  terminal  bud ; 

the  inner  ones  are  divided  into  a  number      j  leayes ;  f  s;porocarps .  x  point 
of    loculi   arranged   in   two   rows.      On     of  branching  of  petiole. 


220 


PTERIDOPHYTA. 


germination,  water  is  absorbed,  the  two  sides  separate  slightly,  aa 
valves  (Fig.  220  A),  and  a  long  vermiform  mass  of  gelatinous, 
parenchymatous  cells  (Fig.  220),  swollen  by  the  water,  emerges, 
bearing  a  large  number  of  sori  arranged  pinnately.  Each  sorus 
(sr)  is  covered  by  a  thin  indusium.  (The  thin  covering  may  be 
considered  an  indusium  physiologically,  though  not  morpho- 
logically). 

Marsilia  quadrifolia,  in  Europe.     Many  species  are  found  in  Australia.    The 


FIG.  220.— Marsilia  salvatrix :  A  the 
•eporocarp  commencing  to  germinate;  B 
a  more  advanced  stage  of  germination. 


FIG.  221. — Pilularia  globulifera  (natural 
size) :  s  sporocarps ;  b  leaves ;  fc  the 
growing  point;  r  roots. 


nutritious  sporocarps  of  M.  salvatrix  were  the  means  of  saving  the  Burke  ex- 
pedition in  the  interior  of  Australia,  and  hence  this  species  lias  earned  its  specific 


Pilularia  has  linear  leaves,  without  lamina.  The  sporocarps  are 
spheroid  (Fig.  221),  brown  and  hard,  and  situated  near  the  base  of 
the  leaves.  They  are  2-4  chambered  and  open  by  a  corresponding 
number  of  valves. 


PTERIDOPHYTA. 


221 


Class  2.    Equisetinae    (Horsetails.) 

The  characteristics  of  this   class  have  been  described  on  page 
204. 

It  is  divided  into  two  sub-classes  : — 

1.  THE  ISOSPOROUS  EQUISETINJE.     To  this  sub-class  belong,  with 
certainty,  only  the  EQUISETACE^E  now  existent,  which   are    repre- 
sented by  only  one  genus,  Eqtiisetum. 

2.  The  HETEROSPOROUS  EQUISETINAE.     Forms  which   are  now  ex- 
tinct. 

Sab-Class  1.     Isosporous  Equisetinae. 

Order.     Equisetaceae  (Horsetails). 

The  sexual  generation.     The  pro  thallium  is  green  and  leaf- 
like,  as  in  the  majority  of  Ferns,  but  irregularly  branched  and 


Fio.  222.— Equi*etum  arvense.  Tde  prothallium  highly 
magnified.  A  Male ;  s,  s  autheridia.  B  Portion  of  a 
female,  cut  through  vertically  ;  ce  ce  archegonia,  the  central 
one  is  fertilised  ;  h  h  root-hairs. 


FIG.  223.- — EquifetummaX' 
tmttm.  Spermatozoids  :  a 
shows  them  still  enveloped 
by  the  mother-cell. 


curled.  It  is  often  unisexual.  The  male  pro thallia  bear  antheridia 
only,  and  are  smaller  and  less  branched  (Fig.  222  A)  than  the 
female ;  the  latter  may  attain  a  diameter  of  |  an  inch,  and  bear 
archegonia  only  (Fig.  222  B).  The  antheridia  and  the  archegonia 
resemble  those  of  the  Ferns,  but  the  spermatozoids  (Fig.  223)  are 


222 


PTERIDOPHYTA. 


larger  and  less  twisted.  On  the  last  curve  is  situated  a  more  or 
less  elongated  appendage  of  cytoplasm  (Fig.  223  c). 

The  asexual  generation.  The  embryo  is  similar  to  that  of 
the  Ferns.  The  fully  developed  Equisetum.  is  a  perennial  herb, 
with  widely  creeping  (in  some  species  tuberous)  rhizome,  from 
which  extend  erect,  aerial,  most  frequently  annual  shoots. 

The  vegetative  aerial  STEMS  are  divided  into  a  number  of  inter- 
nodes  by  the  whorls  of  leaves  (Fig.  224).  The  internodes  are  hol- 
low, the  cavities  being  separated  from  each  other  by  the  transverse 


FIG.  221.— Equisetum  arvense  :  a  fertile  branch  with  cone  ;  b  vegetative  shoot;  c  cone  ; 
d  sporophylls. 

partitions  of  the  solid  nodes.  The  lower  portion  of  the  internode, 
which  is  encased  by  the  leaves,  has  much  thinner  and  softer  cell- 
walls,  so  that  the  stem  is  easily  separated  into  segments  just  above 
the  nodes.  Each  internode  has  a  large  number  of  ridges  and 
furrows,  and  bears  at  its  apex  a  whorl  of  leaves  whose  number  and 
position  correspond  to  the  ridges  of  the  internode.  As  in  the 
case  of  other  Verticillate  plants,  the  whorls  are  placed  alternately, 
one  above  the  other  ;  the  same  arrangement  is  also  found  in  the 
ridges  on  two  successive  internodes.  In  addition  to  the  large 
air-cavity  in  the  centre  of  each  internode  (the  central  cavity),  a 


PTERIDOPHYTA.  223 

whorl  of  tubular  air-passages  is  found  in  the  cortex  of  the  stems, 
opposite  the  furrows  (vallecular  canals).  There  is  also  a  similar 
air-passage  (carinal  canals)  in  each  of  the  vascular  bundles,  which 
are  placed  in  a  ring,  one  opposite  each  ridge,  and  therefore  alter- 
nating with  the  vallecular  canals.  The  vascular  bundles  are  col- 
lateral as  in  the  majority  of  Flowering-plants,  but  poorly  deve- 
loped. The  xylem  of  each  bundle  consists  of  two  groups  of 
annular  or  spiral  vessels,  close  to  the  outer  border  of  the  carinal 
canal,  and  two  groups  of  scalariform  tracheids,  each  placed  on  a 
radius  passing  through  a  group  of  spiral  vessels.  The  phloem  is 
placed  between  these  four  groups,  each  of  which  has  only  a  few 
vessels.  The  stiffness  of  the  stems  is  mainly  due  to  the  large 
amount  of  silica  in  the  cell-walls  of  the  epidermis,  and  to  the 
sclerenchymatous  cells  of  the  ridges. 

All  LEAVES  are  situated  in  whorls.  The  VEGETATIVE  are  simple, 
undivided,  1-nerved,  and  are  united  into  toothed  sheaths  (Fig. 
224  a,  &).  The  branching  of  the  stems  in  some  species  (E.  arvense) 
is  very  abundant.  The  branches  break  through  the  base  of  the 
leaf -sheaths  (Fig.  224  6),  and  generally  alternate  with  the  teeth 
(leaves). 

The  FERTILE  LEAVES  (sporophylls)  are  different  from  the  barren 
ones.  They  are  free,  shield-like,  each  one  having  a  short  stalk 
bearing  usually  an  hexagonal  plate  (Fig.  224  ciT),  and  closely  com- 
pressed into  an  ear  or  cone  (Fig.  224  a,  c).  The  Equisetums  thus 
present  an  advance  in  development  distinctly  beyond  that  of  the 
Ferns,  which  is  further  emphasized  by  the  circumstance  that  a 
transition  from  the  sheath-leaves  to  the  fertile-leaves  is  found  in 
the  involucre  or  annulus,  a  "  collar  "  of  specially  modified  leaves 
situated  at  the  base  of  the  cone  (Fig.  224  a  and  c).  The  cone 
may  be  considered  as  a  very  rudimentary  flower,  and  the  annulus 
may  be  regarded  as  a  very  early  stage  in  the  formation  of  a,  flower 
(perianth).  See  page  235. 

The  SPORANGIA  are  situated  on  the  underside  of  the  sporophylls, 
one  at  each  angle ;  they  are  sac-like,  and  open  inwardly  by  a 
longitudinal  cleft  (Fig.  224  d).  An  annulus  is  wanting;  but  in 
the  wall  of  the  sporangium,  as  in  the  pollen-sacs  of  the  Flower- 
ing-plants, a  layer  of  cells,  with  annular  or  spiral  thickenings,  is 
developed,  which  assists  in  the  dehiscence  of  the  sporangium. 

The  SPORES  are  green;  the  walls  composed  of  three  distinct 
layers,  of  which  the  outer  is  gradually  separated,  except  at  one 
point,  and  becomes  split  into  four  long  bands  (elaters)  (Fig.  225). 

W.  B.  Q 


224  PTERIDOPHYTA. 

The  elaters  are  extremely  hygroscopic,  coiling  round  the  spore 
when  moistened,  and  expanding  as  soon  as  dry,  presenting  a  most 
lively  object  under  the  microscope  when  breathed  upon  and  allowed 
to  dry.  The  second  layer,  when  germination  commences,  becomes 
detached  from  the  inner  wall,  which  is  formed  of  the  exospore 
and  endospore. 

The  order  has  become  much  reduced,  and  ut  the  present  time 
includes  only  one  genus,  Equisetum,  with  about  twenty-five  species, 
which  are  distributed  over  the  entire  globe,  particularly  in  damp 
situations.  In  SOME  SPECIES  the  barren  shoots  are  green  and  very 
much  branched,  but  the  fertile  ones  are  unbranched,  pale  brown, 
and  possess  no  chlorophyll  (E.  arvense,  Field-Horsetail,  Fig.  22  J, 
and  E.  maximum).  IN  OTHERS  the  fertile  and  barren  shoots  are 
alike  green,  and  either  both  unbranched  (E.  Memale),  or  branched 
(E.  palustre,  E.  limosum,  etc).  The  fertile  shoots  of  E.  sihaticum, 


A 

FIG.  225.  -  Spores  of  Equisetum  :  A  rtnmp.  with  elaters  (e)  coiled  round  tbe  spore ;  B  dry, 
wish  elaters  expanded. 

up  to  maturity,  resemble  those  without  chylorophyll  of  E.  arvense, 
but  after  that  period  they  produce  green  branches,  and  thus  re- 
semble the  barren  ones. 

EXTINCT  ISOSPOROUS  EQUISETIN.E.  In  addition  to  several  true  species  of 
fossilized  Equisetums,  the  order  of  the  CALAMITES,  which  no  doubt  is  closely 
allied  to  the  Equisetinse,  is  also  found  in  the  fossil  state.  These  were  gigantic 
forms,  attaining  about  twenty  times  the  size  of  those  of  the  present  clay,  and 
stems  of  nearly  10-12  metres  in  height  are  known.  They  reached  the  culmina- 
ting point  of  their  development  in  the  Carboniferous  period,  and  died  out 
towards  the  close  of  the  Palaeozoic.  The  stems  had  hollow  internodes  and 
a  ternating  grooves,  similar  to  their  relatives  of  the  present  day.  The  leaves 
must  either  have  bet-n  absent  or  very  perishable,  since  they  have  not  been 
identified  with  certainty.  If  the  determinations  of  certain  remains  of  cones 
which  of  late  have  been  discovered  are  correct,  they  were  heterosporous  and 
had  two  kinds  of  sporangia  as  in  the  following  sub-class.  A  cambium  forma- 
tion and  an  increase  in  thickness  has  been  found  in  the  stems. 

Their  USES  are  very  limited.  A  few  species,  such  as  E.  hiemale  are  used  for 
polishing  on  account  of  the  hard  siliceous  cell-waUs  of  the  epidermis,  found  in 
all  species  of  Equisetum. 


PTEKIDOPHYTA. 
Sub-Class  2.     Heterosporous  Equisetinae. 


225 


The  two  orders  which  come  under  this  head  are  united  by  the  characteristics, 
that  the  verticillate  leaves  are  not  united  into  sheaths  (Fig.  226),  and  that  be- 
tween each  whorl  of  fertile  leaves  there  is  also  a  whorl  of  barren  ones.  The 


FIG.  226.— A.  fragment  of  Annularia. 

fertile  whorls  in  ATJNULAKI^;  are  situated  about  midway  between  the  barren  ones 
(Fig.  227),  but  in  ASTEKOPHYLLITE.E  they  occur  immediately  above  a  barren 
whorl  (Fig.  228)  and  contain  only  half  as  many  members  as  the  latter.  The 
lower  whorls  bear  macrosporang'a  with  one  macrospore,  the  upper,  micro- 
sporangia  with  many  microspores. 


FIG.  227. — Fragment  of  Annularia  longi- 
/olia,  with  sporangia ;  the  leaves  have 
partly  fallen  off :  a  barren  whorls  ;  s  fer- 
tile whorls. 


FIG.  228.— Fragment  of  cone  of  Astero- 
phyttites  (Volkmannia  elongata)  -.  a  and  s 
as  in  Fig.  227. 


The  ANNUIABI^E  were  distichous  (Fig.  226),  and  presumably  floating  plants. 
The  ASTEKOPHYLLITE^E  had  verticellate  branches.  These  also  died  out  after  the 
Carboniferous  period,  at  the  close  of  the  Palaeozoic. 


226 


PTERIDOPHYTA. 


Class  3.     Lycopodinee  (Club-Mosses), 

The  characteristics  of  this  class  have  been  given  on  page  205v 
It  consists  of  two  sub-classes,  one  embracing  isosporous,  the  other 
heterosporous  forms. 

Sub-Class  1.      Lycopodieae  (ISOSPOROUS  Lycopodina3). 

One  kind  of  spore.  Prothallium  large,  partly  green.  Leaves- 
without  ligule. 


PIG..  229.— Lycopodium  annotinum:   A    em-  FIG.    230. — Lycopodium  clavatum? 

byro  (nat.  size),  with  prothallium  (pr),  one  portion  of  astern,  bearing  cones  (a) ; 

embryo  is  broken    off;    JB  the  prothallium  s,  a  spore;  7i  sporangium  in  the  axil 

(slightly  magnified) ;   C  section  through  the  of  a  leaf,  s. 
prothallium  and  embryo  in  the  direction  a-b 
sf  A,  and  vertically  in  the  plane  of  the  paper. 

Order  1.  Lycopodiaceae.  The  PROTHALLIUM  is  DOW  known  in 
nearly  all  the  species  ;  it  is  always  more  or  less  tubercular,  and 
bears  both  antheridia  and  archegonia. 

In  L.  annotinum  the  prothallium  is  a  relatively  large  mass  of  cells,  without 
chlorophyll,  and  subterranean,  in  which  the  antheridia  and  archegonia  are 
embedded  (Fig.  229).  In  the  widely  distributed  tropical  species,  L.  cernuum,. 


PTERIDOPHYTA.  227 

and  in  L.  inundatum,  it  is  a  small  tubercular  body  which  has  a  subterranean 
portion,  with  either  little  or  no  chlorophyll;  and  an  aerial  green  portion. 
The  prothallia  of  L.  phlegmaria  and  others  live  saprophytically  in  the  crevices 
of  the  bark  of  trees  ;  they  are  partly  filamentous,  branched,  and  possess  no 
chlorophyll. 

The  asexual  generation.  PERENNIAL  PLANTS.  The  stem 
branches  monopodially  (often  apparently  dichotomously),  and  is , 
thickly  covered  by  small,  simple,  triangular  or  scale-like  leaves. 
The  leaves  are  spirally  arranged  in  some  species  (Figs.  229,  230), 
and  in  others,  whose  stem  is  compressed  with  unequal  sides,  oppo- 
site (Fig.  231).  The  roots  of  Lycopodium  are  dichotomously 
branched. 

The  SPORANGIA  in  Lycopodium  are  situated  singly  at  the  base  of 
the  leaves,  almost  in  their  axils  j  they  are  reniform,  unilocular  and 


FIG.  232. — Lycopodium  clavatum.    A 
tetrahedral  spore  seen  from  above, 
FIG.    231.  —  Lycopodium    complttnatum  :    a  where  the  three  borders  join  ;  and  a 

leaves  on  the  edges  of  the  stem ;  d  leaves  on  tetrad  of  bilateral  spores,  still  lying 

the  sides.  in  the  mother-cell. 

open  like  a  mussel-shell  by  two  valves  (Fig.  230  fc).  The  sporangia 
are  developed  from  a  group  of  surface  cells.  The  archesporium  is 
formed  from  one  hypodermal  cell  (or  perhaps  a  cell -row). 

The  fertile  leaves  are  collected  upon  definite  regions  of  the  stem. 
They  are  either  similar  to  the  barren  ones,  and  then  the  fertile 
portions  of  the  stem  pass  gradually,  without  any  break,  into  the 
barren  portion  (L.  selago)  ;  or  they  differ  from  the  barren  leaves, 
and  are  then  collected  into  special  apical  cones  (Fig.  230  a).  The 
SPORES  are  tetrahedral  or  bilateral  (Fig.  232). 

About  100  species,  chiefly  tropical. 

Five  species  of  Lycopodium  are  found  in  Great  Britain.  L.  clavatum  and  L. 
selago  are  common  in  mountainous  districts.  L.  annotinum  is  common  in  the 


228 


PTERIDOPHYTA. 


Highlands  of  Scotland.  The  other  genus  of  the  order  is  Phylloglossum,  with 
one  species,  P.  drummondi  (Australia,  Tasmania,  and  New  Zealand),  a  small 
plant  only  a  few  centimetres  high,  with  two  tubers,  and  about  eleven  linear 
leaves  at  the  base  of  the  stem  which  is  terminated  by  a  cone  of  sporophylls. 
— FOSSIL  Lycopodiaceas  in  the  Carboniferous  period. 

OFFICINAL:  "Lycopodium,"  the  spores  of  L.  clavatum. 

Family  2.  Psilotaceae.  The  sporangia  are  placed  on  the  apex  of  short,  two- 
leaved  stems,  as  2-3,  seldom  four,  small  capsules.  Small  herbs,  with  angular 
stems;  leaves  small,  simple,  and  one  nerved.  Only  four  species.  —  Psiloium 
(Madagascar,  Moluccas,  Sandwich  Islands,  etc.)  is  destitute  of  roots,  their  place 
being  supplied  by  special  underground  stems  which  bear  a  few  modified  leaves, 
very  much  reduced,  especially  when  buried  deeply  in  the  soil.  Three  species. 
— Tmesipteris  (Australia),  one  species. 

Sub-Class  2.     Selaginelleae  (HETEROSPOROUS  Lycopodinse). 
Micro-  and  macro-spores.  The  prothallia  are  very  much  reduced, 


especially   the  male;  the  female   does    not  leave  the   spore. 
leaves  are  ligulate. 


The 


A  D  C 

FIG.  233.  —  Germination  of  the  microspores  of  Sclfaginclla :  A  the  spore  rendered 
transparent,  seen  from  above.  In  the  iutci'ior  is  seen  the  prothallinm  (/),  and  the  first 
divisions  of  the  antheridium  (a,  b,  c,  d) ,  in  B  the  spore- wall  is  removed  and  all 
the  spermatozoid-mother-cells  formed;  in  C,  the  microspore  has  opened  and  the  spermato- 
zoids  and  the  mother-cells  are  escnping  togetl,  er. 

The  sexual  generation.  In  the  MICROSPORES  are  formed: 
(1)  a  very  small  "  vegetative  "  cell  (/  in  Fig.  223  A,  B),  and  (2) 
a  cell  many  times  larger  (the  antheridium)  which  divides  into  a 
number  (4-8)  of  peripheral  cells,  enclosing  2-4  inner  cells,  which 
last  form  a  large  number  of  spermatozoid-mother-cells  in 
Selaginetta  but  only  4  in  Isoetes.  On  germination,  when  the 
spore- wall  is  ruptured,  the  spermatozoids  and  spermatozoid-mother- 
cells  are  ejected  into  the  water. 

The  SPERMATOZOIDS  in  Selaginella  are  elongated  and  club-shaped, 
with  two  cilia  (Fig.  234) ;  but  in  Isoetes  lacustris  they  are  spirally- 
twisted  threads  which  differ  from  all  other  spermatozoids  by 


PTER1DOPHYTA. 


229 


having  a  bunch  of  cilia  at  each  end ;  the  other  species  of  Isoetes 
have  cilia  only  at  the  anterior  end. 

The  MACROSPORES.  Shortly  after 
the  macrospores  have  been  set  free, 
or  in  Selaginella,  while  still  en- 
closed in  the  sporangium  of  the 
mother- pi  ant,  they  germinate  and 
soon  become  filled  with  the  cellular 
tissue  of  the  prothallium,  and  even 
in  Selaginella  the  archegonium  be- 
gins to  be  formed  before  the  rupture  of  the  spore-cell-wall  has 
commenced  (Fig.  235  A). 

The  ARCHEGONIA  are  constructed  on  the  same  plan  as  those  of  the 


FIG. 234.— Spermatozoidsof  Selaginella: 
b  with  a  remnant  of  cytoplasm. 


FIG.  233. — Macrospore  of  Selaginella :  A  longitudinal  section,  before  the  rupture  of  the 
wall,  six  weeks  after  being  sown.  The  endosperm  (e)  has  not  yet  filled  the  entire 
chamber.  Cell-formation  is  still  proceeding  in  the  lower  part  of  the  spore.  The  endosperm 
nnd  prothallium  (//)are  separated  by  a  distinct  line  (diaphragm).  C  G.;rminatiug  macro- 
spore  seen  from  outside :  s  wall  of  the  spore  ;  CB  archegonia. 


FIG.  238. — Archegonia  of  Selaginella  :  A  unripe,  in  longitudinal  section;  c  the  central 
cell;  k  neck-canal-cell,  which  is  wedged  in  between  the  two-storied  neck-cells;  B  ripe; 
•u  ventral  canal-cell ;  C  seen  from  above,  open.  It  will  be  noticed  that  the  neck  is  formed 
of  two  tiers  of  four  cells  each. 


230 


PTERIDOPHYTA. 


other  Arch  ego niatae,  but  are  quite  embedded  in  the  pro  thallium 
(Figs.  235  ce,  236). 

The  asexual  generation  varies  very  much  in  the  different 
orders. 

Order  1.  Isoetaceae  (Quill-  worts).  The  only  known  genus, 
Isoetes  (Quill-wort),  has  an  extremely  short,  tuberous,  unbranclied 
stem  with  very  short  internodes  (Fig.  237).  The  STEM  is 
remarkable  as  being  the  only  one  among  the  Vascular  Cryptogams 
which  increases  in  thickness  (see  page  202).  The  meristematic 


239 
FIG.  237.— Isoetes  lacustris  (slightly  diminished):  st  the  stem  ;  v  roots  •  b  leaves. 

FIG.  238.— Isoetes  lacustris.  Longitudinal  section  through  the  hase  of  the  leaf  with  a 
microsporangium.  The  edge  of  the  groove,  in  which  the  microspangium  is  placed,  is 
continued  as  a  thin  covering  which  envelopes  the  sporangium.  The  inferior  edge  of  the 
ligular  groove  (L)  forms  a  lip  (J) ;  t,  sterile  cell-rows  (trabeculae)  which  divide  the 
sporangium  into  compartments;  I  vascular  bundle. 

FIG.  239. — Selaginella  inosqualifolia  Cone  in  longitudinal  section  ;  microsporangia  are 
seen  on  the  left  side,  macrosporangia  on  the  right  (most  frequently  each  with  four 
macrospores). 

cells  are  situated  round  the  axial  cylinder,  and  form,  especially, 
parenchymatous  tissue  in  two  or  three  directions,  giving  rise  to 
2-3  grooves  in  which  the  dichotomously-branched  BOOTS  are 
produced.  The  LEAVES  are  arranged  spirally  in  a  close  rosette. 


PTERIDOPHYTA.  231 

They  are  awl-shaped  and  have  at  the  base  a  semi-amplexicaul 
sheath,  with  a  groove  (/cwea),  in  which  a  sporangium  is  situated 
(Fig.  238).  The  ligule  is  a  foliar  outgrowth  from  the  upper  edge 
of  the  groove. — The  MACROSPORANGIA  (each  with  a  number  of 
macrospores),  are  situated  on  the  outer  leaves,  the  MICROSPORANGIA 
(Fig.  238),  on  the  inner  ones.  Between  each  cycle  of  fertile 
leaves  there  are  a  number  of  imperfect  or  barren  ones,  as  in  the 
case  of  the  female  plant  of  Cycas.  The  spores  are  liberated  by  the 
decay  of  the  sporangium.  The  two  kinds  of  sporangia  develope  at 
the  commencement  in  the  same  way.  The  archesporium  is,  at 
first,  a  hypodermal  layer  of  cells  which  grow  out  in  the  direction 
perpendicular  to  the  surface  of  the  leaf,  and  divide  by  a  number  of 
walls  parallel  to  this  direction,  forming  a  sporogenous  mass  of 
cells.  Some  of  the  cell-rows  of  this  sporogenous  mass  lose  their 
rich  protoplasmic  contents,  and  are  arrested  in  their  growth  ;  thus 
incomplete  divisional  walls  of  sterile  cells,  "  trabeculce"  arise  in  the 
sporangium,  dividing  it  into  a  number  of  compartments  one  above 
the  other  (Fig.  238  £).  (The  trabeculee,  according  to  Groebel,  play 
the  same  part  as  the  nutritive  cells  of  the  sporangium  of  Riella  ; 
the  tapetal  cells,  as  in  the  Ferns,  are  in  a  great  measure  dissolved 
at  a  later  period.)  The  sporogenous  cell-rows,  in  the  micro- 
sporangia,  give  rise  to  a  large  number  of  spore-mother-cells,  but  in 
the  macrosporangia  only  one  spore-mother-cell,  with  tapetum,  is 
developed  from  each  fertile  archesporial  cell. 

The  two  native  species,  and  several  others,  are  aquatic  plants, 
the  remaining  species  are  land  plants,  or  are  amphibious.  About 
50  species.  In  temperate  and  tropical  regions. — FOSSIL  species 
in  the  Tertiary  period. 

Order  2.  Selaginellaceae.  This  order  contains  only  one  genus, 
Selaginella.  The  STEM,  in  the  majority  of  species,  is  dorsiventral, 
long  and  slender,  and  apparently  branches  dichotomously,  but  in 
reality  monopodially,  with  well  developed  lateral  shoots.  The 
LEAVES  are  small,  round,  or  ovate,  in  the  majority  of  species 
arranged  in  whorls  of  two  leaves  each ;  these  whorls,  however, 
are  not  decussate,  but  are  considerably  inclined  towards  each 
other,  an  arrangement  by  which  four  rows  of  leaves  are  produced, 
each  whorl  having  one  large  and  one  small  leaf.  The  two  leaves 
in  each  whorl  are  of  unequal  size,  the  smaller  one  being  placed 
on  the  upper  surface  and  the  larger  on  the  lower  surface  of  the 
stem  (Fig.  240).  Some  species  have  spirally-arranged  leaves,  more 
resembling  the  arrangement  in  the  Lycopodiums. 


232  PTERIDOPHYTA. 

The  FERTILE  LEAVES  most  frequently  differ  from  the  barren  ones, 
and  are  collected  into  spike-like  cones  (a  kind  of  flower;  Fig.  239). 
Micro-  and  macro-sporangia  are  found  in  the  same  cone  (Fig.  239). 
Each  sporangium  arises  from  a  group  of  superficial  cells  of  the 
stem,  directly  over  the  leaf  on  which  it  will  be  situated  later  on. 
Each  sporangium  has  a  hypodermal,  unicellular  archesporium, 
and  contains  a  layer  of  tapetal  cells  ;  these  are  dissolved  later, 
when  the  spores  are  ripe,  and  not  before  as  in  the  Ferns.  In  the 
very  early  stages  of  their  development, 
the  micro-  arid  macro-sporangia  are 
precisely  similar,  and  the  differences 
between  them  arise  later  on.  In  the 
microsporangium  all  the  spore-mother- 
cells  divide,  and  each  forms  four  tetra- 
hedrically -arranged  microspores  (Fig. 
204)  ;  but  in  the  macrosporangium 
only  four  macrospores  are  formed,  by 
the  division  of  a  single  mother-cell, 
while  the  remaining  spore-mother-cells 
are  aborted.  It  is  rarely  that  the 
macrosporanjria  contain  2  or  8  macro- 

FIG.  2-10.— Selaginella  martensii: 
K  lower  leaves ;  r  upper  leaves.  spores. 

For  the  GERMINATION  OF  THE  SPORES,  see  pages  228,  229.  The  protballium 
arises  in  the  macrospore  (/-/,  in  Fig.235^i),  probably  by  division  of  the  meniscus- 
shaped  protoplasmic  mass,  which  is  marked  off  at  the  apex  of  the  spore ; 
primordial  cells  are  thus  formed  which  later  on  are  surrounded  by  a  cell-wall. 
In  six  to  seven  weeks  after  sowing,  the  spore-wall  is  ruptured  by  the  growing 
prothallium,  which  already  has  developed  archegonia  (Fig.  235  ce-ce).  The 
prothallium  so  formed  does  not  occupy  the  entire  cavity  of  the  spore,  but  ftfur 
to  five  weeks  after  sowing,  the  large- celled  parenchyma  is  developed  in  the 
lower  portion  of  the  spore  by  free  cell-formation;  this  has  been  termed  by  Pfeffer, 
"  endosperm."  since  it  is  similar  to  the  endosperm  of  Flowering-plants.  Goebel, 
however,  has  termed  it  "  secondary  prothallium,"  as  the  homology  with  the 
endosperm  of  the  Angiosperms  is  very  doubtful. 

The  FERTILISED  OOSPHERE  divides  into  an  upper  (hypobasal)  and 
a  lower  (epibasal)  cell;  from  the  latter  alone  the  embryo  is  deve- 
loped with  its  root,  stem,  foot,  and  two  cotyledons,  and  the  former 
gives  rise  to  an  organ  which  appears  in  this  instance  for  the  first 
time,  but  which  occurs  in  all  Flowering-plants,  viz.  the  suspensor. 
This  forces  the  embryo  down  into  the  "  endosperm,"  which  is 
entirely  or  partially  absorbed  by  the  embryo.  In  the  case  of  the 
Flowering-plants  the  embryo  is  developed  with  its  longitudinal 


PTEEIDOPHYTA.  233 

axis  in  the  elongation  of  the  suspensor,  but  in  Selaginella  the  em- 
bryo is  situated  transversely  to  it. 

Selaginella  (300-400  species),  is  essentially  tropical,  only  one 
species  living  in  the  North  (S.  spinulosa),  but  others  grow  in 
Central  and  South  Europe. 

Order  3.  Lepidodendracese  are  extinct,  tree-like  Lycopods,  which  are 
found  especially  in  the  Lower  and  Middle  Carboniferous.  Vegetatively  they  are 
most  nearly  related  to  Lycopodium,  but  the  stem  attained  much  larger  propor- 
tions (about  eleven  metres  in  height  and  one  metre  in  thickness),  and  had  a 
cambium  by  which  it  increased  in  thickness.  It  was  regularly  dichotomous, 
and  closely  studded  with  spirally- placed  leaves,  which  left  behind  them  peculiar 
rhombic  scars.  The  large  cones  resemble  Pine-cones,  and  bore  sporangia 
much  larger  than  any  which  are  now  produced  (the  male  ones  as  much  as 
2  cm.'s  in  length).  The  macrosporangia  were  situated  at  the  base,  and  the 
microsporangia  at  the  apex. 

Order  4.  Sigillariacese.  These  are,  presumably,  another  group  of  extinct 
tree-like  Lycopods  (especially  in  the  Middle  Carboniferous).  The  name  has  been 
derived  from  the  seal-like  scars,  which  the  fallen  leaves  have  left  behind  in 
longitudinal  rows  on  the  grooved  stem.  The  rhizomes  of  these  plants  were 
formerly  termed  Stigmaria,  and  placed  in  a  separate  genus. 

Order  5.  Sphenophyllacese  form  an  entirely  extinct  group.  They  do  not 
definitely  belong  to  any  of  the  three  large  classes  of  Vascular  Cryptogams,  but 
it  is  perhaps  best  to  place  them  in  juxtaposition  to  these.  They  were  herbaceous 
plants  with  verticillate,  wedge-shaped  leaves,  with  nerves  branching  dicho- 
tomously  into  equally  strong  branches.  Micro-  and  macro-sporangia  were 
formed  in  the  same  cone  ;  and  were  situated  in  the  axils  of  the  leaves,  as  in  the 
Lycopods. 


The  Transition  from  the  Cryptogams  to  the 
Phanerogams. 

All  the  plants  considered  in  the  preceding  chapters  are  included 
in  the  term  CRYPTOGAMS;  all  in  the  following  chapters  under  the 
head  of  PHANEROGAMS  (see  page  3).  Hoffmeister's  pioneer  works 
(1851,  Vergleichende  Untersuchungen  der  hoheren  Kryptogamen,  etc.) 
and  the  numerous  researches  published  later  by  other  investigators, 
have  closed  the  gap  which  was  formerly  thought  to  exist  between 
these  plants  ;  so  that  we  now,  in  the  series  :  Bryophyta — Pterido- 
phyta— Grymnospermge — Angiospermce  see  the  expression  of  a 
single  line  of  development  in  accordance  with  a  definite  plan.  The 
forms  through  which  this  gradual  development  has  taken  place 
have  in  course  of  time,  however,  to  a  great  extent  died  out,  and 
only  single  links  of  the  chain  connecting  the  lowest  to  the  highest 
still  remain. 

THE  ALTERNATION  OF  GENERATIONS,  which  we  found  indicated  in 
certain  Thallophytes,  can  be  proved  with  the  greatest  clearness  in 
all  the  higher  Cryptogams,  from  the  Mosses  upwards ;  it  is  also 
found  in  the  Phanerogams,  but  not  in  such  a  pronounced  degree, 
because  one  of  the  generations  is  so  far  reduced  that  it  has  almost 
given  up  its  independence.  For  the  sake  of  greater  clearness,  we 
will  begin  with  the  comparison  of  the  sporophyte,  asexual  (second) 
generation. 

The  asexual  (2nd)  generation  of  the  Cormophytes. 

The  asexual  generation  which  follows  from  the  further  develop- 
ment of  the  fertilised  oosphere,  is,  in  the  Mosses,  only  the  sporo- 
gonium  (according  to  one  theory  it  is  perhaps  homologous  with  a 
spore-bearing  leaf,  situated  upon  a  short  stem,  see  p.  187)  ;  in 
Filicince,  Egtiisetince,  and  Lycopodince,  on  the  other  hand,  it  is  a 
highly  developed  plant  differentiated  into  stem,  leaf,  and  true  root, 
and  bearing  the  sporangia  on  its  leaves.  The  MODIFICATION  OF  THE 
SHOOT  is  very  slight  in  Filicince.  The  first  leaves  of  the  embryo 
are  very  simple  in  form  (Fig.  205),  but  after  a  certain  age  all  the 
leaves  which  arise  are  essentially  alike.  The  fertile  leaves  do  not 

234 


CRYPTOGAMS   AND    PHANEROGAMS.  235. 

differ  from  the  barren  ones,  and  are  found  associated  with  them, 
and  their  formation  does  not  limit  the  growth  in  length  of  the. 
stem.  It  is  only  in  a  few  of  the  true  Ferns,  and  in  the  Hydro- 
pteridege,  that  the  fertile  leaves  differ  considerably  from  the 
barren  ones.  A  division  of  labour  in  which  certain  leaves  are  set 
apart  for  nutrition,  and  others  for  reproduction,  is  found  more 
pronouncedly  in  the  Equisetincv  and  Lycopodince,  for  in  these 
groups,  with  a  few  exceptions,  the  fertile  and  barren  leaves  are 
very  dissimilar ;  the  former  are  collected  in  special  ear-like  cones, 
which  terminate  the  further  growth  of  the  short  stems  on  which 
they  are  borne.  In  connection  with  the  cone,  leaves  are  sometimes, 
developed  which  form  a  transition  from  the  barren  to  the  fertile, 
ones  (the  "  annulus"  in  Equisetacece),  and  in  these  cases  the  first 
indication  of  a  flower  with  perianth  or  floral-leaves  is  to  be  traced. 
Among  the  Cryptogams  the  highest  division  of  labour  is  found 
in  Selaginella  and  Isoetes,  which  have  the  two  kinds  of  sporangia 
borne  on  different  leaves.  The  division  of  labour  (modification) 
is,  however,  still  more  pronounced  in  the  Phanerogams :  the  leaves 
which  bear  the  micros porangia  ("  pollen-sacs  ")  have  quite  differ- 
ent forms  from  those  which  bear  macrosporangia  (the  "  nucellus  " 
in  the  ovule),  the  former"  are  termed  stamens,  the  latter  carpels; 
in  certain  instances,  too,  there  is  even  a  contrast  between  the 
"  male  plants"  and  the  "  female  plants."  Moreover,  a  division  of 
labour,  in  a  much  greater  degree,  takes  place  in  the  leaves  which 
do  not  directly  take  part  in  reproduction,  and  it  is  thus  possible 
in  many  plants  to  draw  a  sharp  line  not  only  between  stamens  and 
carpels,  but  also  between  four  or  five  distinct  kinds  of  leaves,  which 
differ  in  form,  structure,  and  corresponding  functions,  and  which 
appear  in  regular  sequence  on  the  shoot:  namely,  between  "  scale- 
leaves  "  and  "  foliage-leaves,"  l  both  of  which  occur  in  the  Crypto- 
gams, and  the  "  floral-leaves,"  including  the  bracts  and  leaves  of  the 
"perianth,"  which  latter  often  differ  from  each  other  in  form  and 
colour,  and  are  then  separated  into  sepals  and  petals.  The  leaves — •. 
stamens  and  carpels — which  bear  the  sporangia  are  termed  sporo- 
phylls,  and  the  shoot,  or  extremity  of  a  shoot,  whose  leaves  are 
modified  into  sporophylls,  is  terminated  in  its  further  growth  by  their, 
production,  and  is  known  as  a  flower.  The  flower  which  is  most 

1  Floral-leaves  (hypsophyllary  leaves)  are  here  adopted  as  an  equivalent  of 
the  term  "  Hochblatter,"  to  signify  leaves  on  the  floral-shoot  other  than  foliage 
or  sporangia-bearing  leaves.  The  term  bract  is  applied  only  to  leaves  in  whose 
axil  a  flower  is  borne,  and  bracteoles  to  leaves  borne  on  the  flower-stalk  (pedicel). 


236 


CRYPTOGAMS   AND  PHANEROGAMS. 


completely  furnished  has  calyx,  corolla,  stamens,  and  carpels  ar- 
ranged in  this  order.  If  the  only  sporophylls  present  are  stamens, 
then  it  is  said  to  be  a  male  (staminate)  flower,  and  if  only  carpels, 
then  SL  female  (pistillate)  flower,  and  in  both  these  cases  the  flowers 
are  unisexual,  or  diclinous.  If  stamens  and  carpels  are  both  present 
in  the  same  flower,  it  is  termed  hermaphrodite.  Diclinous  plants 
in  which  the  female  flowers  are  situated  on  one  plant,  and  the 
male  flowers  on  another,  are  termed  dioecious ;  and  those  in  which 
the  same  plant  bears  the  two  kinds  of  flowers  are  termed  monce- 
vious.  When  the  male,  female,  and  hermaphrodite  flowers  are 
found  in  the  same  species,  the  plant  is  said  to  be  polygamous. 

The  sporangia-bearing  leaves  —  Sporophylls.  In  the 
Mosses  the  asexual  generation  is  only  represented  by  the  sporo- 

gonium,  and  if  the  theory  is 
correct  which  considers  the 
sporogonium  to  be  an  em- 
bryo consisting  of  a  rudi- 
mentary stem  and  terminal 
leaf,  then  the  spores  are  pro- 
duced on  the  leaves  in  these 
plants.  The  sporangia  in 
the  Filicinae  are  situated  in 
groups  (sori)  on  the  back  or 
on  the  edge  of  the  leaves. 
The  number  of  sporangia  in 
the  sorus  diminishes  very 
greatly  in  the  Marattiacere 
and  Gleicheniaceas  (three  to 
four  in  the  latter,  Fig.  213). 
In  the  Equisetina?  the  spor- 
angia are  situated  in  a  small 
number  on  the  under-side  of 
shield-like  leaves,  and  in 
Lycopodinae,  singly,  in  the 
axils  of  the  fertile  leaves,  which  are  alike  and  bear  either  micro- 
or  macro-sporangia.  In  the  Phanerogams  there  is  a  great  differ- 
ence between  the  stamens  and  carpels. 

Stamens.  In  the  lowest  Phanerogams  (Cycadeoi)  there  are 
many  indications  of  relationship  to  the  Ferns.  The  stamens  are 
flat  and  broad,  and  have  on  the  back  -many  pollen-sacs  (micro- 
sporangia)  arranged  in  small  groups  (true  sori),  which  even  have 


FIG.  241. — C'j/cas  :  a  stamen  (nat.  size)  scon 
from  the  under  side  ;  b  four  pollen-sacs,  not  yet 
open,  forminjr  a  "  sorus  "  ;  c  three  open  pollen- 
sacs  ;  d  a  pollen-grain. 


CRYPTOGAMS  AND  PHANEROGAMS. 


237 


a  small  "  placenta,"  similar  to  the  one  possessed  by  the  Ferns, 
and  open  towards  the  inside  by  a  longitudinal  cleft  (Fig.  241, 
compare  Fig.  213).  A  section  of  the  Coniferce  agree  more  closely 


FIG.  212.-  Stamens  of  Araucaria  (pcllin-  FIG.  243.— Male  flower  of  Taxus. 

sacs  long  and  pendulous). 

with  the  Equisetaceae,  in  having  a  few  (three  to  eight)  pollen- 
sacs  arranged  on  the  under-side  of  more  or  less  shield-like  leaves 
(Figs.  242,  243,  compare  with  Fig.  224  a,  c,  d).  In  the  Abietaceos 
the  number  of  sporangia  is  diminished  to  two,  which  are  placed 
also  on  the  lower  side  (Fig.  267)  of  a  stamen.  The  number  of 
pollen-sacs  (microsporangia)  in  the  Angiosperms  is  nearly  always 
four  to  each  stamen ;  they  are  longitudinal  projections  which  are 


f 


C    b 


FIG.  2ii. — A  Cross  section  through  a  quadrilocular  anther  in  different  stages  of  develop- 
ment :  s  the  searn  where  it  bursts  open  ;  vf  vascular  bundle  ;  fc  connective.  B  A  stamen. 
C  Another  stamen  seen  from  the  front  (/)  and  from  the  back  (6). 

placed  in  pairs  on  each  side  of  the  central  line  of  the  stamen,  two 
on  the  edge,  and  the  other  two  generally  on  the  side  which  is 
turned  inwards ;  the  pollen-sacs  generally  dehisce  longitudinally 


238 


CRYPTOGAMS    AND   PHANEROGAMS. 


(quadrilocular  anthers,  Fig.  244).  A  few,  for  instance  Orchiclacena 
and  Asclepiadaceas,  have  only  two  pollen-sacs  (bilocular  anthers)  ; 
and  in  others,  such  as  Solatium  and  the  Ericaceae,  they  open  by 
pores ;  in  Lauraceee  and  Berberidaceae,  by  valves.  The  part  of  the 
stamen  which  bears  the  pollen-sacs  is  termed  'the  anther.  Most  fre- 
quently this  is  supported  by  a  stalk  known  as  the  filament. 

Carpels.  The  simplest  forms  of  carpels  are  found  in  Cycas. 
In  this  genus  both  the  foliage  and  fertile  leaves  are  pinnate,  and 
hence  present  great  similarity;  the  ovules  (macrosporangia)  are 
situated  on  the  margin  of  the  central  portion,  just  as  the  sporangia 
are  placed  on  the  edge  of  the  fertile  leaf  of  Ophioglossum  (Fig. 
245,  compare  with  Fig.  209).  The  carpels  of  the  other  Cycade$» 
present  greater  divergence  from  the  foliage-leaves,  being  peltate, 
for  instance,  in  Zamia  and  Ceratozamia  (Fig.  246).  The  ovules. 


FIG.  215.— A  carpel  of  Cycas  revoluta  with  5 
ovules  (s),  at  half  to  one-third  nat.  size. 


FIG.  246.— Carpel  with  2  ovules  of 
Ceratozamia  robusta  (). 


in  the  Coniferaa  are  situated  on  the  upper  side  and  near  the  base 
of  the  ovuliferous  scales,  almost  in  the  same  position  as  the  spor- 
angia in  the  Lycopodinae  (Figs.  269,  272,  273  H,  compare  Figs. 
230,  239).  In  Taxus  the  uninclosed  ovule  is  placed  on  the  apex  of 
a  shoot  (Fig.  264).  In  all  these  plants  the  ovules  are  not  enclosed 
by  the  carpels,  that  is,  they  are  not  enclosed  in  chambers  formed 
by  the  turning  in  of  the  walls  of  the  carpel,  and  hence  the  name- 


CRYPTOGAMS   AND   PHANEROGAMS.  239 

Gyinnospermce  is  given  to  them.  In  the  higher  Flowering-plants, 
the  Angiospermce,  the  ovules  are  distinctly  situated  on  the  edge,  the 
upper  surface,  or  base  of  the  carpel ;  but  the  carpel  closes  round 
the  ovules  which  are  therefore  enclosed  in  a  chamber — the  ovary. 
In  a  few  cases,  for  example  in  the  Polygonaceae,  an  ovule  is  situated 
apparently  on  the  apex  of  the  stem  itself,  as  in  the  Yew  ;  in  other 
cases,  as  in  the  Primulacese,  many  ovules  are  apparently  devel- 
oped on  the  apex  of  the  stem,  which  seems  to  have  been  specially 
adapted  as  a  placenta,  but  it  is  also  possible  and  correct  in  these 
cases  to  suppose  that  the  ovules  are  in  reality  developed  on 
the  carpels.1  A  single  fully-developed  carpel  or  a  collection  of 
carpels  joined  together  is  termed  the  pistil.  The  extremity  of  the 
carpel,  which  is  specially  developed  to  catch  the  pollen-grains  and 
form  a  suitable  nidus  on  which  they  may  germinate,  is  called  the 
stigma.  The  united  edges  of  a  carpel  which  bear  the  ovules  are 
termed  the  ventral  suture.  The  back  of  the  carpel  forms  the  dorsal 
suture.  The  Marsiliaceae  take  a  position  among  the  Hydropterideae 
analogous  to  that  occupied  by  the  Angiosperms;  the  sporangia  are 
in  a  corresponding  manner  enveloped  in  a  closed  leaf. 

The  collection  of  stamens  in  a  flower  is  termed  the  andrcecium, 
and  all  the  carpels,  whether  individually  free  or  united  into  one 
pistil,  the  gynceceum. 

The  Sporangia.  The  asexual  generation  of  the  Mosses  is  the 
sporogonium,  in  which  the  spores  arise  in  tetrads  from  the  mother- 
cells.  The  sporangia  in  the  Filicince  take  their  origin  either  from 
a  single  cell  (Leptosporangiatse)  or,  what  probably  may  be  regarded 
as  an  older  stand-point,  from  a  group  of  cells  (Eusporangiatge).  In 
both  cases  there  may  be  distinguished  in  a  mature  sporangium 
three  tissues,  which  have  different  significance  (Fig.  204) :  (1)  an 
external  layer,  the  sporangium-wall,  most  frequently  composed  of 
one  layer  of  cells  made  up  of  cells  of  dissimilar  structure,  so  that  on 
desiccation  the  wall  is  ruptured  and  the  sporangium  opens  in  a 
definite  manner ;  (2)  an  internal  group  of  cells,  consisting  of  the 
spore-mother- cells ,  developed  from  an  archesporium,  and  which  by 
division  into  four  gives  rise  to  the  spores ;  (3)  a  layer  of  cells 
lying  between  the  two  already  mentioned,  which  is  dissolved  before 
maturity.  The  intermediate  cellular  layer,  which  directly  sur- 
rounds the  spore-forming  cells,  is  in  form  and  contents  more  worthy 
of  note  than  the  others,  and  is  termed  the  tapetum.  The  construc- 

1  It  may  be  here  remarked  that  another  explanation  is  possible,  based  on  the 
study  of  the  development  (K ) . 

\V.  B.  R 


240 


CRYPTOGAMS    AND    PHANEROGAMS. 


tion  of  the  sporangium  in  the  JBquisetince  and  Lycopodince  is  in  the 
main  the  same. 

In  the  PHANEROGAMS  the  Microsporangia  are  termed  Pollen- 
sacs.  They  take  their  origin  from  a  large  group  of  cells,  which,  in 
the  Angiosperms,  lie  immediately  beneath  the  epidermal  cells  of  the 
anther.  In  the  developed,  but  not  yet  mature,  sporangium  (pollen- 
sac)  there  are  to  be  found  :  (as  in  the  Vascular  Cryptogams)  (1)  an 
internal  group  of  mother-cells  which  give  rise  to  the  pollen-grains 


FIG.  217.— Development  of  an  anther.  A  Transverse  section  of  a  young  anther  of  Doroni- 
cum'macrophyllum.  The  formiitim  of  the  4  pollen-sacs  commences  by  divisions  of  th« 
hypodermal  cells  (at  m,  for  instance).  These  cells  divide  by  periclinal  walls  into  external 
cells  which  only  take  part  in  forming  the  anther-wall;  and  internal  cells,  which  corres- 
pond to  the  Archesporium,  and  from  which  the  spores  are  derived.  These  spore-forming 
cells  are  drawn  with  thicker  walls  in  B-E.  The  commencement  of  the  vascular  bundle  is 
seen  in  the  centre.  B  An  older  stage ;  the  pollen-sacs  already  project  considerably.  It 
is  the  cells  in  the  hypodermal  layer  which  are  active  and  in  which  tangential  divisions  par- 
ticularly occur  ;  /  v  vascular  bundle.  C  A  corresponding  longitudinal  section.  D  Trans- 
verse section  through  an  older  anther,  the  thickness  of  the  wall  outside  the  mother-cells 
of  the  pollen-grains  is  already  increased,  and  it  becomes  still  thicker  by  the  division  of  the 
hypodermal  cells:  its  most  external  layer  of  cells  but  one,  becomes  transformed  into  the 
"  fibrous  cells."  E  Transverse  section  of  a  still  older  pollen-sac  of  Menyanthes  ;  sm  are 
the  mother-cells  of  the  pollen-grains  surrounded  bythetapetum  (t),  external  to  the  tapetum 
is  the  anther-wall,  which  is  still  far  from  being  fully  developed.  The  sub-epidermal  layer 
becomes  "fibrous,"  and  the  cells  lying  inside  it  become  dissolved,  together  wit  h  the  tapetum. 

(microstores),  in  this  case  also  formed  in  tetrads  ;  (2)  a  group  of 
cells  surrounding  these,  of  which  the  internal  ones  form  a  tapetal 
layer,  similar  to  that  in  the  Vascular  Cryptogams  ;  the  tapetum 


CRYPTOGAMS    AND    PHANEROGAMS. 


241 


and  some  of  the  cells  surrounding  it  in  this  group,  become  dissolved 
before  maturity ;  the  more  external  ones,  on  the  other  hand,  are 
provided  with  peculiar  thickenings,  and  form  the  "  fibrous  "  layer 
by  the  aid  of  which  the  dehiscence  of  the  anther  takes  place ; 
(3)  an  external  layer,  the  epidermis,  enclosing  all  the  other  layers 
(Fig.  247). 

In  some  Coniferas  (Cupressus,  Thiija,  and  several  species  of  Juniperu*)  the 
microsporangia  (pollen-sacs),  which  are  situated  on  the  under  side  of  the  stamen, 
are  covered  by  a  thin  structure  which  seems  to  be  a  continuation  of  the  lamina 
and  which  is  supposed  to  be  homologous  with  the  indusium  of  the  Ferns. 

The  Ovule  in  the  Phanerogams  arises  most  frequently  on  a, 
projecting  portion  of  the  carpel,  termed  the  placenta.  The  ovules 
(compare  the  sporangium  of  the  Eusporangiatas  and  especially  the 


X         1LC 


FIG.  248.— Development  Of  the  ovule  in  the  Red  Currnnt,  Kibes  rubrum,  arranged  alpha- 
betically in  the  order  of  development.  A  Is  the  youngest  stage,  E  the  oldest,  tt  Inner 
integument;  ie  outer  integument ;  ncnucellus;  m  archespore  (mother-cell  of  the  embryo- 
sac). 

pollen-sac)  take  their  origin  from  a  group  of  cells  which  lies  beneath 
the  epidermis  (Fig.  248  A,  B).  First  of  all  a  small  papilla  is 
formed,  which  is  later  on  provided  with  a  vascular  bundle  and 
becomes  the  funicle ;  this  probably  has  the  same  value  as  the 
projections  ("placenta")  on  which  the  sori  in  the  Ferns  are 
attached.  Only  one  macrosporangium  (nucellus  ;  Fig.  248  nc} 
is  developed  at  the  apex  of  the  funicle.  This  arises  by  a  process  of 
•cell-division  exactly  .corresponding  to  that  by  which  the  pollen- 


242 


CRYPTOGAMS   AND    PHANEROGAMS. 


sacs  are  formed  (Fig.  248  C-E),  with  this  difference  only,  that 
while  a  great  many  cells  may  be  distinguished  in  each  pollen  - 
sac,  which  forms  pollen-grains  by  tetrad-division,  only  a  few  are 
found  in  the  ovule,  and  all  these  moreover  are  suppressed,  with  one 
single  exception  which  developes  into  the  macrospore  (embryo- 
sac)  without  undergoing  a  division  into  tetrads.  The  wall  of  the 
embryo-sac,  in  the  Gymnosperms,  may  be  thick  and  divided  into- 
two  layers  and  partly  cuticularized,  as  in  the  spores  of  the  Cryp- 
togams which  are  to  be  set  free.  In  the  Angiosperms,  on  the  other 
hand,  the  wall  is  extremely  thin. 

The  pollen-sac  thus  stands  in  the  same  relation  to  the  nucellu& 
as  the  microsporangium  does  to  the  macrosporangium  :  in  the 
pollen-sacs  and  microsporangia  a  number  of  spores  arise  by  the 
tetrad-division  of  several  mother-cells  ;  in  the  nucellus  and  macro- 
sporangium,  a  reduction  of  the  cells  already  formed  takes  place  to 
such  an  extent  that  the  number  of  macrospores  becomes  one 
(Salvinia,  Marsilia,  Phanerogams)  or  four  (Selaginella),  or  rarely  a 
large  number  as  in  Isoetes. 

In  the  Ferns,  as  stated  on  page  210,  etc.,  indusia  covering  the 
sori  very  often  occur.  Horsetails  and  Club-Mosses  have  no 
indusium;  but  in  all  Phanerogams  cupular  or  sac-like  structures 
(integuments}  are  found  which  envelop  the  nucellus.  These  de- 
velope  from  the  upper  end  of  the  funicle  (ii  and  ie,  in  Fig.  248  ;. 
y  and  i,  in  Fig.  249)  and  enclose  the  nucellus  on  all  sides  as  a- 

sac,   leaving  only  a 


small  channel  at  the 
apex  of  the  nucellus 
—  the  micropyle  — 
(Fig.  249)  through 
which  the  pollen- 
tube  proceeds  to  the 

A  an   erect  ovule    em':)I'JO  -  Sac. 
(orlhotropous)  ;  B  reversed  (anah-opous);  C  curved  (campylo-    ovules   of   the   Gym- 
tropous)  :  fc  the  nucellus  (shaded  in  all  the  figures)  ;  s  the  em-    nosperms   have    On!  V 

f  ? 

integument 

251    264    269 

274)  and  the  same 
is  the  case  with  the  majority  of  the  SympetalaB  and  a  few  Cho- 
ripetal^;  but  the  Monocotyledons  and  most  of  the  Choripetalfe- 
have  two  integuments  (Fig.  249). 

In  shape  the  integuments    resemble  very   closely   the    cupular 


FIG.  2  19.  -Various  forms   of  ovules 


bryo-sac  ;    c?i  the  base  of  the  ovule  (chalaza)  ;  y  and  i  the 
external  and  internal  integuments,  the  dotted  line  denotes    One 
the  place  where  the  scar  (hilum)  will  form  when  the  seed 
is  detached  from  the  f  uuicle. 


CRYPTOGAMS   AND    PHANEROGAMS.  243 

indusium  of  the  Hjmenophyllaceae,  certain  Cyatheacese  (Fig. 
212  E),  and  Salvinia  (Fig.  218)  ;  that  they  are  really  homologous 
with  these  is  probable,  but  is  not  proven.  Some  authorities 
regard  them  as  structures  found  only  in  the  Phanerogams. 

The  ovule  is  thus  a  "  monangic  "  (i.e.  reduced  to  1  sporangium, 
the  nucellus}  sorus,  situated  on  afunicle,  and  enclosed  by  one  or  two 
cupular  indusia,  the  integuments.  Some  of  the  ovules  are  erect 
(prihotropous) ,  others  curved  (campylotropous) ,  the  majority  reversed 
(anatropous)  (Fig.  249). 

[Goebel  (1884  and  earlier)  with  Strasburger  considered  the  entire  ovule  of  the 
Phanerogams  as  homologous  with  the  macrosporangium,  the  integuments  how- 
ever, as  new  structures  in  contradistinction  to  the  Ferns  :  the  funicle  then  corre- 
sponds to  tbe  stalk  of  the  sporangium.  The  integuments  of  the  ovule  (according 
to  Goebel,  1-582)  differ  from  the  indusium  of  the  Fern-like  plants  in  being  devel- 
oped from  the  basal  portion  of  the  nucellus  and  are  not,  as  in  the  Ferns  and 
Isoetes,  a  portion  (outgrowth)  of  the  leaf  which  bears  the  sporangia  (K).] 

The  nucellus  is  the  only  macrosporangium  which  never  opens; 
the  macrospore  remains  enclosed  in  it,  and  the  macrosporangium  remains 
attached  to  the  mother-plant.  It  is  therefore  essential  that  the 
method  of  fertilisation  which  is  employed  should  be  very  different 
from  that  of  the  Cryptogams.  The  pollen-grains  must  be  transferred 
to  the  ovule,  and  retained  either  by  a  drop  of  mucilage  at  the  micro- 
pyle  (Gymnosperms)  or  by  the  stigma  on  the  carpels  (Angiosperms). 
Fertilisation  by  spermatozoids,  which  are  ciliated  and  freely 
motile  has  been  discovered  in  the  Cycadese  and  in  Ginkgo. 

Many  other  modifications,  unknown  in  plants  of  more  simple 
structure,  take  place,  for  instance,  in  the  shoots  which  bear  the 
fertile  leaves;  especially  in  the  form  of  the  stem  or  thalamus  (hypo- 
gynous,  perigyrious,  epigynous)  ;  in  the  development  of  the  peri- 
anth which  stands  in  intimate  connection  with  the  special  means 
employed  to  effect  fertilisation ;  with  respect  to  the  different 
grades  of  union  found  in  the  leaves ;  in  the  union  of  the  flowers 
into  aggregations  of  a  higher  order  (inflorescences),  and  at  the  same 
time  the  production  of  "  floral-leaves  "  (page  235). 

The  sexual  generation.     The  Fertilisation. 

The  sexual  generation  in  the  Mosses  is  relatively  well  developed, 
because  not  only  the  protonema,  but  all  the  other  vegetative 
parts  of  the  Moss-plant,  in  addition  to  the  archegonia  and  anther- 
idia,  belong  to  it.  In  the  groups  which  follow,  a  gradual  but  in- 
creasing reduction  of  the  sexual  generation  takes  place,  and  at  the 


244  CRYPTOGAMS   AND    PHANEROGAMS. 

same  time  an  indication  of  sex  is  found  in  the  pro  thai  Ha,  which 
finds  expression  in  the  forms  of  the  spores  themselves.  In  the 
majority  of  cases  among  the  isosporous  Vascular  Cryptogams,  the 
sexual  generation — pro  thallium — is  a  green,  leafy  expansion  which 
can  sustain  itself  by  the  assimilation  of  carbonic  acid,  and  by  the 
absorption  of  nutriment  from  the  soil  by  means  of  root-hairs.  In 
some  plants  (Ophioglossacece,  Lycopodium  annotinum)the  prothallium 
is  a  subterranean,  pale,  tubercular  body,  but  in  these  instances  it 
is  relatively  large.  In  the  heterosporous  Vascular  Cryptogams  and 
in  the  Phanerogams,  the  prothallium  is  much  more  reduced,  both 
as  regards  its  size,  and  also  with  respect  to  the  number  and  struc- 
ture of  the  antheridia  and  archegonia. 

1.  The  Microspores.  Among  the  Hydropterides  the  con- 
tents of  the  microspore  divide  into  three  cells,  from  the  lower  one 
of  which  a  small  lenticular  cell  is  cut  off,  while  from  the  two 
upper  or  from  the  upper  one  only  (as  in  Azolld)  a  very  simple 
antheridium  is  developed.  The  male  prothallium  is  thus  reduced 
in  this  group  to  a  very  few  cells.  In  the  heterosporous  Lycopo- 
diacese  also,  the  prothallium  is  represented  by  one  small  lenticular 
cell,  and  only  one  antheridium  is  present  which  gives  rise  in 
Selaginella  to  a  large  number  of  spermatozoids  (Fig.  233),  but  in 
Isoetes  to  only  four.  The  spermatozoids  are  set  free  through 
rupture  of  the  microspore- wall,  by  the  swelling  of  the  wall  of  the 
antheridium.  When,  however,  the  microspores  are  not  liberated 
from  the  sporangium  (Salvinia),  the  upper  cell  of  the  prothallium 
elongates  and  perforates  the  walls  of  both  the  microspore  and  the 
sporangium  (Fig.  214)  in  order  to  protrude  the  antheridium,  thus 
resembling  the  protrusion  of  the  pollen-tube  in  the  Phanerogams. 

In  the  Phanerogams,  the  microspores  are  termed  pollen-grains. 

In  the  GYMNOSPERMS  the  male  prothallium  is  represented  by  a 
small  cell  cut  off  laterally  by  a  curved  wall  from  the  large  pollen- 
cell  (Figs.  250,  267),  the  vegetative  cell  of  the  prothallium.  The  large 
cell,  by  the  protrusion  of  the  endospore,  grows  out  into  a  tubular 
body  known  as  the  pollen-tube  (Fig.  250)  into  which  the  vegetative' 
nucleus  passes.  The  small  cell,  corresponding  to  the  rudiment  of 
the  antheridium,  divides  into  1,  2,  or  3  small  cells,  the  innermost, 
or  when  only  one  is  present  this  single  one,  being  the  antheridial 
cell.  This  divides,  either  after  cutting  off  an  inner  stalk-cell  or 
directly,  into  two  generative  cells,  which  in  Cycadacece  and  Gink- 
goaccce  are  liberated  from  the  pollen-tube  as  spermatozoids,  or,. 
as  in  the  higher  Grymnosperms,  pass  into  its  anterior  end  as  naked 


CRYPTOGAMS  AND  PHANEROGAMS. 


245 


cell  without  cilia.  After  the  rupture  of  the  membrane  at  the 
tip  of  the  pollen-tube  they  pass  over  into  the  archegonium,  reach 
the  oosphere  and  complete  the  fertilization,  the  sperm-nucleus 
fusing  with  the  nucleus  of  the  oosphere. 

In  the  ANGIOSPERMS  the  reduction  pro- 
ceeds still  further.  The  pollen-grain  contain  s 
two  cells,  a  vegetative  and  a  small  an- 
theridial  cell,  but  these  are  not,  as  in  the 
last  group,  separated  by  a  true  cell- wall ;  a 
membrane  at  most  being  formed  between 
them.  Both  these  cells  pass  into  the  pollen- 
tube,  but  the  vegetative  cell  disappears 
about  the  time  the  pollen-tube  reaches  the 
ovule :  while  the  antheridial  cell  divides 
into  two  :  one,  the  sperm-nucleus,  coales- 
cing with  the  nucleus  of  the  oosphere,  the 
other  uniting  with  the  definitive  nucleus. 

The  Gymnosperms  prove  in  yet  another 
point  that  they  are  more  closely  related  to 
the  Cryptogams  than  are  the  Angiosperms. 
When  the  pollen -grain  begins  to  germinate 
the  external  wall  ruptures  as  in  the  Cryp- 
togams (Fig.  250),  but  in  the  Angiosperms  special  germ-pores  are 
formed  in  the  cell-wall  f6r  the  emeigence  of  the  pollen-tube. 

2.  The  Macrospores.  The  prothallium  in  Salvinia  and  Marsilia 
is  still  rather  large,  green,  and  capable  of  the  independent  assimi- 
lation of  carbon.  It  projects  more  or  less  from  the  macrospore  and 
bears  (in  Marsilia  only  one,  in  Salvinia  several)  archegonia,  which 
however  are  embedded  to  a  greater  degree  in  the  prothallium,  and 
are  more  reduced  than  the  archegonia  of  the  true  Ferns  and  Horse- 
tails (Figs.  215,  216).  The  prothallium  is  still  more  reduced  in 
Isovtes  and  Selaginella  ;  partly  because  it  is  smaller  and  is  in  a  higher 
degree  enclosed  in  the  spore,  it  also  contains  less  chlorophyll,  or 
is  entirely  without  chlorophyll,  and  in  consequence  incapable  of 
independent  existence,  whilst  the  number  of  archegonia  is  less  ;  and 
partly  because  the  archegonia  are  themselves  reduced,  the  cells  of 
the  neck  are  fewer  and  embedded  to  the  level  of  the  surface 
of  the  prothallium  without  any,  or  with  only  a  very  slight 
projection  (Figs.  235, 236). — Finally,  the  prothallium  with  its  arche- 
gonia begins  to  develope  in  Selaginella  while  the  macrospore  is  still 
within  its  sporangium,  and  before  it  is  set  free  from  the  mother- 


FIG.  250.— I  Pollen-grains 
of  Cupressus  ;  at  the  top  is 
seen  one  prothallinm-cell. 
II  Germinating;  c  pollen- 
tube  ;  a  the  exine ;  b  the 
inline. 


246 


CRYPTOGAMS   AND    PHANEROGAMS. 


plant.     After  the  spores  are  set  free  and  germination  has  com- 
menced, the  spore-wall  ruptures  and  the  prothallium  is  exposed. 

The  GYMNOSPERMSgo  still  further.  The  macrospore  (embryo-sac) 
germinates  and  forms  internally  a  cellular  tissue,  designated  in 
former  times  by  the  name  of  albumen  (endosperm),  which  is  homo- 
logous with  the  prothallium.  It  always  remains  enclosed  in  the 
embryo-sac,  and  is  aparenchymatous  mass  containing  a  large  supply 
of  nourishment.  In  the  upper  part  of  the  endosperm  a  number 
of  archegonia  are  developed  which  are  in  the  main  constructed  in 


FIG.  251.  —Longitudinal  sec- 
tion of  ovule  of  Abies  cana- 
densis.  Inside  the  integumen< 
(i)  is  seen  the  nucellns,  n ; 
m  the  micropyle.  In  the  in- 
terior  of  the  nucellus  is  seen 
an  oval  mass  of  cells,  the  en- 
dosperm, and  at  its  top  two, 
archegonia,  c.  The  ovule  is 
turned  in  such  a  way  that  the 
micropyle  points  upwards, 
but  usually  it  turns  down- 
ward in  the  Abietinea. 


FIG.  252.— The  ape :  of  the  nacellus  (n)  of  an  ovule  o£ 
Abies:  I  long-shaped  cells  which  guide  the  pollen-tube; 
s  the  wall  of  the  macrospore  (embryo-sac)  ;  7i  the  neck- 
cells  of  the  archegonium  ;  fc  the  ventral  canal-cell;  and 
c  the  central  cell  (oosphere).  The  archegonia  of  the 
Cryptogams  should  be  compared  with  this  (see  pages  181, 
208,  216). 


the  same  manner  as  those  in  the  Cryptogams,  but  are  still  more 
reduced,  the  neck  consisting  only  of  4,  2,  or  1  cell  (Figs.  251,  252). 
The  ventral  canal-cell  is  also  formed,  in  the  majority,  as  a  small 
portion  cut  off  from  the  large  central  cell  just  beneath  the  neck  ; 
the  larger  remaining  portion  becomes  the  oosphere.  When  the 
pollen-tube  has  passed  down  to  the  oosphere  (Fig.  253)  and  fer- 
tilisation has  been  effected,  the  oospore  commences  a  cell-formation, 
the  final  result  of  which  is  the  formation  of  an  embryo  (the  asexual 
generation)  which  is  provided  with  a  thinner,  lower  end,  termed 
the  suspensor.  The  embryo  is  forced  more  or  less  into  the  endo- 


CRYPTOGAMS    AND    PHANEROGAMS. 


247 


sperm  in  which  it  may  rest  for  a  longer  or  shorter  time,  and 
generally  is  developed  to  such  an  extent  that  it  has  a  distinct 
primary-root  (radicle)  and  stem  (plumule)  with  one  or  more 
«mbryo-leaves  (cotyledons). 

When  the  oosphere  has  been  fertilised  its  nucleus  sinks  down  to  its  lower  end, 
and  by  repeated  division  into  two,  forms  four  cells  lying  in  one  plane  (Fig.  253, 
see  base  of  the  left  archegonium).  Three  tiers  of  cells  are  now  formed  by  trans- 
verse division  of  these  four.  It  is  the  intermediate  one  of  these  which  elongates 
and  forms  the  suspensor,  or  four  suspensors,  if  they  separate  from  each  other, 
which  push  the  lowermost  four  cells  deep  do.va  into  the  endosperm.  It  is 
from  these  four  lower  cells  that  the  embryo  (or  four  embryos  when  the  suspen- 


Fio.  25  i.— Embryo-sac  of 
Car  ex  prcecox:  syn  synergi- 
das ;  7ob  the  oosphere ;  c  the 
central  nucleus;  ant  the 
antipodal  cells. 

Fin.  253. —The  apex  of  the  nucellus  (n)  of  Abies  in  longi- 
tudinal section  :  c,  c  the  oospheres  of  the  two  archegonia  ; 
the  embryo-formation  has  commenced  at  the  bottom  of 
the  left  archegonium ;  s  wall  of  the  macrospore  ;  p  pollen- 
grains  ;  r  pollen-tubos. 

sors  separate)  is  developed,  but  never  more  than  one  embryo  attains  full  devel- 
opment. As  several  archegonia  are  contained  in  one  and  the  same  ovule,  all  of 
which  are  capable  of  forming  embryos,  there  is  the  possibility  that  several 
embryos  may  be  developed  in  a  seed  (polyembryony),  but  usually  only  one 
•embryo  attains  perfect  development. 

At  the  same  time  that  the  embryo  is  being  developed,  other 
•changes  are  taking  place  in  the  ovule,  especially  in  the  integument 
which  becomes  the  shell  of  the  seed  (testa).  The  endosperm  grows, 
and  the  embryo-sac  supplants  the  cells  of  the  nucellus.  The  seed 


248 


CRYPTOGAMS    AND    PHANEROGAMS. 


is  now  formed,  and  it  consists  in  its  most  complete  development, 
as  in  this  instance,  of  three  parts  : 

(1)  The  testa  of  the  seed,  formed  by  the  enveloping  integuments, 
with  the  remainder  of  the  tissue  of  the  nucellus  lying  outside  the 
embryo-sac  (the  macrosporangium). 

(2)  The  endosperm  or  prothallium. 

(3)  The  embryo. 

The  reduction  in  the  ANGIOSPERMS  is  carried  to  the  extreme 
limit.  In  the  embryo-sac  (the  macrospore)  the  nucleus  by  con- 
tinued division  produces  a  prothallium  consisting  of  primordial 
cells  (Fig.  254).  In  the  upper  end  of  the  embryo-sac  (which  is 
nearest  the  micropyle)  are  three  cells,  two  of  which  are  termed 
the  "co-operating  cells"  (synergidcR)  and  the  third  is  the  oosphere. 
Three  others  are  placed  at  the  opposite  end  of  the  embryo-sac  and 
are  therefore  termed  the  "  antipodal  cells."  Finally,  a  large  cell  is 
also  formed,  which  occupies  the  space  between  the  two  groups  and 
whose  cell-nucleus,  the  central  definitive  nucleus,  lies  in  the  centre 
of  the  embryo-sac.  These  primordial  cells  are  the  slight  rem- 
nant of  the  prothallium.  The  entire  structure  of  the  archegonium, 
with  its  neck  and  canal-cells,  has  disappeared,  and  nothing  is  left 
but  the  indispensable  oosphere.  When  the  oosphere  has  been 

fertilised,  and  has  com- 
menced the  cellular  divi- 
sions which  lead  to  the 
formation  of  the  embryo 
(Fig.  255),  the  synergidae 
and  antipodal  cells  are 
absorbed,  and  a  cell-for- 
mation begins  by  a  new 
process  which  emanates 
from  the  definitive  nu- 
cleus and  by  which  a  pa- 
renchymatous  cell-tissue, 
the  nutritive-tissue,  arises 
which  may  perhaps  be 
considered  as  homologous 
with  the  endosperm  of  the 
Gymnosperms.  The  dif- 
ference is  that  the  nutritive-tissue  of  the  Angiosperms  is  formed 
in  two  parts  with  an  intervening  interruption  ;  the  primary 
nutritive-tissue  is  first  formed,  and  after  fertilisation  is  absorbed, 


FIG.  255.— Diagrammatic  longitudinal  section 
through  an  anatropous  ovule  shortly  after  fertili- 
sation ;  a  and  i  are  the  two  integuments ;  /  the 
funicle  ;  fc  the  nucellus;  S  the  embryo-sac,  with  the 
incipient  formation  of  nutritive-tissue;  E  tie 
embryo;  P  the  pollen-tube  passing  through  the 
rnicropylo  (n)  to  the  oosphere. 


CRYPTOGAMS   AND    PHANEROGAMS.  249 

with  the  exception  of  one  cell,  which  continues  the  development 
and  gives  rise  to  the  nutritive-tissue  proper,  which  is  formed  in 
the  first  instance  of  primordial  cells,  and  later  on  of  a  cellular 
tissue  ;  this  nutritive-tissue  formed  in  the  embryo-sac  is  termed 
"  endosperm";  in  a  few  instances1  a  tissue  which  is  derived  from 
the  nucellus  functions  as  nutritive-tissue,  and  is  termed  "  peri- 
sperm."  In  many  plants  the  seeds,  when  ripe,  contain  a  very- 
rich  nutritive-tissue,  in  addition  to  the  embryo,  for  the  purpose  of 
its  nourishment  during  germination.  These  are  termed  albu- 
minous (endospermous)  seeds,  in  distinction  to  the  ex-albu- 
minous, or  those  in  which  the  nutritive- tissue  is  stored  in  the 
embryo  itself,  before  it  is  completely  developed,  and  used  for  its 
sustenance. 

In  addition  to  the  changes  which  fertilisation  produces  in  the 
ovule  itself,  it  also  gives  the  impetus  to  a  series  of  changes  in  the 
entire  shoot  which  bears  the  ovule.  The  perianth,  stamens,  and 
style,  generally  wither,  because  the  part  they  play  is  at  an  end  ; 
the  wall  of  the  ovary  grows  and  becomes  the  wall  of  the  fruit 
(pericarp).  The  entire  gyncecium  of  a  flower,  transformed  as  a 
consequence  of  fertilisation,  is  termed  a  fruit.  It  consists  of  two 
parts,  the  pericarp  and  the  seeds,  and  according  to  the  nature  of 
the  pericarp,  the  fruit  is  termed  a  capsule,  nut,  berry,  or  drupe. 

The  chief  characteristic  of  the  Phanerogams  does  not  lie  in  the 
formation  of  the  flower  (although  they  may  quite  properly  be 
termed  "  Flowering-plants  "),  because  Equisetums  and  Lycopods 
have  reproductive  shoots  as  highly  differentiated  as  those  of 
certain  Gymnosperms  and  other  Phanerogams.  As  regards  the 
SEXUAL  GENERATION  the  characteristics  are  found : — (1)  in  its 
great  reduction  ;  (2)  in  the  transmission  of  the  microspore  (pollen- 
grain)  to  the  macrosporangium,  and  its  germination,  with  the 
formation  of  a  pollen-tube  (antheridium),  the  protoplasm  of  which 
is  not  differentiated  into  spermatozoids2;  (3)  in  the  fact  that  the 
macrospore  (embryo-sac)  never  leaves  its  sporangium  (nucellus)  ;. 
and  further  in  the  Angiosperms,  (4)  in  the  peculiar  development 
of  the  nutritive-tissue  in  two  parts ;  and  (5)  in  the  great  reduc- 
tion of  the  archegonium. 

As  regards  the  ASEXUAL  GENERATION  the  characteristic  feature  is- 
that  this  generation  is  formed  whilst  the  sporangium  is  still 
attached  to  the  mother-plant,  and  for  a  long  time  is  nourished  by 
it ;  and  that  after  the  sporangium  has  become  detached  from  the- 

1  .tiperaceos,  Nymph  aeaceas.         2  Except  in  Cycads  and  Ginkgo. 


250  CRYPTOGAMS   AND    PHANEROGAMS. 

mother-plant,  it  spends  a  longer  or  shorter  resting  period  as  the 
embryo  in  the  seed  (enveloped  by  the  testa),  and  does  not  make  its 
appearance  nntil  the  "germination  "  of  the  seed.  In  addition  the 
shoot  which  bears  sporangia  undergoes  greater  modification  than 
in  the  case  of  the  Flowerless-plants. 

The  Phanerogams  are  separated  into  two  Divisions  as  follows  : — 
Division  4.  Gymnospermae.  The  ovules,  as  well  as  the  seeds, 
are  borne  naked  on  the  surface  of  open  carpels,  or  on  the  apex  of 
a  stem  (ovary  wanting).  The  pollen-grains  are  conveyed  by  the 
wind  to  the  ovules,  and  caught  by  drops  of  mucilage,  secreted  by 
the  micropyle.  A  "stigma"  is  wanting.  The  entire  female  prothal- 
lium, (the  endosperm},  which  serves  for  the  nourishment  of  the 
embryo,  is  formed  before  fertilisation.  The  archegonia  are  em- 
bedded in  the  upper  part  of  the  prothallium.  The  pollen-grains  ar°. 
" multicellular"  i.e.  there  is  always  in  their  interior  a  distinct 
prothallium,  formed  by  1-3  cells,  and  a  larger  cell  which  gives 
rise  to  the  pollen-tube. 

Division  5.  Angiospermse.  The  carpels  surround  the  ovules 
and  form  an  entirely  closed  chamber  (ovary),  in  which  the  ovules 
mature  and  ripen  into  seeds.  The  surface  of  a  portion  of  the 
apex  of  the  carpel  is  transformed  into  the  "  stigma,"  which,  by  a 
sticky  fluid  and  also  by  hair- structures,  is  capable  of  retaining  the 
pollen-grains  conveyed  to  it  by  the  wind,  or  more  frequently  by 
insects.  The  pollen-tube  grows  from  the  stigma,  through  the 
"conducting  cellular  tissue"  (style),  to  the  ovules.  The  pollen- 
grains  contain  two  cells,  a  vegetative  and  a  free  generative  cell. 
The  latter  passes  into  the  pollen-tube  and  there  divides  into  two, 
one  of  which  is  the  sperm-nucleus.  The  female  prothallium, 
which  is  intended  to  serve  as  nutritive-tissue,  is  formed  after 
fertilisation.  Archegonia  are  wanting. 


DIVISION   IV. 

GYMNOSPERM^:. 

The  following  characters  should  be  added  to  those  already  given 
on  page  2  : — 

The  Gymnosperms  comprise  only  trees  or  shrubs.  The  flowers 
are  always  unisexual  and  destitute  of  perianth  (except  Gnetacece)  ; 
the  female  plant  of  Cycas  is  the  only  one  which  has  no  flower. 
The  MALE  FLOWERS  are  constructed  on  the  same  type  as  the  cones  of" 
the  Horsetails  and  Club-Mosses,  and  are  'most  frequently  long  shoots 
(Figs.  243,  258,  260  A,  267  J)  bearing  a  number  of  spiral  or  verti- 
cillate  stamens.  The  FEMALE  FLOWERS  are  of  a  more  varied  struc- 
ture (see  the  orders).  The  OVULE  is  orthotropous  (except  Podo- 
carpus  which  is  anatropous)  and  projects  from  the  carpel  up- 
rightly, inverted,  or  horizontally;  it  has  usually  only  one  integument 
(compare  however  Taxacese)  which  proceeds  from  the  upper  part 
of  the  nucellus,  so  that  the  embryo-sac  in  part  is  placed  below 
the  integuments  (Figs.  251,  264).  The  drop  of  mucilage  which 
catches  the  pollen-grains  dries  up  and  draws  the  pollen-grain 
through  the  micropyle  to  a  space  just  above  the  nucellus — the 
pollen-chamber — in  which  the  germination  of  the  pollen-grain 
commences. 

In  each  seed,  only  one  of  the  many  embryos  which  are  formed 
proceeds  to  its  full  development.  The  seed  is  always  endospermous> 
and  the  embryo  has  one,  two,  or  a  whorl  of  several  cotyledons. 
A  vigorous  primary  root  is  developed  on  germination.  THE 
VASCULAR  BUNDLES  in  the  stern  are  arranged  in  a  ring,  and  increase 
in  thickness  takes  place  by  a  closed  cambium-ring  which  forms 
bast  (phloem)  externally,  and  wood  (ocylem)  internally  with  distinct 
annual  rings,  as  in  the  Dicotyledons.  Only  certain  of  the  Cycadea3 
deviate  from  this  arrangement.  The  secondary  wood  is  very  uni- 
form, as  it  is  almost  exclusively  formed  of  trjacheides  with  bordered 
pits,  but  true  vessels  are  wanting ;  this  also  indicates  a  relationship 
with  the  Pteridophyta  (see  page  202). 

The  Grymnosperms  are  biologically  lower  than  the  Angiosperins ; 
they  are  wind-fertilised,  and  without  extra  floral-nectaries. 


252  GYMNOSPEKM.-T:. 

This  Division  embraces  three  classes:  CTCADEJB,  CONIFERS,  and 
GNETEJ;.  It  is  no  doubt  monophyletic,  and  has  taken  its  origin 
from  heterosporous  Ferns,  now  extinct,  most  nearly  related  to  the 
Ophioglossaceae  and  Marattiaceoe.  The  Cycadeee  appear  to  be  the 
oldest  class.  The  Conifers  are  related  to  these  through  Ginkgo. 
The  Gnetacea?  are  more  isolated.  The  Division  is  not  continued  into 
the  higher  Flowering-plants  ;  it  has  evidently  attained  its  highest 
development,  and  is  now  in  a  retrograde  condition.  The  similarity 
which  ha's  often  been  pointed  out  between  certain  Coniferse  and 
Lycopodinae  is  only  in  analogous  resemblances,  and  does  not  entitle 
one  to  suppose  that  there  is  a  nearer  relationship,  or  that  the 
former  take  their  origin  from  the  latter. 

Class  l.    Cycadeas. 

The  stem  is  very  rarely  ramified.  The  leaves  are  large,  pinnate, 
and  arranged  spirally.  The  flowers  are  dioeciotts,  without  perianth. 

There  is  only  one  order,  the  Cycadacese. — In  habit  they 
resemble  the  Ferns,  especially  the  Tree-Ferns  (compare  Figs.  207 
and  256).  The  stem  is  tubercular  (Fig.  258),  or  cylindrical  (Fig. 
256),  but  not  very  tall  (as  much  as  about  12  metres),  and  very 

itf 


FIG.  256. Cycis  circinalis  (female  plant).    The  carpels  are  seen  hanging  from  the  top  of 

the  stem.  Three  leaves  with  the  leaflets  still  rolled  up  project  almost  vertically  into  the 
air,  from  the  centre  of  the  crown. 

rarely  ramified.  [In  Ceylon,  unbranched  specimens  of  Cycas  are 
rarely  met  with  in  the  wild  state.  The  stems  of  C.  circinalis 
occasionally  branch  in  greenhouses.] 

The  LEAVES  are  arranged  spirally,  and  so  closely  together  that 
no  free  stem-surface  is  left  between  them,  and  have  only  a  slight 
sheath  (which  is  not  amplexicaul,  as  in  the  Palms) .  They  are  com- 


CVCADE^E. 


253 


pound  (most  frequently  pinnate ;  in  Boieenia, 
bipinnate)  ;  in  some  genera  the  leaves  are 
rolled  up  in  various  ways,  resembling  the 
vernation  in  Ferns  (Fig.  257)  ;  they  are 
leathery  and  perennial.  In  some,  stipules  are 
present,  as  in  the  Marattiaceae.  Groups  of 
scale-leaves  alternate  in  the  majority  with 
groups  of  foliage-leaves. 

The  FLOWERS  are  without  perianth.  The 
MALE  FLOWER  is  sometimes  an  enormous  collec- 
tion of  stamens  (Fig.  258),  which  are  flat  in 
some  (Cycas,  Fig.  241),  shield-like  in  others 
(Zamia,  Ceratozamia)  like  the  sporophylls  in 
Horsetail  (Fig.'  259)  ;  but  in  all,  the  pollen- 
nacs  are  situated  in  large  and  varying  numbers  on  the  back  of 
the  stamens,  and  arranged  in  groups  of  2-5,  like  the  sporangia  in 
the  sori  of  the  Ferns  (Fig.  241  5,  c).  On  germination  the  micro- 


Fio.  257.  — Ci/cas  cir- 
cinilis.  Part  of  a  young 
leaf  with  circinate 
leaflets. 


FIG.  2-~8.— A  male  plant  of  Stungeria  paradoxa  (about 
j|  nat.  size). 


FTG.  259. — Female  cone 
of  Zamia  integrifolia  (\-$ 
nn.t.  size).  The  male  cone 
is  very  similar  externally. 

spore  contains  a  bicellular  antheridium  in  the  upper  end  of 
the  pollen-tube?  the  vegetative  nucleus  being  found  near  its 
lower  end.  The  inner  of  these  two  cells  is  the  antheridial  cell ; 
this  divides  and  forms  two  spermatozoids,  with  cilia  at  the 
anterior  end,  which  swim  about  in  the  pollen- chamber,  and 
finally  penetrate  the  archegonium.  FEMALE  FLOWERS  are  wanting 
in  Cycas,  because  the  carpels  do  not  terminate  the  apical  growth  of 
the  stem.  After  a  group  of  foliage-  and  of  scale-leaves,  a  group  of 
carpels  is  developed,  which  are  pinnate  and  resemble  the  foliage- 
leaves,  bearing  on  their  edges  a  number  of  ovules  (most  fre- 


254  GYMNOSPERMJ:. 

quently  5-6)  (Figs.  245,  256)  ;  the  same  stem  produces  succes- 
sively scale-leaves,  foliage-leaves,  and  carpels.  The  differentiation 
is  not  much  more  advanced  than  in  certain  Ferns  (Struthiopteris, 
Blechnuni),  where  barren  and  fertile  leaves  of  different  form 
regularly  alternate.  The  other  genera  have  female  flowers;  the 
carpels  are  shield-like  in  Zamia  and  Ceratozamia  (Fig.  246), 
and  collected  into  cone-like  flowers,  which  terminate  the  growth 
of  the  stem  (Fig.  259).  The  number  of  ovules  in  these  instances 
is  two  to  each  carpel. 

The  SEEDS  are  large  (most  frequently  2-6  centimetres  long)  and 
plum-like  ;  the  external  layer  of  the  testa  is  fleshy,  while  the  in- 
ternal one  is  hard  and  horny.  There  are  two  systems  of  vascular 
bundles  in  the  testa,  one  outside,  the  other  inside  the  stone.  The 
embryo  is  straight,  attached  to  the  end  of  the  suspensor,  which  is 
often  long,  filamentous,  and  rolled  up ;  it  has  one  or  two  cotyledons. 

The  embryo  in  Ceratozamia  and  others  is  very  slightly  developed,  at  the  time 
when  the  ripe  seed  is  detached  from  the  carpel ;  and  it  is  not  until  after  sowing 
that  its  further  development  and  germination  proceed.  This  calls  to  mind 
the  Cryptogams,  especially  Selaginella,  whose  macrospores  are  thrown  off  filled, 
•with  endosperm ;  but  the  oosphere  is  not  fertilised  till  after  the  separation  of  the 
macrospore  from  the  parent-plant,  while  in  the  Cycadeaa  fertilisation  is  effected 
before  the  separation.  In  Cycas  the  testa  may  rupture,  and  the  endosperm 
grow  and  become  green  in  the  light,  even  though  no  embryo  has  been  formed. 
This  also  is  an  indication  of  its  prothalloid  nature. 

Gum-passages  are  present  in  all  organs.  Collateral  vascular  bundles,  with 
spiral  and  scalariform  tracheides,  are  found  ;  and  normal  thickening  takes  place 
by  means  of  a  cambium.  An  exceptional  mode  of  growth  is  found  in  Cijcus  and 
Encephalartos,  the  cambium  ceases  to  divide  after  a  time  and  is  replaced  by  a 
new  cambium  which  arises  in  the  cortical  parenchyma  just  outside  the  bast,  and 
which  forms  a  new  ring  of  xylem  and  phloem.  This  may  be  repeated  so  that  a 
number  of  concentric  rings  are  produced.  In  Ceratozamia,  structures  resem- 
bling corals  extend  from  the  roots  in  a  vertical  direction  and  appear  on  the 
surface  of  the  soil;  these  are  peculiar  roots,  in  which  a  symbiotic  Alga 
(Anal/ana)  is  found. 

The  Cycadete  were  formerly  (from  the  Coal  period  to  the  Later  Cretaceous) 
far  more  numerous  than  at  the  present  day.  They  appear  to  have  been  most 
numerous  in  the  Trias  and  Jurassic.  The  remnant  (75  species)  which  have 
persisted  to  the  present  time  are  found  in  all  tropical  countries.  Cycas  (Trop. 
and  Sub-trop.,  Eastern  Hemisphere)  ;  Dioon  (Mexico)  ;  Macrozamia  (Australia) ; 
EncepJialartos  (Trop."  and  S.  Africa)  ;  Stangeria  (Fig.  258,  Sub-imp.  South  and 
East  Africa) ;  Bowenia  (Trop.  Australia) ;  Ceratozamia  (Mexico,  New  Granada, 
Western  Brazil)  ;  Microcycas  (Cuba) ;  Zamia  (Trop.  and  Sub-trop.  N.  America.) 

USES.  Sago  is  made  from  the  starch-containing  pith  of  Cycas  revoluta  and 
circinalis.  The  leaves  are  often  used  at  funerals  and  church  festivals,  under 
the  name  of  "  palm-branches." 


CONIFEEJE.  255 

Class  2.    Coniferae  (Pine-trees). 

The  stem  branches  freely.  The  leaves  are  entire,  relatively 
small,  linear  or  reduced  to  scales.  The  flowers  are  without  peri- 
anth. The  ovules  naked.  It  is  seldom  that  the  female  flower  is 
reduced  to  only  one  carpel. 

Whilst  the  Cycadese  principally  resemble  the  Ferns,  the  Conifers 
partly  resemble  the  Lycopods,  and  partly  the  Equisetums — the 
former  especially  in  the  needle-  or  scale-like,  leathery,  simple,  and 
often  perennial  leaves  ("  evergreen  plants  "),  which  never  possess 
stipules  (Figs.  263,  270,  272).  Ginkgo  deviates  from  this,  being 
no  doubt  the  oldest,  and  the  Conifer  which  stands  nearest  to  the 
Cycadea3  (Fig.  260).  The  resemblance  to  the  Equisetums  is 
especially  owing  to  the  fact  that  the  stem  ramifies  abundantly,  and 
often  very  regularly,  forming  a  pyramid  with  verticillate  branches. 
In  addition-  to  the  foliage-leaves,  scale-leaves  (bud-scales)  are 
present  in  the  majority  of  species. 

-The  FLOWERS  are  monoecious  or  more  rarely  dioecious.  Perianth 
is  wanting.  The  stamens  of  the  catkin-like  male  flowers  (Fig.  267, 
J)  are  of  different  forms,  but  as  a  rule  more  or  less  shield-like. 
As  in  the  Cycadese,  the  pollen-sacs  are  in  all  cases  situated  on  the 
underside.  There  are,  as  a  rule,  two  pollen-sacs  (the  Abietacea3, 
Fig.  267),  or  3-5,  (the  Cupressacese  and  Taxaceae,  Fig.  243) ;  a 
few  have  more,  e.g.  Araucaria  (Fig.  242)  ;  they  dehisce  by  clefts. 

If,  in  commencing  our  consideration  of  the  female  /lower,  we 
begin  with  that  of  Ginkgo,  we  shall  observe  in  the  corner  of  a  scale- 
or  foliage-leaf  a  small  flower,  which  consists  of  two  carpels,  each 
bearing  one  ovule,  and  reduced  almost  to  the  ovule  itself  (Fig.  260 
C,  D).  The  flower  in  Podocarpus  is  still  further  reduced,  viz.  to  a 
single  carpel  with  one  ovule,  which  is  anatropous  and  has  two  in- 
teguments. This  ovule  is  situated  in  the  axil  of  a  cover-scale  (c,  in 
Fig.  262  D),  and  several  female  flowers  of  this  description  are  col- 
lected in  a  small  cone,  the  stalk  and  bracts  of  which  become  fleshy 
(Fig.  262  C).  The  external  integument  also  becomes  fleshy  (an  aril). 
Dacrydium,  which  is  clearly  related  to  Podocarpus,  has  an  external 
integument  which  developes  more  independently  as  a  fleshy  aril 
(Fig.  262  B,  J51).  Microcachrys  also  is  clearly  allied  to  these : 
the  bracts  are  more  fleshy,  and  the  ovule  (i.e.  the  female  flower) 
is  protruded  beyond  the  bract  (Fig.  262  A,  A1).  Taxus  stands 
in  a  more  isolated  position :  a  flower  which  has  been  reduced  to 

an  ovule  is  situated,  in  this  instance,  on  the  apex  of  a  secondary 
W.B.  S 


256  GYMNOSPERMJ;. 

branch  which  is  studded  with  floral-leaves  (Figs.  263,  264)  ;  an  ex- 
ternal integument  is  developed  on  all  sides  and  surrounds  the  seed 
as  a  scarlet  aril.  According  to  this  conception  the  aril  corresponds 
to  an  external  integument,  and  the  Taxoideas  thus  possess  a  partly 
dichlamydeous  ovule.  Only  Ginkgo  and  Cephalotaxus  appear  to 
deviate  from  this,  as  in  these  there  is  only  one  integument  (unless 
the  small  outgrowth  indicated  by  ar,  in  Fig.  260  D,  really  is  a 
rudimentary,  external  integument)  ;  in  CYCADE^,  to  which  Ginkgo 
is  most  closely  related,  there  is  likewise  only  one  integument.  But 
in  these  genera  the  testa  is  differentiated  into  two  layers,  and  the 
seed  resembles  a  drupe ;  like  the  Cycadeae  there  is  an  external 
fleshy  covering  and  an  internal  hard  one,  and  these  two  layers 
may  probably  be  considered  homologous  with  the  two  integuments. 
This  theory  is  also  borne  out  by  the  arrangement  of  the  vascular 
bundles  in  Cephalotaxus  and  Podocarpus,  which  present  the  xylem 
in  the  fleshy  external  layer  to  the  outside  of  the  testa,  which  is 
therefore  the  upper  side  of  the  integument  (Celakovsky). 

The  coalescence  of  the  integuments  into  one  is  only  slight- in 
Torreya,  more  pronounced  in  Podocarpus  and  strongest  in  Cephalo- 
taxus and  Ginkgo.  Celakovsky  terms  these  ovules  "  holochlamy- 
deous." 

If  we  pass  from  these  to  the  order  PINOIDE.E,  we  find  the  female 
flowers  collected  into  catkin-like  cones,  which  have  been  considered 
from  various  points  of  view  to  be  sometimes  single  flowers,  at 
other  times  compound  inflorescences.  The  structure  in  ABIETACEJ; 
is  as  follows :  a  number  of  spirally  arranged,  scale-like  leaves, 
cover-scales  (Figs.  267,  268),  are  situated  on  a  long  axis.  In  the 
axil  of  each  cover- scale  a  larger  leaf -like  projection,  the  ovuliferous 
scale,  is  borne,  which  turns  the  upper  side  towards  its  cover-scale 
(which  is  shown  by  the  fact  that  the  wood  of  its  vascular 
bundles  is  turned  downwards  and  towards  the  wood  in  the  bundles 
of  the  cover-scale  :  Fig.  269).  Two  ovules,  with  micro pyles  turned 
towards  the  central  axis,  and  with  apparently  only  one  integument 
(Fig.  268),  are  situated  on  the  dorsal  side  of  each  ovuliferous 
scale,  i.e.  the  side  turned  away  from  the  cover-scale.  The  ovuli- 
ferous scales  grow  after  fertilisation,  into  the  woody  or  leathery 
"cone-scales,"  which  are  usually  much  larger  than  the  cover-scales. 
This  ovuliferous  scale  with  its  axis  may,  according  to  Celakovsky, 
be  considered  as  a  dwarf-branch  which  is  situated  in  the  axil  of 
the  cover-scale,  and  bears  two  ovules  (in  the  same  way  as  in 
Ginkgo,  one  long-stalked  flower,  reduced  to  two  ovules,  is  situated 


CONIFERJE.  257 

in  the  axil  of  a  leaf),  And  in  this  case  the  external  integument  of  the 
ovules  is  expanded  into  leaf-like  bodies,  which  have  united  to  form 
one  "  symphyllodium  "  (ovuliferous  scale)  which  is  inverted  so  that 
its  dorsal  side  is  turned  upwards  and  bears  the  nucellus  and 
the  other  integument  ("  hemichlamydeous  "  ovules).  The  carpel 
itself  is  therefore  in  this  instance  extremely  reduced.  The  keel, 
or  (in  Pinus)  "  mucro"  (Fig.  268  jB),  which  is  found  in  several 
genera,  represents  then  a  third  carpel,  which  is  sterile.  In  the 
other  orders  of  the  PinoideaB  the  cover-scales  and  ovuliferous  scales 
grow  more  and  more  together  and  finally  form  one  structure,  which 
also  is  termed  a  "  cone-scale,"  although  from  its  development  it 
cannot  be  homologous  with  the  cone-scales  of  the  Abietaceas. 
This  connation  is  least  in  the  TAXODIACEJ  and  ARAUCARIACE.E  and 
may  be  traced  on  the  upper  surface  of  the  "  cone-scale  "  by  the 
presence  of  a  stronger  or  slighter  ridge  or  pad,  the  free  portion 
of  the  ovuliferous  scale  (Figs.  256,  266,  269).  It  is  most  strongly 
pronounced  in  the  CUPRESSACEJ;,  in  which  the  two  scales  form  one 
single  structure,  the  cone-scale  (Fig.  274).  The  vascular  bundles 
in  the  under  portion  corresponding  to  the  cover-scale,  have  the 
xylem  towards  the  upper  side  as  usual  in  leaves,  whilst  the  bundles 
present  in  the  upper  side  of  the  cone-scale,  which  thus  represents 
the  ovuliferons  scale,  turn  their  xylem  downwards.  The  hemi- 
chlamydeous ovules  are  then  situated  on  the  upper  side  of  this 
cone-scale.  According  to  this  theory  the  CUPRESSACEJ  appear  to  be 
the  youngest  type,  a  view  which  corresponds  with  their  vegetative 
structure.  If  there  is  only  one  ovule  in  these  orders  as  in  Agathis 
(Fig.  265)  and  Araucaria,  then  the  flower  is  reduced  to  a  single 
carpel  and  one  ovule,  as  in  the  case  of  Dacrydium  and  Microcachrys. 
If  two  or  more  ovules  are  present,  then  the  same  number  of  carpels 
may  be  supposed  to  exist,  the  external  integuments  of  their 
ovules  being  developed  into  leaf-like  structures  which  collaterally 
coalesce  to  form  a  "  symphyllodium,"  or  are  suppressed. 

According  to  this  theory,  which  is  based  on  the  researches  of 
Celakovsky,  the  female  flowers  of  the  Coniferte  may  be  classed 
thus  : — 

1.  In  all  cases  situated  in  the  axil  of  a  bract  and  collected  into 
cones,  with  numerous  flowers  or  with  few  or  one  flower.     In  Ginkgo 
only,  are  they  situated  in  the  axil  of  foliage-  or  scale-leaves. 

2.  It  is  only  in  Taxus  that  bracteoles  are  present. 

3.  They  are  formed  only  from  rudimentary  carpels,  in  which  the 
stem  takes  no  part. 


258  GTMNOSPEBMJ:. 

4.  The   number  of  carpels  in  each  flower  varies   from   one   to 
many,  most  frequently  three,  of  which  the  central  one  remains 
sterile. 

5.  Each  carpel   bears    only  one    ovule.       The  flower  which  is 
formed  of  only  one  carpel  appears  to  consist  of  only  one  ovule. 

6.  The    ovule    has    in    Taxaceae    either    a   double    integument 
(Podocarpese,  Taxeae),  of  which  the  external  is  the   "  aril,"  or,  as 
in  the  Cycadeae,  a  single  one,  which  is  homologous   with  the  two 
united  together. 

7.  The   external   integument   in  the  Pinoideae  is   expanded  to 
form  a  leaf -like  structure — the  ovuliferous  scale — and  bears  on  its 
dorsal  side  the  ovules,  which  are  thus  only  provided  with  one,  and 
that  the  inner,  integument. 


This  later  interpretation  of  the  female  cones  in  the  Conifera  is  more  pro- 
bably correct  than  the  older  ones  ;  that,  however,  which  appeared  in  the  former 
issues  of  this  book,  may  also  be  stated.  It  was  to  the  effect  that  each  catkm- 
like  female  cone  is  in  reality  a  single  flower  ;  the  cone-scales  in  the  Cupressacefe 
were  single  leaves,  namely  carpels,  which  bore  the  ovules  on  the  side  which  is* 
turned  upwards;  the  division  into  two  parts  which  makes  its  appearance  in  the 
other  orders,  and  becomes  most  prominent  in  the  Abietaceae,  was  compared 
with  the  division  of  a  leaf  into  a  barren  and  a  fertile  portion,  which  is  fouudi 
especially  in  Ophioglossacese  and  Marsiliacese,  or  with  the  ligule  in  Isoetes. 


POLLINATION  is  accomplished  by  means  of  the  wind.  At  the- 
period  of  pollination  the  leaves  are  always  so  widely  separated 
from  one  another,  that  the  ovules  can  catch  the  pollen-grains- 
carried  to  them  by  the  wind  ;  this  is  often  effected  by  the  mucila- 
ginous drops  which  appear  at  the  micropyle,  and  by  the  evapo- 
ration of  which  the  pollen-grains  are  brought  in  contact  with 
the  nucellus.  The  entire  cone  grows  considerably  as  soon  as- 
fertilisation  has  taken  place,  and  the  cone-scales  in  Pinoidese 
close  together  so  that  the  seeds  while  maturing  are  enclosed,  and 
it  is  not  until  the  seeds  are  ready  for  distribution  that  the  cone- 
scales  again  become  separated.  In  the  Pinoideay,  the  fully  deve- 
loped ovuliferous  scales  are  hard  and  woody ;  and  in  this  condition 
the  collection  of  female  flowers  is  termed  a  cone.  In  the  Taxoidese, 
true  cones  are  the  exception.  2-15  cotyledons  are  present,  ar- 
ranged in  a  whorl. 

The  characteristic  feature  of  this  class  is  the  abundance  of 
resin,  which  is  to  be  found  in  isolated  cells  (especially  in  ther 


CONIFER*.  259 

cortex),  partly  in  intercellular  glands  or  passages  (both  in   the 
cortex  and  wood).     Taxus  is  the  only  genus  which  has  no  resin. 

There  are  about  350  species,  mostly  from  the  Northern  Temperate  zone 
(especially  North  America  and  Siberia),  where  they  grow  gregariously  and  form 
the  most  northern  forests.  The  Juniper,  Scotch  Fir,  and  Yew  are  natives  of 
Great  Britain. 

This  class  may  be  divided  into  two  families  : — 

1.  Taxoideae.     The  ovules  have   either  one  integument,  the 
external  part  of  which  is  fleshy,  and  the  internal  hard  and  stone- 
like  ;  or  two  integuments,  of  which  the  external  is  the  fleshy  and 
coloured  "aril."     "Ovuliferous  scales"  are  wanting.     The  cones 
are  never  woody,  but  are  generally  sacculent,  the  bracts  become 
fleshy,  or  cones  usually  are  not  developed.     The  seeds  project  more 
or  less  freely  beyond  the  bracts. 

2.  Pinoideae.     The  ovules  have  two  integuments,  the  external 
one  of  which  is  leaf-like  and  becomes   developed  as  the   "  ovuli- 
ferous  scale  "  ;  if  there  are  several   of  these  in  each  flower  they 
unite  and  form   a  "  symphyllodium."     This  may  remain  free  or 
unite  with  the    bract.     The    cones    are    most  frequently  woody, 
rarely  succulent.     The  seeds  are  hidden  among  the  cone-scales. 

Family  1.     Taxoideae. 

This  family,  considered  to  be  most  nearly  related  to  the  Cycadeee, 
also  made  its  appearance  at  a  very  early  period.  There  is  only 
one  order.  [See  note  on  page  272.] 

Order.     Taxaceae.     The  characters  have  been  given  above. 

A.  CEPHALOTAXEJ:  is  the  oldest  group,  presumably  the  connect- 
ing link  between  the  Cycadeae  and  the  other  Coniferaa.  The  flower 
consists  normally  of  two  ovules.  Aril  wanting.  One  integument. 
Seeds  drupaceous. — The  flowers  in  Ginkgo  biloba  (Salisburia} 
are  situated  in  the  axil  of  foliage-  or  scale-leaves.  The  stamens 
bear  only  two  pollen-sacs  (Fig.  260  A).  The  female  flower  has 
two  ovules,  placed  together  at  the  end  of  a  long,  bare  axis  (Fig. 
260  (7) .  Round  the  base  of  the  ovule  a  small  collar  (ar,  in  Fig. 
260  D)  is  found,  which  may  probably  be  considered  homologous 
with  the  collar-like  outgrowth  which  surrounds  the  base  of  the 
Cycas- ovule.  The  seed  resembles  a  Plum,  and  has  a  fleshy  external 
coat,  surrounding  a  hard  internal  layer.  The  embryo  is  developed 
after  the  seed  has  fallen  off.  The  Ginkgo- tree  has  long-stalked, 
fan- shaped  leaves,  more  or  less  indented,  with  dichotomous  veins 


260 


GYMNOSPERM.S. 


resembling  certain  Ferns — the  Adiantums.     It  is  a  native  of  East 
Asia,  and  the  only  surviving  species  of  a  genus  which  in  earlier 


Fio.  260. — Ginlcgo  (nat.  size)  :  A  a  branch  with  a  small  flowering  dwarf-branch  (male 
flower);  £  a  leaf;  C  a  flower  with  two  ovules ;  Da  ripe  seed;  ar  collar. 

times   was  very  rich   in  species,  and   distributed  over  the  entire 
Northern  Hemisphere.    Cephalotaxus  (Eastern  Asia)  is  related  to  it. 
B.  PODOCARPE^E.      The  female  flower  is  reduced    to  one   ovule, 
placed  in  the  axil  of  a  bract,  or  a  little  forward  upon  it.     The  ovule 
has   an  aril    (2  integuments). — Phyllocladus 
(Fig.  261),  from  New  Zealand  and  Tasmania, 
has  obtained  its  name  from  its  flat,  leaf-like 
branches,  the  leaves  proper  being   scale-like 
(/).     The  ovules   stand  erect  in  the  axil  of 
the    scale-like   leaves    (c),    and    several    are 
collected   at  the   end    of   short    branches. — 
Microcachrys     tetragona     (Tasmania)     has    a 
small    female    catkin   with  several   spirally- 
placed,  fleshy  bracts,   at  the  end  of  which 
the    inward  and  downward  turned  ovule  is 
attached  (Fig.  262  A,  A').     The  ripe  cones 
are    red,    succulent,    and    resemble    Straw^ 
berries.  —  In     Dacrydium     (Tasmania,    New 
FIG.  201.  -  Phyllocladus      Zea]aild  Malaysia)  the  female  cone  has  most 

glaucus  •.    a  branch    with  ,  •'         ' 

female  flowers  (nat.  size).      frequently  only  1-2   (-6)  bracts,  which   re- 


CONIFERS. 


261 


semble  the  vegetative  leaves  ;  they  have  also  a  fleshy  aril  (Fig. 
262  1»,  J3'). — Podocarpus  (40  species,  East  Asia,  S.  Temp.)  ;  the 
bracts  of  the  female  flowers  become  fleshy,  and  unite  together ; 
only  1  or  2  are  of  use  in  supporting  the  flowers.  The  ovules 


A 


FIG.  2G2.— A  Microcachrys  :  female  cone  (f).  A'  A  single  carpel  with  its  ovule.  J?" 
Dacrydium  :  branch  with  female  flower  (T).  B'  The  flower;  cp  the  bract;  ar  the  aril;  ov 
ovule.  C  Podocarpus  :  female  flower  with  2  ovules.  D  Another  female  flower  with  1  ovule, 
in  longitudinal  section. 

project  high  above  the  apex  of  the  bract,  and  are  anatropous,  the 
micro pyle  being  turned  downwards  (Fig.  262  C,  D}.  An  aril 
commences  to  develope  in  the  flowering  period  as  an  external 
coating,  and  later  on  it  becomes  fleshy  and  coloured. 

C.  TAXEJ:.  The  female  flower  is  reduced  to  one  ovule,  which  is. 
situated  terminally  on  an  axis  which  bears  2-3  pairs  of  opposite, 
scale-like  bracteoles  ;  on  this  account  the  Taxea?  form  a  very 
isolated  group  among  the  Conifer®. 
—Taxus  (T.  baccata,  the  Yew-tree). 
Dioecious.  The  female  flower  con- 
sists of  only  one  ovule,  placed  at  the 
end  of  a  short  secondary  branch 
(Fig.  264),  which  is  studded  with 
scale-like  leaves.  The  aril  when 
ripe  is  thick,  fleshy,  and  scarlet 
(sometimes  yellow),  and  only 
loosely  envelopes  the  seed  (Fig. 
263).  The  leaves  are  scattered, 

FIG.   263.  —  Taxus  baccata  :  branch 

flat,  linear,  and  pointed  (Fig.  263,  with  two  ripe  seeds  (nat.  size). 


262 


GYMNOSPEms. 


264).  The  short  male  flowers  have  5-8  pollen-sacs,  pendent  from 
the  stamens,  and  are  surrounded  at  their  bases  by  scale-like 
bracteoles  (Fig.  243).  Torreya  (4  species,  K  America  and  Japan) 
is  closely  allied  to  Taxus.  The  aril  ultimately  fuses  with  the 
woody  inner  integument,  and  hence  the  ovule  becomes  drupaceous, 
as  in  Cephalotaxese. 


Fis.  2Gi.— Tan-.us  Ituccata :  A  shoot  of  Taxus  with  female  flowers  at  the  time  when  tho 
ovules  are  ready  for  pollination.  B  Leaf  with  flower  in  its  axil  (nat.  size).  C  Longitudinal 
median  section  through  a  female  shoot ;  i»  growing  point  of  primary  shoot ;  a  commence- 
ment of  aril;  i  integument;  wnucellus;  m  micropyle. 

USES.  Taxus  baccata  is  usually  planted  in  gardens,  especially  in  hedges. 
Its  wood  is  very  hard  and  is  used  for  wood-carving.  The  shoots  are  poisonous, 
but  not  the  aril,  which  is  often  eaten  by  children  and  by  birds. 

Family  2.     Pinoideae. 

The  four  orders  differ  from  one  another  partly  in  the  arrange- 
ment of  tho  leaves  (Cupressacece  have  opposite  or  verticillately 


CONIFERS.  263 

placed  leaves,  flowers,  and  inflorescences  ;  in  the  others  they  are 
placed  spirally),  but  chiefly  in  the  greater  or  less  degree  of  union 
which  takes  place  between  the  female  flower  (the  leaf -like  "  sym- 
phyllodiurn")  and  its  supporting  cover-scale,  and  in  the  position  of 
the  ovules  (the  micropyle  being  turned  upwards  or  downwards). 
The  "  cone-scales "  in  Abietacece  are  formed  by  "  symphyllodia  " 
alone,  in  the  others  by  their  union  with  the  cover-scale. 

Order  I.  Araucariaceae.  This  order  most  frequently  has 
solitary  ovules,  turned  downwards  and  attached  to  the  centre  of  the 
cone-scales.  In  Agathis  (Dammara}  the  arrangement  is  the  most 
simple,  a  winged  seed  (Fig.  265),  which  hangs  freely  downwards, 
being  borne  in  the  centre  of  the  undivided  cone-scale.  In  Arau- 
caria,  the  stamens  with  the  free,  pendulous  pollen-sacs  have  been 
represented  in  Fig.  242  ;  the  ovuliferous  scale  is  united  for  nearly 


VA 

265.  266. 

FIG-.  265. — AgatMs  (Dammara)  australis.  Cone-scale  with  the  seed.  A  Longitudinal 
section  ;  A'  from  within  ;  fo,  fv'  vascular  bundles  ;  v  wing. 

FIG.  266. — Cunninghamia  sinensis.  Cone-scale  with  three  ovules,  interior  view :  d  cover- 
scale  ;  /  ovuliferous  scale. 

its  whole  length  with  the  bract,  and  projects  from  its  apex  in  the 
shape  of  a  sheath-like,  dentate  scale,  resembling  the  ligule  in 
Isoetes,  and  may  therefore  be  termed  a  "  ligule."  Araucaria  (S. 
America,  Australia)  has  often  rather  broad  leaves  (A.  brasiliensis) . 
The  ovuliferous  scale  in  Cunninghamia  is  more  distinct,  and 
stretches  transversely  over  the  entire  cover-scale ;  it  bears  three 
inverted  ovules  (Fig.  266)  (Eastern  Asia). 

Dammara -resin,  which  is  used  for  varnish,  is  obtained  from  Agathis 
(Dammara)  species  (New  Zealand,  Philippine  Islands). 

Order  2.  Abietaceae  (Pine  and  Fir  Trees).  The  leaves 
are  spirally  arranged  and  needle-like.  The  flowers  are  monoecious. 


264 


GTMNOSPEEMJi. 


The  male  flowers  are  long,  and  catkin-like,  with  numerous  stamens, 
each  bearing  two  oblong  pollen-sacs.  The  pollen-grains  are  most 
frequently  tri-lobed,  having  two  bladder-like  appendages,  formed 
as  outgrowths  of  the  exospore,  to  assist  in  their  distribution  by  the 
wind  (Fig.  267  N~).  The  bracts  are  arranged  spirally.  The  union 
between  the  bract  and  the  ovuliferous  scale,  which  is  found  in  the 
preceding  order,  is  not  in  this  instance  so  complete ;  these  scales 
make  their  appearance  as  two  free  parts,  and  are  attached  only 
at  their  bases  (Fig.  268) ;  the  lower  portion,  that  is  the  cover- 
scale,  in  most  instances  remains  quite  small  (Fir,  Red  Pine,  and 
others),  it  is  only  in  the  "Noble  Pine  "  (Abies)  and  Pseudotsuga 


FIG.  207. — A-G  Pseudotsuga  douglasii :  A  cone,  B  cone-scale,  with  the  inner  side 
turned  forward  ;  the  points  of  the  cover-scale  are  seen  behind  it;  C-G  transitions  from 
the  acicular  leaf  to  the  cover  scale,  from  the  base  of  a?  cone.  If  Pinus  montana.  Young 
ovuliferous  scale,  with  the  inner  side  turned  forward ;  the  ovules  are  now  in  the  stage  for 
pollination.  J-M  Abies  alba  :  J  male  cone  ;  b  bud-scale ;  a  anthers  •  K  LM  individual 
anthers.— Pinus  montana:  N  pollen-grain;  the  two  lateral  expansions  are  the  aii- 
bladders ;  in  the  upper  part  of  the  interior  of  the  grain  a  vegetative  cell  may  be  seen, 
and  in  the  centre  the  large  cell-nucleus. 


m,  that  it  attains  a  greater  length  than  the  ovuliferous 
scale  (Fig.  267,  B-G).  On  the  other  hand  the  upper  part,  the 
ovuliferous  scale  (the  vascular  bundles  of  which  have  the  bast 
turned  upwards),  grows  strongly  and  elongates,  especially  after 
fertilisation,  becoming  woody  or  leathery  ;  it  is  commonly  termed 
the  "  cone-scale"  but  is  in  reality  only  homologous  with  a  part  of 


CONIFERS.  265 

the  "  cone-scale"  in  the  other  order  of  Pinoideae.  On  the  side 
of  the  ovuliferous  scale,  turned  towards  the  axis,  are  situated  two- 
ovules  with  micropyles  directed  inwards.  The  seeds  are  most 
frequently  provided  with  a  false  wing  (a  tissue-like  part  of  the 
surface  of  the  ovuliferous  scale).  Cotyledons,  more  than  2, 
verticillate.  Fertilisation  does  not  take  place  until  some  time  after 
pollination.  In  Pinus,  for  instance,  the  pollen-tube  only  penetrates 
the  nucellus  for  a  short  distance  during  the  year  of  pollination,  and 
then  ceases  its  further  growth,  fertilisation  not  taking  place  until 
after  the  middle  of  the  next  year;  whilst  the  seeds  ripen  about  a 


Fie.  2GS.— A  AUcs:  c  the  cover-scale  ;  s  ovnliferous  scale,  or  "cone-scale";  sk  ovules  in 
a  young  condition.  B  Pinus  :  ovuliferous  scale  with  two  ovules  (s) ;  m  the  two-lobed 
micropyle;  c  "  mucro  "  ;  b  the  cover-scale  behind.  C  Abies  :  ripe  "  coue-scale  "  with  two 
seeds  (su) ;  /  wing  of  seed. 

yenr  and  a  half  after  pollination.     In  the  Larch   and   others,  the 
seeds  are  mature  in  the  autumn  succeeding  pollination. 

Abies  (Fir).  The  leaves  are  often  (e.g.  Ab.  pectiriata)  displaced 
into  2  rows,  flat  and  indented  at  the  apex,  with  2  white  (wax- 
covered)  lines  on  the  under  surface,  in  which  the  stomata  are  situ- 
ated. The  leaf-scars  are  nearly  circular  and  do  not  project.  The 
cones  are  erect.  The  cover-scales  and  the  ovuliferous  scales  separate 
from  the  axis,  to  which  they  remain  attached  in  other  genera. — 
Tsuga  has  leaves  like  Abies,  but  by  the  slightly  projecting  leaf-scars,  and  cones 
•with  persistent  scales,  it  forms  the  transition  to  Picea. — PseucLotsuga  has  leaves 
similar  to  those  of  Abies  and  persistent  carpels  as  in  Picea,  but  the  cover-scales 
grow  as  in  Abies  and  project  beyond  the  ovuliferous  scales  (P.  doitglasii,  Fig. 
267).  These  two  genera  are  considered  as  sub-genera  of  Abies. — Picea.  The 
leaves  project  on  all  sides,  square  and  pointed  ;  the  leaf-scars  are 


266  GYMNOSPERM^l. 

rhombic,  on  projecting  leaf-cushions.  The  cones  are  pendulous. 
The  cover-scales  are  much,  shorter  than  the  leathery,  persisting 
ovuliferous  scales. — The  genus  Larix  (Larch)  differs  from  all  the 
others  in  having  deciduous  leaves  (the  three  preceding  have 
leaves  which  persist  for  eleven  to  twelve  years).  It  has  long- 
branches  with  linear  foliage-leaves  and  short,  thick,  perennial 
dwarf -branches,  which  each  year  form  a  new  rosette  of  foliage- 
leaves,  similar  to  those  on  the  long-branches.  The  male  flowers 
and  the  erect  cones  resemble  those  of  Picea,  and  are  borne  on 
dwarf-branches. — Cedrus  (Cedar)  resembles  Larix  to  some  extent, 
but  has  persistent  leaves  (C,  libani,  C.  deodara). — Pinus  (Pine)  has 
long-branches  and  dwarf-branches.  The  leaves  of  the  long- 
branches  are  scale-like  and  not  green;  the  dwarf-branches  have 
very  limited  growth,  and  persist  for  three  years;  they  arise  in 
the  axils  of  the  scales  borne  on  the  long-branches  of  the  self-, 
same  year,  and  each  bears  2-5  foliage-leaves,  they  are  also  sur- 
rounded at  the  base  by  a  number  of  membranous  bud-scales. 
The  cone-scales  have  a  thick,  rhomboid  extremity  (the  u  shield  "J. 

The  buds  which  develope  into  long-branches  arise  at  the  apex  of  other  long- 
branches,  and  being  very  close  together,  form  false  whorls.  The  female  cones 
occupy  the  position  of  long-branches,  and  take  about  two  years  for  their  develop- 
ment. The  male  flowers  arise  close  together,  and  form  a  spike-like  inflorescence 
at  the  base  of  a  long-branch  of  the  same  year.  The  male  flowers  occupy  the 
position  of  dwarf-branches,  so  that  a  female  cone  may  be  considered  to  be  a 
modified  long-branch,  and  a  male  cone  a  modified  dwarf-branch.  The  main 
axis  of  the  seedling  has  needle-like  leaves,  similar  to  those  found  on  the  older 
parts,  and  on  dwarf-branches ;  it  is  not  until  some  time  later  that  the  dwarf- 
branches  are  developed  and  the  permanent  arrangement  attained. 

USES.  Several  species  are  commonly  cultivated  in  this  country,  partly  on 
heaths  and  moors,  and  partly  in  plantations  and  as  ornamental  trees,  such 
as  Mountain  Pine  (Pinus  montana,  Cen.  Ear.);  Austrian  Pine  (P.  laricio, 
Eur.);  Scotch  Fir  (P.  silvestris,  Eur.) ;  Weymouth  Pine  (P.  strobus,  N.  Am.) ; 
common  Ked  Pine  (Picea  excelsa,  Cen.  and  N.  Eur.)  ;  White  Pine  (P.  alba,  N. 
Am.)  ;  Abies  pectinata  (Common  Fir,  S.  and  Cen.  Eur.) ;  A.  nordmanniana 
(Crimea,  Caucasus);  A  lalsamea  (N.  Am.);  Tsuga  canadeiisis  (N.  Am.); 
Pseudotsurja  doitglasii  (W.N.  Am.) ;  Larch  (Larix  europcea,  Alps,  Carpathians) ; 
L.  sibirica  (N.E.  Kussia,  Siberia). — The  wood  of  many  species,  especially  Pine, 
on  account  of  its  lightness  and  because  it  is  so  easily  worked,  is  very  well 
adapted  for  many  useful  purposes.  The  wood  of  the  Yew-tree  is  very  hard 
and  is  used  for  ornamental  turning.  Resin  and  Turpentine  (i.e.  Besin  with 
essential  oils,  the  name  being  derived  from  the  Terebinth-tree,  from  which 
formerly  a  similar  material  was  obtained)  are  extracted  from  Pinus  laricio  and 
P.  pinaster.  Oil  of  Turpentine  is  obtained  by  distillation  of  turpentine  with 
water;  Tar  by  dry  distillation  of  Pine-wood.  Canada -balsam  is  from  North 


CONIFERS. 


267 


American  Abies-speciea  (A.  balsamea  and  Fraseri).  The  officinal  Turpentine 
is  mainly  obtained  from  Pinus  pinaster  (South  of  France),  P.  tada,  australis, 
utrobus  (Weymouth  Pine)  and  other  North  American  species;  more  recently 
also  from  P.  silvestris  (Scotch  Fir),  maritlma,  laricio,  Picea  excelsa,  and  others ; 
Venetian  Turpentine,  from  Larch  (S.  Eur.)  Amber  is  resin  from  a  Tertiary 
plant  (Pityoxijlon  succiniferum),  closely  related  to  the  Pine,  which  grew  especially 
in  the  countries  round  the  South-East  coast  of  the  Baltic.  Pinus  pinea  (the 
Pine,  S.  Eur.)  has  edible  seeds  and  also  P.  cembra  (in  Cen.  Eur.  and  Siberia). 

Order  3.  Taxodiaceae.  The  vegetative  leaves  and  cone-scales 
are  arranged  spirally.  The  ovules  (2-9) 
are  situated  either  at  the  base  of  the 
ovuliferous  scales,  in  which  case  they 
are  erect ;  or  at  their  centre,  when  they 
are  generally  more  or  less  inverted.  The 
ovuliferous  scale  is  more  or  less  united 
with  the  cover-scale,  and  projects  be- 
yond the  surface  of  the  cone-scale,  like  a 
comb  (Fig.  269).  The  vascular  bundles, 
which  extend  into  the  cover-scale,  have 
the  usual  leaf-arrangement,  viz.  the 
wood  placed  above  the  bast ;  while  those 
bundles  which  enter  the  ovuliferous 
scale  have  this  arrangement  of  the 
bundles  reversed. 


FIG.  239.  —  Crypiomeria  ja« 
pontca.  Portion  of  longitudinal 
section  through,  female  flower, 
d  cover-scale';  /  ovuliferous 
scale ;  ou  ovules ;  fv  and  fv' 
vascular  bundles;  the  xylem  is 
indicated  by  a  wavy  line,  and 
the  phloem  by  a  straight  line. 


Taxodium  distichum  (the  North  American  "  Swamp  Cypress ")  has  annual 
dwarf -branches,  with  distichous  leaves,  and  cone-like  "  pneumathodia."  In  the 
Tertiary  period  it  was  very  common  in  the  Polar  regions.  Sequoia  (Welling- 
t<mia)  gigantea  is  the  famous  Californian  Giant-Fir,  or  Mammoth-Tree,  which 
attains  a  height  of  300  feet,  a  diameter  of  36  feet,  and  is  said  to  live  for  1,500 
years.  Cryptomeria  japonica  (Japan,  China)  has  the  least  adnate  ovuliferous 
scales  ;  Gli/pto^trobus  (China) ;  Arthrotaxis  (Tasmania) ;  Sciadopitys  verticillata 
(the  only  species  in  Japan)  has,  like  Pinus,  scale-like  leaves  on  the  long-branches, 
of  which  those  which  are  situated  at  the  apex  of  the  annual  shoots  suppoit 
"  double  needles,"  i  e.  dwarf-branches  similar  to  the  two-leaved  dwarf-branches 
in  Pinus,  but  without  bud-scales,  and  with  the  two  leaves  fused  together  at  the 
edges  into  one  needle,  which  turns  its  upper  surface  away  from  the  long-branch. 

Order  4.  Cupressaceae  (Cypresses).  The  leaves  are  opposite 
or  verticillate,  sometimes  acicular,  but  most  frequently  scale-like 
(Fig.  270).  In  the  species  with  scale-like  leaves,  the  seedlings 
often  commence  with  acicular  leaves  (Fig.  272),  and  branches 
are  sometimes  found  on  the  older  plants  which  revert  to  this 
form,  seeming  to  indicate  that  the  acicular  leaf  was  the  ori- 


268 


GYMNOSPERIO;. 


ginal  form  (atavism).  The  so-called  "  Eetinospora  "  species  are 
seedling-forms  of  Biota,  Thuja,  Chamcecyparis,  which  have  been 
propagated  by  cuttings,  and  retain  the  seedling- form.  The  flowers 
are  monoecious  or  dioecious.  The  male  flowers  are  short,  and 
have  shield-like  stamens,  bearing  most  frequently  several  pollen- 
sacs.  The  cover-scales  and  ovuliferous  scales  are  entirely  fused 
together  and  form  undivided  cone-scales,  opposite  or  whorled;  the 
oculiferous  scales  have  slight  projections  near  the  base  on  which 
l-2-several  erect  ovules  are  developed  (Fig.  274).  Most  frequently 
2  cotyledons. —  Evergreen  trees  and  shrubs. 


Fia.  270. — Cupressus 
gooeniana. 


FIG.  271. — Portion  of  a  branch 
of  Thuja  orientalis  (magnified). 
The  leaf  at  the  base  on  the  right 
has  a  branch  in  its  axil. 


FIG.  272.— Seedling  of 
Thuja  occidentalis.  The 
branch  (g)  is  borne  in 
the  axil  of  the  leaf  s. 


Juniperus  (Juniper).  Dioecious.  The  cone-scales  become  flesliy 
und  fuse  together  to  form  most  frequently  a  1-3  seeded 
"  berry-cone."  J.  communis  (Common  Juniper)  has  acicular  leaves,  borne  in 
whorls  of  three,  and  the  "  berry-cone  "  is  formed  by  a  trimerous  whorl  of  cone- 
scales  (Fig.  273).  J.  sabina  and  J.  virginiana  have  "  berry-cones"  formed  from 
several  dimerous  whorls  of  cone-scales ;  the  leaves  are  connate  and  opposite, 
needle-  and  scale-like  leaves  are  found  on  the  same  plant. 

Cupressus  (Cypress).  Monoecious.  The  cones  are  spherical ;  the 
cone-scales  shield-like,  generally  five-cornered  and  woody  (Fig. 
270),  each  having  many  seeds.  The  leaves  are  scale-like.— Thuja. 


CONIFERS. 


269 


Monoecious.  Cones  oblong.  The  cone-scales  are  dry,  as  in  the 
Cypress,  but  leathery  and  imbricate,  and  not  shield-like ;  each 
cone-scale  bears  2-3  seeds.  The  leaves  are  most  frequently 
dimorphic;  those  leaves  which  are  situated  on  the  edges  of  the 
flat  branches  are  compressed,  and  only  these  bear  buds,  which 
fire  developed  with  great  regularity,  generally  alternately,  on  both 
sides  of  the  branch  ;  those  which  are  situated  on  the  flattened 
surfaces  are  pressed  flat  and  broad,  and  never  bear  branches 
(Fig.  271).  Along  the  central  line  of  each  leaf  there  is  a  resin- 
canal  (Fig.  271). — Cham(ecyparis,Callitris,Libocedrus,  Tlnijopsis  (1  species  : 
T.  dolabrita;  in  Japan). 


FIG.  273.— Branch  of  Juniper  with 
"  berry-cones." 


FIG.  274.  —  Cupressus  lawsoniana.  Lonsi- 
tudinal  section  through  female  cone.  Two 
ovules  (ou)  are  bisected;  /ovuliferous  scales. 


OFFICINAL.  Junipervs  sabina  from  Central  and  South  of  Europe  (the  young 
branches  yield  an  essential  oil).  The  wood  of  J.  communis  is  used  in  the 
production  of  an  essential  oil,  and  J.  oxycedrus  in  the  production  of  empy- 
reumatic  oil.  The  '•  berry-cone  "  of  J.  communi*  is  officinal,  and  is  also  used 
for  gin. — The  wood  of  J.  virginiana  (N,  Am.)  is  known  as  red  cedar,  and  is 
used  for  lead-pencils.  Sandarack  resin  is  obtained  from  Callitris  quadrivalvis 
(N.W.  Africa). 

THE  FOLLOWING  ARE  CULTIVATED  IN  GARDENS: — Thuja  occid  entails  (Arbor  vitae) 
(N.  Am.),  and  orientalis  (China,  Japan);  Juniperus  salina  and  virginiana; 
Thvjopsis  dolabrata  (Japan) ;  Cupressus  laivsoniana  (California),  C.  sempervirens 
(S.  Eur.,  W.  Asia),  and  other  species,  are  grown  especially  in  conservatories, 
and  in  Southern  Europe  particularly  in  cemeteries. — The  Retinospora  species 
which  are  so  often  planted,  do  not  belong  to  an  independent  genus,  but  are 
obtained  from  cuttings,  taken  from  seedling-plants  with  acicular  leaves  (see  page 
267). 


270 


GYMNOSPERMJ!. 

Class  ill.    Gneteae. 


This  class,  independent  of  extinct  forms,  comprises  the  most 
highly  developed  of  the  Gymnosperms,  partly  from  the  circum- 
stance that  a  perianth  of  2-4  members  encloses  the  terminally 


I 


GNETEJ1.  271 

placed  ovule,  which  is  provided  with  one,  or  (in  Gnetum)  two, 
integuments,  and  partly  owing  to  the  fact  that  the  wood  has  true 
vessels.  There  is  only  one  order. 

Order.  Gnetaceae.  The  three  known  genera  differ  very  much  in  appear- 
ance. Wehvitschia  mirabilis  (from  the  deserts  of  South  Western  Africa)  is  the 
oldest  (?)  genus  now  living.  It  resembles  a  giant  radish,  in  that  the  hypocotyl 
is  the  only  part  of  the  main  axis  of  the  stem  which  becomes  developed.  It 
attains  a  circumference  of  upwards  of  four  metres  with  a  length  of  J-f  of  a 
metre.  It  bears  only  two  oblong,  leathery  leaves  (Fig.  275)  which  are  torn  into 
segments  at  the  apex  and  lie  on  the  surface  of  the  soil ;  these  are  the  two  first 
foliage -leaves  which  succeed  the  cotyledons,  and  they  are  remarkable  for  their 
enormous  length  (upwards  of  two  metres)  as  well  as  for  their  long  duration, 
living  as  long  as  the  plant  itself.  In  their  axils  are  situated  the  4-rowed,  spike- 
like  male  and  scarlet-coloured  female  cones,  upon  dichotomous  branches.  The 
perianth  consists  in  the  $  of  2  alternating  pairs  of  leaves,  the  inner  ones  of  which 
are  slightly  united.  The  androecium  likewise  consists  of  2  whorls  :  the  external 
(transverse)  with  2,  the  internal  with  4  stamens  ;  the  lower  halves  of  the  6 
filaments  uniting  to  form  a  cup.  Each  of  the  terminal  anthers  corresponds  to 
a  sorus  of  3  sporangia,  the  sporangia  being  fused  together,  and  opening  at  the  top 
by  one  three-rayed  cleft.  In  the  centre  of  the  $  -flower  there  is  a  sterile 
ovule.  In  the  £  -flower  a  perianth  of  two  connate  leaves  is  present. — Ephedra 
(desert  plants,  especially  in  the  Mediterranean  and  W.  Asia)  at  first  sight  re- 
sembles an  Equisetum;ihe  stems  are  thin,  long- jointed,  and  the  leaves  opposite, 
small,  and  united  into  a  bidentate  sheath ;  <J  -perianth  of  two  connate  leaves 
(median  leaves) ;  2-8  stamens  united  into  a  column.  Each  anther  is  formed 
of  2  sporangia  (is  bilocular).  $  mainly,  as  in  Wehvitschia.  The  seeds  are  sur- 
rounded by  the  perianth  which  finally  becomes  red  and  fleshy.  There  are  30 
species. — Gnetum  has  opposite,  lanceolate,  pinnately-veined,  leathery  leaves. 
They  are  mostly  climbers  (Lianas)  from  Tropical  Asia  and  America.  The  <J  - 
flowers  have  a  tubular  perianth,  (formed  from  two  median  leaves)  which 
surrounds  a  centrally-placed  filament,  bearing  2  anthers.  In  the  ?  -flower 
there  is  a  similar  perianth,  surrounding  an  ovule  provided  with  2  integuments . 
The  perianth  becomes  fleshy  and  envelops  the  hard  seed.  20  species. 

From  the  circumstance  of  Welwitschia  having  $  flowers  which,  besides 
stamens,  possess  also  a  rudiment  of  an  ovule,  Celakovsky  draws  the  inference  that 
the  earliest  Gymnosperms  had  hermaphrodite  flowers  which  from  this  structure 
became  differentiated  entirely  into  3  -  and  £  -flowers,  with  the  exception  of 
Welwitschia  only,  in  which  this  differentiation  was  only  carried  out  in  the  $  - 
flower.  This  theory  has  so  far  been  scarcely  proved. 

Fossil   Gymnosperms. 

The  earliest  continental  plants  which  are  known  belong  to  the  COBDAITACE.E, 
a  group  of  plants  which  existed  as  early  as  the  Silurian  period ;  they  were 
Gymnosperms,  but  it  has  not  yet  been  determined  whether  they  were  Cycads  or 
Conifers.  The  CYCADS,  even  in  the  Coal  period,  were  scarce  ;  they  attained  their 
fullest  development  in  Jurassic  and  Cretaceous  periods,  during  which  they  were 
rich  in  species  and  genera,  and  extended  as  far  as  the  Polar  regions.  In  addition 
W.  B  T 


272  GYMNOSPERMJE. 

to  these,  Taxacese,  Abietaceae,  and  Taxodiaceaa  appeared  in  the  Carboniferous 
period.  The  TAXACE;E  appear  to  have  attained  their  culmination  in  the 
Jurassic  and  Cretaceous  periods ;  Ginkgo  appears  in  the  Rhaetic  ;  Torreya,  in  the 
Cretaceous ;  Taxus  and  Podocarpus  in  the  Tertiary  periods.  The  ABIETACE^: 
also  appear  in  the  Carboniferous ;  Pinus  was  first  known  with  certainty  in  the 
English  Weald  and  in  the  Cretaceous  ;  almost  all  other  contemporary  genera  are 
represented  in  this  latter  period.  The  ABAUCABIACE^;  first  appear,  with  certainty, 
in  the  Jurassic.  The  TAXODIACE^E  may  be  traced  back  as  far  as  the  Carboni- 
ferous (?) ;  Sequoia  is  first  found  in  the  lowest  Cretaceous,  at  tbat  period  it 
spread  throughout  the  entire  Arctic  zone,  and  being  represented  by  a  large 
number  of  species,  formed  an  essential  part  of  the  forest  vegetation.  Sequoia 
played  a  similar  part  in  the  Tertiary  period.  The  CUPRESSACE^E  are  first 
known  with  certainty  in  the  Jurassic,  but  they  appeared  more  frequently  and 
numerously  in  the  Tertiary  period,  in  which  most  of  the  present  living  genera 
were  to  be  found.  The  GNETACEJE,  according  to  a  theory  advanced  by  Renault 
were  represented  in  the  Coal  period  by  the  genus  Steplianospermum,  which  had 
four  ovules  enclosed  by  an  envelope. 

Recent  investigations  have  established  Ginkgoineee  as  a  distinct  Class  inter- 
mediate between  the  Cycadeae  and  the  Conifers,  Ginkgo  biloba  being  the  only 
existing  representative  of  this  Class.  It  is  more  closely  allied  to  the  Cycadeaa. 


FIG.  275A.— Ginkgo  bilobo: 
Motile  Rpermatozoida  in  the 
pollen-tube;  n  nucleus. 

The  pollen-grains  are  spherical,  and  in  these  a  prothallium  is  formed  generally 
composed  of  three  cells,  the  most  internal  of  which  is  the  antheridium.  This 
contains  two  ovoid  spermatozoids  having  at  the  anterior  end  a  three-coiled 
spiral  groove  bearing  numerous  cilia  (Fig.  275A). 


DIVISION  V. 
ANGIOSPERMS. 

See  pages  3  and  224.  To  this  Division  belong  the  majority 
of  the  Flowering  -  plants.  They  are  divided  into  two  parallel 
classes,  the  Monocotyledons  and  the  Dicotyledons,  which  differ 
from  each  other  not  only  in  the  number  of  cotyledons,  which,  with 
a  few  exceptions,  is  one  in  the  former,  two  in  the  latter,  but  also 
in  the  internal  structure  of  the  stem,  the  venation  of  the  leaves, 
the  number  of  the  parts  of  the  flower,  etc.  Assuming  that  these  two 
classes  have  sprung  from  a  common  origin,  it  is  amongst  the  Helobiete  in  the 
first,  and  amongst  the  Polycarpicse  in  the  second  class  that  we  might  expect 
to  find  closely  allied  forms,  which  might  reasonably  be  supposed  to  have  varied 
less  from  this  original  type.  As  for  the  rest,  they  seem  to  stand  quite  parallel, 
without  exhibiting  any  close  relationship.  It  is  scarcely  proved  that  the 
Monocotyledons  are  the  older  class. 

[Our  knowledge  of  the  forms  included  under  the  Angiosperms 
has  recently  been  considerably  increased  by  Treub  (Ann.  d.  Jar. 
Sot.  d.  Buitenzorg,  1891),  who  has  shown  that  the  Casuarinas  differ 
in  many  important  points  from  the  typical  Angiosperms.  Among 
other  characters  the  pollen-tube  is  found  to  enter  the  ovule  near 
the  chalaza  and  therefore  at  the  opposite  end  to  the  micropyle, 
and  Treub  therefore  suggests  that  these  plants  should  be  placed 
in  a  subdivision  termed  Chalazogams. 

According  to  this  view  the  principal  divisions  of  the  Angio- 
spernis  would  be  represented  thus  : — 

Angiospermae. 

Sub-division.  Sub-division. 

CHALAZOGAMES.  POROGAMES. 

Class.  Classes. 

Chalazogames.  Monocotyledones,  Dicotyledones. 

More  recently  Nawaschin  (Bull.  Acad.  Imp.  Sci.  St.  Petersb., 
ser.  iii.,  xxxv.)  has  shown  that  Betula,  and  Miss  Benson  (Trans. 
Linn.  Soc.,  1894)  that  Alnus,  Corylus,  and  Carpinus  also  belong  to 
the  Chalazogams. 

273 


274  ANGIOSPERHE. 

Our  knowledge,  however,  is  still  so  incomplete  that  one  would 
hesitate  to  accord  the  full  systematic  value  which  Dr.  Treub 
attaches  to  his  discovery  until  the  limits  of  the  Chalazogamic 
group  are  better  denned  ;  and  it  would  hardly  be  justifiable  to 
include  the  Casuarinas  and  the  above-noted  genera  in  one 
family.] 

Class  1.    Monocotyledones. 

The  embryo  has  only  one  cotyledon ;  the  leaves  are  as  a  rule  scattered, 
with  parallel  venation ;  the  vascular  bundles  of  the  stem  are  closed, 
there  is  no  increase  of  thickness.  The  flower  is  typically  constructed  of 
five  3-merous  whorls,  placed  alternately. 

THE  EMBRYO  is  generally  small  in  proportion  to  the  abundant 
endosperm  (exceptions,  see  Helobiece),  and  its  single  cotyledon  is 
often  sheath-like,  and  very  large.  On  the  germination  of  the 
seed  either  the  entire  cotyledon,  or  its  apex  only,  most  generally 
remains  in  the  seed  and  absorbs  the  nutritive-tissue,  while  the 
lower  portion  elongates  and  pushes  out  the  plumule  and  radicle, 
which  then  proceed  with  their  further  growth.  The  primary  root 
in  most  cases  soon  ceases  to  grow,  but  at  the  same  time,  however, 
numerous  lateral  roots  break  out  from  the  stem,  and  become  as 
vigorous  as  the  primary  root,  or  even  more  so.  Increase  in 
thickness  does  not  take  place  in  these  roots ;  they  branch  very 
little  or  not  at  all,  and  generally  die  after  a  longer  or  shorter  time. 

THE  STEM  is  frequently  a  corm,  bulb,  or  other  variety  of  under- 
ground stem,  as  the  majority  of  the  Monocotyledons  are  perennial, 
herbaceous  plants ;  it  has  scattered,  closed  vascular  bundles  (Fig. 
276),  and  no  cambium  by  which  a  continuous  thickening  may  take 
place.  The  stem  of  the  Palms,  however,  attains  a  very  consider- 
able thickness,  which  is  due  to  the  meristem  of  its  growing-point 
continually  increasing  in  diameter  for  a  lengthened  period  (often 
for  many  years),  until  it  has  reached  a  certain  size.  In  this  con- 
dition the  growing-point  has  the  form  of  an  inverted  cone,  and  it 
is  only  when  this  cone  has  attained  its  requisite  size  that  the  for- 
mation of  a  vertical  cylindrical  stem  commences.  Certain  tree- 
like Liliaceas,  as  Draccena,  Aloe,  etc.,  have  a  continuous  increase  in 
thickness ;  this  is  due  to  a  meristematic  layer,  which  arises  in  the 
cortex,  outside  the  original  vascular  bundles,  which  were  formed  at 
the  growing-point  of  the  stem.  This  meristem  continues  to  form 
thick-walled  parenchyma  and  new,  scattered  vascular  bundles. 
The  primary  vascular  bundles,  in  the  Palms  and  others,  run  in  a 


MONOCOTYLEDONES. 


275 


curved  line  from  their  entrance  into  the  stem  at  the  base  of  the 
leaf,  towards  the  centre  of  the  stem,  and  then  bend  outwards  and 
proceed  downwards  in  a  direction  more  parallel  to  the  sides  of 
the  stem  (Fig.  277).  The  bundles  formed  later,  in  those  stems 
which  increase  in  thickness,  are  not  continued  into  the  leaves. 

THE  BRANCHING  as  a  rule  is  very  slight,  the  axillary  buds  of  the 
majority  of  the  leaves  never  attaining  development,  e.g.  in  the 
Palms,  bulbous  plants  and  others.  As  the  cotyledon  arises  singly, 


Fie.  276.— Transverse  Bection  of  the 
stem  of  a  Palm:  v  v  is  the  wood  portion, 
b  b  the  bast  portion  of  the  vascular 

bundles. 


FIG.  2/7.— Diagrammatic  represen- 
tation of  the  course  of  the  vascular 
bundles,  from  the  stem  into  the  leaves 
in  a  Monocotyledon. 


the  succeeding  leaves  also  must  be  scattered,  but  they  are  fre- 
quently arranged  in  two  rows  (Grasses,  Iris,  etc).  The  first,  leaf 
borne  on  a  branch  (the  "  Fore-leaf,"  l — the  bracteole,  if  on  a  floral 
shoot)  has  generally,  in  the  Monocotyledons,  a  characteristic  form 
and  position,  being  situated  on  the  posterior  side  of  its  own  shoot, 
and  hence  turned  towards  the  main  axis ;  it  is  sometimes  provided 
with  two  laterally-placed  keels  (Figs.  279  /,  290  0t),  but  the  midrib 
is  often  absent.  It  arises  in  some  cases  from  two  primordia,  which 
at  the  beginning  are  quite  distinct,  and  thus  has  been  regarded  as 
formed  by  two  leaves.  It  is,  however,  only  one  leaf,  a  fact  which  is 
evident  from  several  circumstances,  one  being  that  it  never  supports 
more  than  one  shoot,  and  this  stands  in  the  median  plane  (Fig.  279). 
THE  LEAVES  are  amplexicaul,  and  have  a  large  sheath  but  no 
stipules  ;  the  blade  is  most  frequently  long,  ligulate,  or  linear,  en- 
tire, with  parallel  venation,  the  veins  being  straight  or  curved 
1  "Fore-leaf"  is  adopted  as  a  translation  of  "  Vorblatt." 


276 


MONOCOTYLEDONES. 


(Figs.  300,  309).  Connecting  the  large  number  of  veins  which 
run  longitudinally,  there  are  as  a  rule  only  weak  tran verse  ones. 
It  is  very  rarely  that  other  forms  of  leaves  are  found,  such  as  cor- 
date (Figs.  302,  312),  or  that  the  blade  is  branched,  or  the  vena- 
tion is,  for  example,  pinnate  or  palmate  (Figs.  225,  298)  ;  these 
deviations  are  especially  found  in  the  Araceae,  the  Palms,-  the 
Scitamineae  (Fig.  308),  the  Dioscoreaceae,  and  in  several  aquatic 
plants.  The  incisions  in  the  Palm-leaf  are  derived  by  the  split- 
ting of  an  originally  entire  leaf. 

THE  STRUCTURE  OF  THE  FLOWER  is  generally  as  follows  :  Pr3  +  3, 
A3  +  3,  G3,  rarely  S3  +  P3  with  the  other  members  unchanged.1 
Instead  of  3,  the  numbers  2  and  4  may  occur ;  rarely  others.  In 
all  these  instances  there  are  5  whorls,  which  regularly  alternate 
with  one  another,  most  frequently  in  the  3-merous  flower,  as  in  the 
diagram  (Fig.  278).  This  diagram  is  found  in  the  following 


*• 


278.  279.  230. 

FIG.  278. — Diagram  of   the  ordinary,  regular  flower  in   the  Monocotyledons :  «  is  the 
bract. 

FIG.  279.— Diagram  of  Iris  :  /  the  bracteole  ;  in  its  axil  is  a  shoot  with  its  bracteole. 
FIG.  280.— Diagram  of  Orchis  •.  I  the  lip ;  a-  a-  the  two  staminodes. 

orders :  Liliaceae,  Convallariacese,  Juncacea3,  Bromeliacea3,  Ama- 
ryllidaceee,  Dioscoreacese,  Palmes,  some  Araceee,  and  in  some  small 
orders,  and  may  be  considered  as  the  typical  structure  and  also 
the  starting1  point  for  the  exceptional  orders.  The  ovary  in  many 
Monocotyledons  has  many  ovules,  and  the  fruit  becomes  a  many- 
seeded  berry  or  capsule  ;  this  form  is  no  doubt  the  oldest.  In 
others  the  number  of  seeds  becomes  reduced  to  1,  and  the  fruit 
then  becomes  a  cypsela,  or  a  drupe  (e.g.  Graminece,  Cyperacece, 
Palmce,  etc). 

Deviations  from  this   typical   floral  structure  in  some  instances 

1  Eegarding  these  and  other  abbreviations  see  the  appendix  in  the  book. 


MONOCOTYLEDONES.  277 

may  be  traced  to  suppression,  very  rarely  to  a  splitting  of  certain 
members,  the  typical  relative  positions  not  being  changed.  Thus, 
the  Iridaceaa,  the  Cyperacese,  most  of  the  Gramineae  and  some 
Juncaceaa  deviate  in  having  only  3  stamens  (Fig.  279),  the  inner 
whorl  (indicated  by  *)  not  becoming  developed.  The  Musaceae 
differ  in  the  posterior  stamen  not  being  developed ;  Zingiberacece 
(Fig.  314),  Marantacece,  and  Cannacece,  in  the  fact  that  only  1  of 
all  the  stamens  bears  an  anther,  and  the  others  are  either  sup- 
pressed or  developed  into  petaloid  staminodes,  with  some  perhaps 
cleft  in  addition.  The  Orchideae  deviate  in  having,  generally, 
only  the  anterior  stamen  of  all  the  6  developed  (Fig.  280).  In 
this,  as  in  other  instances,'  the  suppression  of  certain  parts  of 
the  flower  is  often  connected  with  zygoworphy  {i.e.  symmetry  in 
one  plane),  chiefly  in  the  inner  perianth-whorl,  but  also  in  the 
other  whorls.  In  the  Orchids,  the  perianth-leaf  (the  labellum, 
Fig.  28.0  I)  which  is  directly  opposite  the  fertile  stamen,  is  larger 
and  altogether  different  from  the  others.  The  perianth -leaves 
may  also  be  suppressed ;  see,  for  example,  the  two  diagrams  of  the 
Cyperacese  (Fig.  284).  In  some  orders  the  suppression  of  these 
leaves,  which  form  the  basis  of  the  diagram,  is  so  complete  that  it 
is  hard  to  reduce  the  actual  structure  of  the  flower  to  the  theo- 
retical type,  e.g.  the  Grasses  (Fig.  290)  and  Lemna  (Fig.  303).  In 
the  first  family,  which  especially  comprises  water-plants,  a  some- 
what different  structure  is  found  ;  thus  Fig.  282  differs  somewhat 
from  the  ordinary  type,  and  other  flowers  much  more  so;  but  the 
floral  diagrams  which  occur  in  this  family  may  perhaps  be  con- 
sidered as  the  most  probable  representatives  of  an  older  type,  from 
which  the  ordinary  pentacyclic  forms  have  taken  their  origin.  In 
favour  of  this  theory  we  have  the  larger  number  of  whorls,  the 
spiral  arrangement  of  some  of  these  in  the  flower,  with  a  large  and 
indefinite  number  of  stamens  and  carpels,  the  perfectly  apocarpous 
gyiiceceum  which  sometimes  occurs,  etc.,  etc. 

The  Monocotyledons  are  divided  into  7  Families : — 

1.  HELOBIE.E.     This  family  forms  a  group  complete  in  itself.     It  commences 
with  hypogynous,   perfect  flowers,  whose   gyncecium  is  apocarpous  and  ter- 
minates in  epigynous  and  more  or  less  reduced  forms. 

2.  GLUMIFLOKJE.     These  have  as  a  starting  point  the  same  diagram  as  the 
following  families,  but  otherwise  develope  independently. 

3.  SPADICIFLOR^S.      Also   an  independent  branch,  or  perhaps  two  different 
ones  which  terminate  in  much  reduced  forms. 

4.  ENANTIOBLAST^E.    These  ought  perhaps  to  be  amalgamated  with  the  follow- 
ing family. 


278  MONOCOTYLEDONES. 

5.  LILIIFLOBJE.     These  advance  from  forms  with  the  typical  diagram  and 
hypogynous  flower,  to  epigynous  and  reduced  forms. 

6.  SCITAMINE^E  and 

7.  GYNANDR.E.     Two  isolated  families,  which  probably  have  taken  their  ori- 
gin from  Liliiflorae,  and  have  epigynous,  mostly  zygomorphic,  and  much  re- 
duced forms. 

Family  1.     Helobiese. 

To  this  family  belong  only  water-  or  marsh-plants ;  the  endosperm 
is  wanting,  and  they  possess  an  embryo  with  a  very  large  hypocotyl 
prolonged  downwards  and  often  club-like.  The  perianth  is  often 
differentiated  into  calyx  and  corolla  ;  the  flower  is  regular,  and  in 
the  first  orders  to  be  considered,  may.be  reduced  to  the  ordinary 
Monocotyledonous  type  ;  there  are,  however,  usually  found  two 
3-merous  whorls  of  carpels  (Fig.  282),  and'  thus  in  all  6  whorls, 
or  again,  the  number  of  carpels  may  be  indefinite ;  the  number  of 
stamens  also  may  be  increased,  either  by  the  division  of  the 
members  of  a  whorl,  or  by  the  development  of  additional  whorls. 
Syncarps,1  with  nut  or  follicular  fruitlets,  are  very  common,  for 
example,  in  the  first  orders ;  in  the  last  (Hydrocharitaceae)  the 
•carpels  are  not  only  united,  but  the  ovary  is  even  inferior. 

The  primitive  type  appears  to  be  a  hypogynous  flower,  similar  to  that  of  the 
Juncaginaceae  or  Alismaceas,  with  several  3-merous  whorls,  and  free  carpels, 
each  with  many  ovules  'the  green  perianth  in  this  instance  being  no  doubt  older 
than  the  coloured  ones.  If  we  take  a  flower  with  this  structure  as  the  start- 
ing point,  then  the  family  dcvelopes  partly  into  epigynous  forms,  partly  into 
others  which  are  so  strongly  reduced  and  exceptional  that  it  is  scarcely  possible 
to  refer  them  to  the  ordinary  type.  The  family,  tbrough  the  peculiar  Zosterece, 
appears  to  approach  the  Araceae,  in  which  Potamogetonaccce  and  Najadacets  are 
included  by  some  authorities.  However,  the  inclusion  of  Potamogeton,  and  with 
it  liuppia  and  Zannichellia,  in  the  Juncaginaceaa  appears  quite  correct.  It  would 
scarcely  be  right  to  separate  Zosterece  from  these.  Great  stress  has  often  been 
laid  upon  the  similarity  with  the  Ranunculaceae  which  is  found  in  the  Alismaceae, 
but  it  is  scarcely  more  than  an  analogous  resemblance. 

Order  1.  Juncaginaceae.  The  §  ,  regular,  hypogynous  flowers 
have  the  perianth  3  +  3,  sepaloid,  stamens  3  +  3  (with  extrorse  an- 
thers), and  carpels  3  +  3  (free  or  united),  of  which  last,  however, 
one  whorl  may  be  suppressed  (in  Triglochin  maritima  all  6  carpels 
are  developed,  in  T.palustris  the  inner  whorl  is  unfertile).  Inflores- 
cence long  spikes.  Embryo  straight. — Marsh-plants  with  radical, 
rush-like  leaves,  arranged  in  two  rows,  and  often  sheathing  and 
ligulate  ("  squamulge  intravaginales  ")  ;  the  inflorescence  is  a  spike 
or  raceme. — Scheuchzeria.  Carpels  almost  free  ;  in  each  at  least 
1  Syncarp  =  cluster  of  fruits  belonging  to  one  flower. 


HELOBIE&.  279 

two  ovules.  Follicles. — Triglocliin  has  long,  fine  racemes  without 
bracts  or  bracteoles ;  one  ovule  in  each  carpel.  The  carpels  in 
the  two  native  species  are  united,  but  separate  when  ripe  as  a 
schizocarp,  loosening  from  below;  they  open  along  the  ventral 
suture  or  remain  closed ;  a  linear  central  column  remains. 
The  most  reduced  is  Lilcea  (1-2  sp.  Am.)  — Protogynous.  About  10  species. 
Temp.  Fossils  in  Tertiary. 

Order  2.  Potamogetonacese.  The  aquatic  plants  belonging 
to  this  order  are  perennial,  living  entirely  submerged,  or  with 
floating  leaves,  and  preferring  still  water.  The  leaves  are  alter 
nate,  in  some  linear  and  grass-like,  in  others  there  is  an  elliptical 
floating  blade,  supported  by  a  linear  submerged  petiole.  Axillary 
scales.  The  fruit  is  generally  a  syncarp  with  nuts  or  drupes;  the 
embryo  is  curved,  of  very  various  forms. 

Potamogeton  (Pond-weed).  The  rhizome  is  creeping,  sympodial 
(with  two  internodes  in  each  shoot-generation)  ;  the  inflorescence  is 
a  terminal,  many-flowered  spike,  without  floral-leaves ;  below  it  are 
found  2  foliage-leaves  placed  nearly  at  the  same  height,  from  whose 
axils  the  branching  is  continued  cymosely.  The  flowers  are  $  »  4- 
merous,  naked,  and  consist  only  of  4  stamens,  with  the  connectives, 
broadly  developed  at  the  back  of  the  anthers,  resembling  a  perianth, 
and  of  4  free,  sessile  carpels.  They  are  common  plants  in  fresh  water. 
The  spike,  during  the  flowering,  is  raised  above  the  water.  Wind-pollinated  and 
protogynous. — Closely  allied  is  Euppia  (Tassel  Pond-weed),  in  salt  or  brackish 
water.  The  spike  has  only  two  naked  flowers,  each  consisting  of  2  stamens  find 
4  carpels.  The  stalks  of  the  individual  carpels  are  considerably  prolonged.  — 
Zannichellia  (Horned  Pond-weed)  is  monoecious ;  the  ?  -flower  consists  of  4 
(2-9)  carpels,  with  membranous,  bell-shaped  perianth  ;  long  styles  ;  the  $  -flower 
has  1  (-2)  stamens.  Althenia. 

Zostera  (Grass-wrack)  is  an  entirely  submerged,  marine  plant  with 
creeping  rhizome  (with  displacement  of  buds)  and  strap-shaped 
leaves.  The  flowering  shoots  are  sympodia  with  displacement  of 
the  axes  (Fig.  281).  The  inflorescence  is  a  peculiar,  flatly-com- 
pressed spike,  on  owe  side  of  which  the  flowers  are  borne  (Fig.  28  L). 
This  inflorescence  may  be  considered,  no  doubt  correctly,  to  be  derived  from  the 
symmetrical  spike  of  Potamogeton  by  strongly  dorsiventral  development,  and  by 
a  strong  suppression  of  tbe  floral  parts  taking  place  simultaneously.  Two 
rows  of  flowers  are  developed,  but  of  these  one  is  so  pressed  into  the  other  that 
apparently  only  one  is  present.  Each  flower  consists  of  only  1  stamen 
and  1  carpel  situated  at  the  same  height  (Fig.  281)  ;  the  unilocular 
ovary  encloses  1  pendulous  ovule  and  bears  a  bifid  style.  As 
regards  the  perianth  (?)  one  leaf  may  be  preset  (Z.  nana,  Fig. 
281  D).  The  pollen-grains  are  filamentous.  Pollination  takes 


FIG.  281.— Zostera.  A  Diagram  of  the  branching  of  the  floral  shoots:  I,  17  ....  are  the 
successive  shoot-genet ations,  eveiy  other  one  being  shaded  ;  gl  g2  ....  fore-leaves;  sp1 
sp2  ....  spathes  for  the  successive  spikes.  Each  shoot  is  united  for  some  distance 
with  the  parent  axis  (indicated  by  the  half-shaded  internodes).  Each  shoot  commences 
with  a  fore-leaf  turning  towards  the  parent  axis,  g;  succeeding  this  is  the  spathe,  sp  ;  and 
then  the  inflorescence.  The  fore-leaf  supports  a  new  lateral  shoot.  B  Diagram  of  a  shoot, 
IT,  which  is  borne  laterally  in  the  axil  of  the  fore-leaf  glt  on  the  shoot  I,  ga  its  fore-leaf; 
s)ia  its  spathe ;  sti  squamula?  intravaginales.  II  Is  the  gpadix  with  stamens  and  carpels; 
b  a  perianth-leaf  (or  connective  expansion,  similar  to  those  which  occur  in  Potamogeton). 
C  The  upper  portion  of  a  yonng  spadix  with  development  of  flowers.  D  Part  of  n  gpadix 
with  2  flowers  ;  the  parts  which  theoretically  belong  to  one  another  are  connected  by  a 
dotted  line. 

280 


HELOBIE33. 


281 


place  under  water.  Posidonia  and  Cymodocea  are  allied  to  these.  About 
70  species. 

Order  3.  Aponogetonaceae.  Aquatic  plants  with  tuberous  stem.  They  have 
a  single,  petaloid  perianth  (3-2-1 -leaved),  most  frequently  6  stamens  and  3(-6) 
carpe-s.  Straight  embryo. — About  15  species  (Africa,  Madagascar,  Tropical 
Asia  and  Australia). — Aponogeton  distachyus  and  A.  (Ouvirandra)  fenestralis  are 
grown  in  conservatories  ;  the  latter  has  lattice-like,  perforated  leaves. 

Order  4.  Najadacese.  Only  one  genus  Najas  (about  10  species) ;  annual 
fresh  water  plants  with  leaves  in  pairs  and  solitary,  unisexual  flowers.  The  c? 
flower  is  remarkable  in  having  a  terminal  stamen,  which  has  either  4  longi- 
tudinal loculi  or  1  central  one ;  on  this  account  the  stamen  of  Nuias  is  con- 
sidered by  some  authorities  to  be  a  stem  and  not  a  leaf-structure.  The  uni- 
locular  gynoeceum  and  the  single,  erect,  anatropous  ovule  are  also  terminal. 
Pollination  takes  place  under  the  water. 

Order  5.  Alisrnacese.  The  regular,  hypogynous  flowers  are  in 
some  species  unisexual  by  the  suppression  of  either  androecium  or 
gynoeceum  ;  they  have  a  6-merous  perianth,  generally  differentiated 
into  3  .epals  and  3  petals  ;  generally  6  stamens  in  the  outer  whorl 
(by  the  division  of  the  3 ;  Fig.  282)  and 
often  several  3-merous  whorls  inside 
these,  and  6-00  free  carpels  arranged 
cyclically  or  spirally.  Fruit  a  syncarp. 
— Marsh-  or  water-plants  with  radical 
leaves  and  long-stalked  inflorescences. 

A.  Butomece.      Follicles   with   many 
seeds,    u'hich    are    borne    on   nearly    the 
whole  of  the  inner  surface  of  the  cyclic 
carpels  (as  in  Nymph  aeacese).     Embryo 
straight. — Butomus     (Flowering    Rush, 

Fig.  282),  has  an  umbel  (generally  composed  of  3  helicoid  cymes). 
S3,  P3,  stamens  9  (6  +  3,  i.e.  the  outer  whorl  doubled),  G  3  +  3. 
B.  umbellatus ;  creeping  rhizome  with  triangular  Iris-like  leaves. — Hydrocleis. 
Limnocharis. 

B.  Alismece.     Fruit    achenes.     Latex    common    (in  the   inter- 
cellular spaces).     The    flowers  are   arranged  most   frequently    in 
single  or  compound  whorls.     Embryo  curved,  horse-shoe  shaped. — 
Alisma  has  S  3,  P  3,  A  6  (in  1  whorl,  grouped  in  pairs,  i.e.  doubled 
in  front  of  the   sepals),  and  1  ivhorl  of  1-seeded  achenes  on  a  flat 
receptacle.     The  leaves  are  most  frequently  radicle,  long-stalked; 
the  lamina  have  curved  longitudinal  veins,  and  a  richly  branched 
venation.     A.  plantugo. — Elisma  (E.  natans)  has  epitropous  (turned 
inwards)  ovules,  whilst  the  ovules  of  Alisma,  Sagittaria  and  others 
are  apotropous  (turned  outwards). — Echinodorus  (E.  ranunculoides) 


FIG.  282. — Diagram  of  Butomus : 
/bracteole. 


282  MONOCOTYLEDONES. 

has  a  convex  receptacle,  carpels  many,  united  and  capitate. 
Damasonium. — Sagittaria  (Arrow-head)  has  monoecious  flowers, 
several  whorls  of  stamens  and  spirally-arranged  achenes  on  a  very- 
convex  receptacle.  S.  aagittifolia  reproduces  by  tuberous  buds  formed  at 
the  end  of  long,  submerged  branches.  The  leaves,  in  deep  and  rapidly  running 
water,  are  long  and  strap-shaped,  but  in  the  air  arrow-shaped. 

Honey  is  secreted  in  the  flower  and  pollination  effected  by  insects.  Alisma 
plantago  has  12  nectaries.  The  submerged  flowers  of  Elisma  natans  remain 
closed  and  are  self-pollinated.  Butomus  has  protandrous  flowers.  There  are 
about  50  species,  which  mostly  grow  outside  the  Tropics. — Uses  insignificant. 
The  rhizome  of  some  is  farinaceous. 

Order  6.  Hydrocharitaceae.  This  order  differs  chiefly  from 
the  preceding  in  its  epigynous  flowers.  These  are  in  general  uni- 
sexual (dioecious),  and  surrounded  by  a  2-leaved  or  bi-partite 
xpathe  ;  they  are  3-merous  in  all  whorls,  but  the  number  of  whorls 
is  generally  greater  than  5,  sometimes  even  indefinite.  The  peri- 
anth is  divided  into  calyx  and  corolla.  The  ovary  is  unilocular  with 
parietal  placentation,  or  more  or  less  incompletely  plurilocular.  The 
fruit  is  berry-like,  but  usually  ruptures  irregularly  when  ripe.  Em- 
bryo straight. — Most  often  submerged  water-plants,  leaves  seldom 
floating  on  the  surface.  Axillary  scales  (squamulw  intravaginales). 

Hydrocharis.  Floating  water-plants  with  round  cordate  leaves  ; 
S3,  P3  (folded  in  the  bud);  £  -flowers  :  3  (-more)  flowers  inside 
each  spathe  ;  stamens  9-15,  the  most  internal  sterile.  ?  -flowers 
solitary ;  three  staminodes  ;  ovary  6-locular,  with  many  ovules 
attached  to  the  septa ;  styles  6,  short,  bifid.  [The  petals  of  the 
9  -flowers  bear  nectaries  at  the  base.  In  this  and  the  following 
genus  the  pollination  is  without  doubt  effected  by  insects.] 
H.  morsus  ranee  (Frog-bit)  has  runners ;  it  hibernates  by  means  of  special 
winter-buds — Stratiotes ;  floating  plants  with  a  rosette  of  linear, 
thick,  stiff  leaves  with  spiny  margin,  springing  from  a  short 
stem,  from  which  numerous  roots  descend  into  the  mud.  In- 
florescence, perianth,  and  ovary  nearly  the  same  as  in  Hydrocharis, 
but  the  ^-flower  has  12  stamens  in  3  whorls,  of  which  the 
outer  6  are  in  1  whorl  (dedoublement),  and  inside  the  perianth 
in  both  flowers  there  are  numerous  (15-30)  nectaries  (stami- 
nodes?). S.  aloides  (Water-soldier);  in  N".  Ear.  only  ?  -plants. 
— Vallisneria  fpiralis  is  a  tropical  or  sub -tropical  plant,  growing  gregariously  on 
the  mud  in  fresh  water.  The  leaves  are  grass-like,  and  the  plants  dioecious  ; 
the  (?  -flownrs  are  detached  from  the  plant,  and  rise  to  the  surface  of  the  water, 
where  they  pollinate  the  $  -flowers.  These  are  borne  on  long,  spirally-twisted 
peduncles  which  contract  after  pollination,  so  that  the  $  -flower  is  again  drawn 
under  the  water,  and  the  fruits  ripen  deeply  submerged. — Elodea  canadensis 


GLOMIFLOR.E.  283 

is  also  an  entirely  submerged  plant.  The  leaves  are  arranged  in  whorls  on  a 
well-developed  stem.  Only  •$  -plants  in  Europe  (introduced  about  1836  from 
N.  Am.).  This  plant  spreads  with  great  rapidity  throughout  the  country,  the 
reproduction  being  entirely  vegetative.  Hydrilta,  Hnlnphila,  Thalassia,  Enhalus. 
— In  many  of  these  genera  the  number  of  whorls  in  the  flower  is  remarkably  re- 
duced ;  for  example,  in  Vallisneria,  in  the  $  -flowers  to  2  :  Pr  3,  A  (1-)  3,  in  the 
?  to  3 :  Pr  3,  Staminodes  3,  G  3.— About  40  species  ;  Temp,  and  Trop. 

Family  2.     Glurniflorse. 

The  hypogynous  flowers  in  the  Juncacese  are  completely  developed 
on  the  pentacyclic,  trimerous  type,  with  dry,  scarious  perianth.  Even 
in  these  the  interior  whorl  of  stamens  becomes  suppressed,  and  the 
ovary,  which  in  Juncus  is  trilocular  with  many  ovules,  becomes  in 
Luzula  almost  unilocular,  but  still  with  3  ovules.  The  perianth  in 
the  Cyperaceas  and  Grraminese  is  reduced  from  hairs,  in  the  first  of 
these,  to  nothing,  the  flowers  at  the  same  time  collecting  more 
closely  on  the  inflorescence  (spike)  supported  by  dry  bracts  (chaff} ; 
the  number  of  stamens  is  almost  constantly  3  ;  stigmas  linear;  the 
ovary  has  only  1  loculus  with  1  ovule,  and  the  fruit,  which  is  a  cap- 
sule in  the  Juncacese,  becomes  a  nut  or  caryopsis. — The  endosperm 
is  large  and  floury,  the  embryo  being  placed  at  its  lower  extremity 
(Figs.  286  B,  291). — The  plants  belonging  to  this  order,  with  the 
exception  of  a  few  tropical  species,  are  annual  or  perennial  herbs. 
The  stems  above  ground  are  thin,  and  for  the  most  part  have  long 
internodes,  with  linear,  parallel-veined  leaves  which  have  long 
sheaths,  and  often  a  ligule,  i.e.  a  membranous  projection,  arising 
transversely  from  the  leaf  at  the  junction  of  the  sheath  and  blade. 
The  underground  stems  are  short  or  creeping  rhizomes.  The 
flowers  are  small  and  insignificant.  Wind-  or  self-pollination. 

Order  1.  Juncacese  (Rushes).  The  regular,  hermaphrodite, 
hypogynous  flowers  have  3  +  3  brown, 
dry,  free  perianth  -  leaves  projecting 
like  a  star  during  the  opening  of  the 
flower ;  stamens  3  +  3  (seldom  3+0) 
and  3  carpels  united  into  one  gynceceum 
(Fig.  283);  the  ovary  is  3-  or  1-locular; 
there  is  as  a  rule  1  style,  which  be- 
comes divided  at  the  summit  into  3 

stigmas,  often  bearing  branches  twisted 

•    T       /-it-       r»oo\        -n      -±  7  Fift.  283.— Flower  of  Luzula. 

to  the  right  (Fig.  283).    Fruit  a  capsule 

with  loculicidal  dehiscence.     The  embryo  is  an  extremely  small, 

ellipsoidal,  cellular  mass,  without  differentiation  into  the  external 

organs. 


284  MONOCOTYLEDONES. 

Juncus  (Rush)  has  glabrous  foliage-leaves,  generally  cylindrical, 
rarely  flat ;  the  edges  of  the  leaf -sheath  are  free  ("  open  "  leaf- 
sheaths)  and  cover  one  another.  The  capsule,  1-  or  3-locular, 
with  many  seeds. — Luzula  (Wood-Rush)  has  flat,  grass-like  leaves 
with  ciliated  edges ;  the  edges  of  the  leaf-sheath  are  united 
("closed  "  leaf-sheath).  The  capsule  unilocular  and  3-seeded. — 
Prioninm  :  S.  Africa  ;  resembling  a  Tacona. 

The  interior  whorl  of  stamens,  in  some  species,  disappears  partially  or 
entirely  (J.  supinus,  capitatus,  conglomerates,  etc.) 

Some  of  the  numerous  Juncws-species  (e.g.  J.  effusus,  glaucus,  confjlomeratus, 
etc.),  have  false,  lateral  inflorescences,  the  axis  of  the  inflorescence  being  pushed 
to  one  side  by  its  subtending  leaf,  which  apparently  forms  a  direct  continua- 
tion of  the  stem,  and  resembles  it  both  in  external  and  internal  structure. 
The  foliage-leaves  of  this  genus  were  formerly  described  as  "  unfertile  stems," 
because  they  are  cylindrical,  erect,  and  resemble  stems,  and  consequently  the 
stem  was  said  to  be  "leafless":  J.  ejfusus,  glaucus,  conglomerate.  Stellate 
parenchymatous  cells  are  found  in  tbe  pith  of  these  stems  and  in  the  leaves. 
Other  species  have  distinct  terminal  inflorescences  and  grooved  leaves ;  J". 
bufonius  (Toad-rush),  compressus,  and  others.  The  inflorescences  most  often 
present  the  peculiarity  of  having  the  lateral  axes  protruding  above  the  main 
axis.  Their  composition  is  as  follows: — The  flowers  have  either  no  bracteoles, 
and  the  inflorescences  are  then  capitulate  ;  or  they  have  1-sevei'al  bracteoles. 
Each  branch  has  then,  first,  a  2-keeled  fore-leaf  placed  posteriorly  ("  basal- 
leaf  "),  and  succeeding  this  are  generally  several  leaves  borne  alternately  and  in 
the  same  plane  as  the  basal-leaf,  the  two  uppermost  (the  "  spathe-leaves  ")  being 
always  barren  ;  those  which  lie  between  the  basal-leaves  and  the  spathe-leaves 
are  termed  '•  intermediate-leaves."  If  only  branches  occur  in  the  axils  of  the 
basal-leaves,  then  the  succeeding  branches  are  always  borne  on  the  posterior 
side  of  the  axis,  and  form  a  fan  1 ;  if  the  hasal-leaf  is  barren,  and  if  there 
is  only  one  fertile  intermediate-leaf,  then  the  lateral  axes  are  always  on  the 
upper  side,  and  a  sickle'-libe  inflorescence  occurs ;  if  there  are  2  fertile 
intermediate-leaves,  then  a  dichasium  is  formed,  and  in  the  case  of  there 
being  several,  then  a  raceme,  or  spike. 

Juncacece  are,  by  several  authors,  classed  among  the  Liliiflorae,  but  there  are 
so  many  morphological  and  partly  anatomical  features  agreeing  with  the  two 
following  orders,  that  they  may,  no  doubt,  most  properly  be  regarded  as  the 
starting  point  of  these,  especially  of  the  Cyperacecc,  which  they  resemble  in  the 
type  of  flowe*s,  the  inflorescence,  the  type  of  mechanical  system,  and  the  stomata. 

POLLINATION  by  means  of  the  wind.  Cross-pollination  is  often  established  by 
protogyny.  J.  bufonius  has  partly  triandrous  and  cleistogamic,  partly  hexan- 
drous,  open  flowers. — DISTRIBUTION.  The  200  species  are  spread  over  the  entire 
globe,  but  especially  in  cold  and  temperate  countries  ;  they  are  seldom  found  in 
the  Tropics. — USES.  Very  slight ;  plaiting,  for  instance. 

Order  2.      Cyperacese.      The  majority    are  perennial  (seldom 

1  "  Fan  "  and  "  sickle  "  are  adopted  as  terms  for  these  inflorescences  from  the 
•German  "fachel"  and  " sichel." 


GLUMIFLORJ;.  285 

annual)  herbs  living  in  damp  situations,  with  a  sympodial  rhizome 
and  grass-like  appearance.  The  stems  are  seldom  hollow,  or  have 
swollen  nodes,  but  generally  triangular,  with  the  upper  internode 
just  below  the  inflorescence  generally  very  long.  The  leaves  are 
often  arranged  in  3  rows,  the  leaf-sheath  is  closed  (very  seldom 
split),  and  the  ligule  is  absent  or  insignificant.  The  flowers 
are  arranged  in  spikes  (spikelets)  which  may  be  united  into  other 
forms  of  inflorescences  (chiefly  spikes  or  racemes).  The  flowers 
are  supported  by  a  bract,  but  have 
no  bracteoles.  In  some  genera  the 
perianth  is  distinctly  represented 
by  six  bristles  corresponding  to 
six  leaves  (Figs.  284  A,  286  A)  ; 
in  others  it  is  represented  by  an 
indefinite  number  of  hairs  (Fig.  FlG>  284.-Diagram  of  structure  of: 
284  S),  and  very  frequently  it  is  -4  Scirpus  silvaticus;  B  Eriophorum 
altogether  wanting.  The  inner  an9ustifolium' 
whorl  of  stamens  is  absent,  and  the  flower  has  therefore  3  stamens 
(rarely  more  or  less  than  3),  the  anthers  are  attached  by  their  bases 
to  the  filament  (innate)  and  are  not  bifid  (Figs.  286).  Gynceceum 
simple,  formed  of  3  or  2  carpels ;  1  style,  which  is  divided  at  the 
extremity,  as  in  the  Juncacese,  into  3  or  2  arms ;  the  single  loculus 
of  the  ovary  contains  one  basal,  erect,  anatropous  ovule ;  the 
stigmas  are  not  feather-like.  Fruit  a  nut,  whose  seed  is  generally 
not  united  with  the  peiicarp.  The  embryo  is  small,  and  lies  at 
the  base  of  the  seed  in  the  central  line,  surrounded  on  the  inner  side 
by  the  endosperm  (Fig.  286  B).  On  germination  the  cotyledon 
not  remain  in  the  seed. 


A  regular  perianth,  with  6  scale-like  perianth-leaves  in  2  whorls,  is  found  in 
Oreobolus.  In  Scirpus  littoralis  the  perianth-leaves  are  spreading  at  the  apex, 
and  divided  pinnately. 

The  branching  of  the  inflorescence  is  often  the  same  as  in  the  Juncaceae,  and 
supports  the  theory  that  these  two  orders  are  related.  In  Rhynchospoia  and 
others,  the  "spikelets"  are  really  only  "  spike-like  "  and  to  some  extent  compound. 

A.     SCIRPES.     HERMAPHRODITE  FLOWERS. 

1.  Spikelets  cylindrical,  the  bracts  arranged  spirally  (in 
many  rows).  The  lower  ones  are  often  barren,  each  of  the  others 
supports  a  flower. — Scirpus  (Club-rush).  The  spikelets  are 
many-flowered ;  the  perianth  is  bristle-like  or  absent,  and  does  not 
continue  to  grow  during  the  ripening  of  the  fruit  (Fig.  286  A). 
Closely,  allied  to  this  is  Heleocharis,  with  terminal  spikes. — 


MONOCOTYLE  BONES. 


Eriophorum  (Cotton-grass)  differs  chiefly  in  having  the  perianth- 
hairs  prolonged,  and  forming  a  bunch  of  white,  woolly  hairs  (Fig. 
284  B). 

Cladium  and  Ehynchospora  (Beak-rush)  differs  especially  in  the /«<;- flowered, 
compound  spikelets  which  are  collected  into  small  bunches  ;  the  latter  has 
received  its  name  from  the  fact  that  the  lowermost  portion  of  the  style  remains 
attached  to  the  fruit  as  a  beak. 

2.  Spikelets  compressed,  the  bracts  arranged  only  in  two  rows ; 
the  other  characters  as  in  the  first-mentioned.  Cyperus  (spikelets 
many -flowered)  ;  Schosnus  (Bog-rush)  ;  spikelets  few-flowered ;. 
S.  nigricans  has  an  open  sheath. 

B.     CARICE^;.     UNISEXUAL  FLOWERS. 

In  the  (£ -flowers  there  is  no  trace  of  a   carpel,  and  in  the  ? 
no  trace  of   a  stamen.     Floral-leaves  in    many    rows.       In  some 
(Selena,  certain  (7are#-species) ,  <£-  and  $  -flowers  are  borne  in  the 
same  spikelet,  the  latter  at  the  base  or  the  reverse  ;  in  the  majority 
each  spikelet  is  unisexual. 

Carex  (Fig.  285)  has  naked,  most  frequently  monoecious  flowers. 
The  $ -spikes,  which  are  generally  placed  at  the  summit  of  the 
whole  compound  inflorescence,  are  not  compound ;  in  the  axil  of 
each  floral-leaf  (bract)  a  flower  is  borne,  consisting  only  of  a  short 


285. 


236. 


FIG.  285.— Carex  •.  ^A  diagram  of  a  male  flower  ;  B  of  a  female  flower  with  3  stigmas ;  C  of 
a  female  flower  with  2  stigmas ;  D  diagrammatic  figure  of  a  female  flower ;  E  similar  one  of 
the  androgynous  (false)  spikelet  of  Elyna.  The  <J  is  here  represented  placed  laterally;  it 
is  terminal,  according  to  Pax. 

FIG.  286.— A  Flower  of  Scirpus  lacustris.    B  Seed  of  Carex  in  longitudinal  section. 

axis  with  three  stamens  (Fig.  285  A).  The  ?  -spikes  are  compound ; 
in  the  axil  of  each  floral-leaf  is  borne  a  very  small  branch  (Fig. 
285  D,  a)  which  bears  'only  one  leaf,  namely,  a  2-keeled  fore-leaf 


GLUMIFLOR^E.  287 

(utriculus,  utr.  in  the  figures)  which  is  turned  posteriorly  (as. 
the  fore-leaves  of  the  other  Monocotyledons),  and  being  obliquely 
sheath-like,  envelopes  the  branch  (in  the  same  manner  as  the  sheath 
of  the  vegetative  leaves),  and  forms  a  pitcher-like  body.  In  the 
axil  of  this  leaf  the  ?  -flower  is  situated  as  a  branch  of  the  3rd 
order,  bearing  only  the  2-3  carpels,  which  are  united  into  one 
gynceceum.  The  style  protrudes  through  the  mouth  of  the  utri- 
culus.  The  axis  of  the  2nd  order  (a  in  Fig.  285  D)  may  sometimes  elongate  as 
a  bristle-like  projection  (normally  in  Uncinia,  in  which  it  ends  as  a  hook, 
hence  the  name) ;  this  projection  is  in  most  cases  barren,  but  it  sometimes 
bears  1-several  bracts  which  support  male-flowers ;  this  is  normal  in  Elyna 
(or  Kohresia)  and  Schaenoxiphium  ;  the  axis  (a  in  285  E)  bears  at  its  base  a 
female-flower  supported  by  the  utriculus,  and  above  it  a  male-flower  supported 
by  its  bract. 

POLLINATION  by  means  of  the  wind.  Protogynous.  Sometimes  self-pollinated. 
The  order  embraces  nearly  3,000  species,  found  all  over  the  world.  Carex 
and  Scirpus  are  most  numerous  in  cold  and  temperate  climates,  and  become 
less  numerous  towards  the  equator.  The  reverse  is  the  case  with  Cijperus 
and  other  tropical  genera.  They  generally  confine  themselves  to  sour,  swampy 
districts ;  some,  on  the  other  hand,  are  characteristic  of  sand-dunes,  such  as 
Sand-star  (Carex  arenaria).  There  are  about  70  native  species  of  Carex. 

USES.  In  spite  of  their  large  number,  the  Cyperaceae  are  of  no  importance 
as  fodder-grasses,  as  they  are  dry  and  contain  a  large  amount  of  silica ;  hence 
the  edges  of  many  of  the  triangular  stems  or  leaves  are  exceedingly  sharp 
and  cutting.  Ct/perus  esculentus  has  tuberous  rhizomes,  which  contain  a  large 
amount  of  fatty  oil  and  are  edible  (earth-almonds)  ;  it  has  its  home  in  the 
countries  of  the  Mediterranean,  where  it  is  cultivated. 

Cijperus  papyrus  (W.  Asia,  Egypt,  Sicily)  attains  a  height  of  several  metres, 
and  has  stems  of  the  thickness  of  an  arm  which  were  used  by  the  ancient 
Egyptians  fdr  making  paper  (papyrus).  Some  serve  for  plaiting,  mats,  etc. 
(Scirpus  lacustris,  etc.).  Isolepi?  is  an  ornamental  plant. 

Order  3.  Gramineae  (Grasses).  The  stems  are  cylindrical, 
generally  hollow  with  swollen  nodes,  that  is,  a  swelling  is  found  at 
the  base  of  each  leaf  which  apparently  belongs  to  the  stem,  but  in 
reality  it  is  the  swollen  base  of  the  leaf.  The  leaves  are  exactly 
alternate  ;  the  sheath  is  split  (excep.  .Bromws-species,  Poa  pratensis, 
P.  trivialis,  Melica,  Dactylis,  etc.,  in  which  the  sheath  is  not  split), 
and  the  edges  overlap  alternately,  the  right  over  the  left,  and  vice 
versa ;  the  ligule  is  nearly  always  well  developed.  In  general,  the 
flowers  are  hermaphrodite  ;  they  are  borne  in  spikelets  with  alternate 
floral-leaves,  and  the  spikelets  themselves  are  borne  in  either 
spikes  or  panicles.  The  two  (seldom  more)  lowest  floral-leaves  in 
each  spikelet  (Fig.  289  0Y,  nY)  are  barren  (as  the  covering-leaves 
in  many  umbels  and  capitula)  ;  these  are  termed  the  glumes.  The 

W.  B.  U 


288 


SIOXOCOTYLEDONES. 


succeeding  floral-leaves,  each  of  which  supports  one  flower  as  its 
bract,  are  called  the  outer  pales  (nl) ;  these  sometimes  each  bear  an 
"  awn  "  (a  bristle-like  body  which  projects  in  the  median  line  either 
from  the  apex  or  the  back)  ;  sometimes  the  upper  ones  are  barren. 
Each  flower  has  a  bracteole,  which  is  placed  on  the  inside  opposite 
the  main  axis  ;  it  is  thin,  binerved  or  two-keeled,  and  never  has  an 
awn;  it  is  known  as  the  inner  pale  (01).  Immediately  succeeding 
the  bracteole  are  :  («)  some  small,  delicate  scales  (lodicules,  Figs. 
287  D,  288  C,  290  L) ;  (6)  three  stamens  with  anthers  versatile, 
so  as  to  be  easily  moved,  and  usually  notched  at  each  end  (Fig. 

287  C) ;  and  (c)  a  simple  gynceceum  formed  of  one  carpel  with  two 
styles   having   generally   spirally -branched  stigmas    (Figs.    287    D, 

288  0).     The  ovary  is  unilocular,  and  contains  one  ascending  or 
pendulous,  anatropous  ovule.     Fruit  a  nut,  whose  seed  is  always 

firmly  united  with  the  thin  pericarp  ("caryopsis  ").     The  embryo  is 


FIG.  287.— TrUicum:  A  axis  (rachis)  uf  ear  showing  the  notches  where  the  spikelets 
wtre  inserted;  Ban  entire  spikelet;  Ca  flower  with  the  pales;  Da  flower  without  the 
pales,  showing  the  lodicules  at  the  base  ;  E  glume ;  F  outer  pale  ;  G  inner  pale  ;  H  fruit; 
J  longitudinal  section  of  fruit. 

larger  than  in  the  Cyperacere  and  is  placed  at  the  base  of  the  seed, 
but  on  the  outer  convex  surface  of  the  pericarp   (Figs.  287  I,  288 


GLIIMIFLOK2E. 


289 


288  D,  291),  outside  the  endosperm;  plumule  large  with  several 
leaf-primordia.  On  germination  the  cotyledon  remains  in  the  seed. 
The  majority  of  Grasses  are  annual  or  perennial  herbs  ;  tree- 
like forms  being  only  found  in  the  Tropics,  for  example,  the 
Bamboos;  they  branch  (in  tufts),  especially  from  the  axils  of  the 
basal -leaves,  while  those  which  are  borne  higher  on  the  stem  are 
separated  by  longer  internodes  and  have  no  vegetative  branches  in 


JI 


FIG.  289.— Diagramatic  outline  c.f 
a  spikelet:  n  T  lower  glume;  0  Y 
upper  glume;  n  I  upper  pale;  0  I 
the  inner  pale ;  Z-l  lodicules ;  it 
stamens;  I-I  main  azes;  II  lateral 
axes. 


FIG.  23?.— Bi-omus  mollis:  A  inflorescenco ; 
iB  the  uppermost  flower  of  a  spikelet,  with 
its  axis  turned  forward  j  in  front  is  seen  the 
two-keeled  inner  pale  (bracteole)  and  the 
stamens  protrude  between  this  and  the  outer 
rpale  (bract) ;  C  an  ovary  with  the  2  stigmas 
on  its  anterior  side,  the  2  lodicules,  and  the 
3  stamens ;  D  the  fruit  seen  from  the  dorsal 
•side;  JB  the  same  from  the  ventral  side. 

their  axils,  though  a  few  forms,  like  Bambusa   and   Calamagrostis 
lanceolata,  produce  branches  in  these  axils. 

Only  a  few  Grasses  have  a  solid  stem,  such  as  Maize,  Sugar-cane,  and  Andro- 
pogon.  The  blade  is  flat  in  the  meadow-grasses,  but  the  Grasses  which  live  on 
•dry  places  ("  prairie-grass  ")  exposed  to  the  sun,  often  have  the  blade  tightly 


290 


MONOCOTYL  EDONES. 


rolled  up  and  almost  filiform  or  bristle-like,  with  anomalous  anatomical 
structure.  A  closed  tubular  sheath  is  found  in  Melica  uniflora,  Bromus -species, 
Poa  pratensis  and  trivialis,  Briza  and  some  GZ?/cm"a-species.  The  sheath  is 
developed  for  the  purpose  of  supporting  the  young  internodes  while  their  growth 
is  proceeding  at  the  base.  The  "  nodes  "  (the  swollen  joints  which  are  seen  on 
stems  of  Grasses)  are  not  really  part  of  the  stem  but  are  formed  by  the  base  of 
the  leaf-sheath.  They  play  a  part  in  assisting  the  haulms  to  regain  a  vertical 
position  when  laid  prostrate  by  wind  or  rain.  The  awn  on  the  pale  is  homo- 
logous with  the  blade  of  the  Grass-leaf,  and  the  pale  itself  is  the  sheath.  The 
arrangement  of  the  leaves  in  the  spikelet  is  similar  to  that  in  Ci/perus  and  other 
Cyperaceae,  their  floral-leaves  being  borne  in  several  rows  in  Streptochceta.  More 
than  two  barren  "glumes"  are  found  in  Streptochcsta,  several  Phalarideae  and 
others.  The  spikelets,  too,  are  again  arranged  in  two  rows  in  the  axils  of  sup- 
pressed floral-leaves.  The  inflorescence  becomes  a  "  compound  spike  "  (ear;  when 


FIG.  290.— Diagram  of  the  Grass-flower  .- 
ni  outer  pale ;  0i  inner  pale ;  l-I  lodi- 
cules. 


FIG.  291.— Longitudinal  section  of  an 
Oat-grain .-  a  the  skin  (pericarp  and 
testa);  b  the  endosperm;  c  the  cotyledon; 
d  the  plumule. 


the  spikelets  are  sessile.  In  the  majority  of  instances  the  spikelets  are  borne 
on  long  stalks ;  when  these  branch,  then  the  secondary  branches,  and  similarly 
all  branches  of  higher  order,  are  placed  so  far  down  upon  the  mother-axis  that 
they  all  appear  to  be  of  equal  value  and  to  arise  in  a  semicircle  from  the  mother- 
axis  itself,  though  in  reality  they  arise  from  each  other  (Panicle,  Fig.  288  A). 
Sometimes  the  main  axis  and  branches  of  different  orders  unite  together  as  in 
Alipecurus,  Phleum,  and  some  other  Grasses,  and  hence  the  single  (short- 
stalked)  spikelets  appear  to  arise  singly  and  spirally,  or  without  any  definite 
order,  directly  from  the  main  axis,  with  the  production  of  a  cylindrical  inflor- 
escence bearing  "spikes"  on  all  sides,  that  is,  a  "spike-like  panicle." — Many 
inflorescences  are  somewhat  dorsiventral.  The  flower  is  rarely  unisexual  (Zea 
mc.is)  or  barren.  Considerable  difficulty  is  experienced  in  reducing  the  Grass- 


GLUMIFLORJ;.  291 

flower  to  the  ordinary  3-merous  Monocotyledonous  type.  Some  authorities 
consider  the  lodicules,  which  are  present  in  all  Grasses  but  absent  in  the 
Cyperaceaa,  to  be  homologous  with  a  perianth.  According  to  a  more  recent 
theory  they  are  bracteoles,  and  hence  the  Gramineaa,  like  many  of  the  Jun- 
caceaa,  have  2-3  bracteoles  placed  in  two  rows  in  the  median  plane.  If  this 
theory  be  correct,  the  flower  is  naked.  The  lodicules  expand  quickly  and 
cause  the  opening  of  the  flower  (i.e.  the  two  pales  become  separated  from  each 
other).  Generally  only  3  stamens  belonging  to  the  outer  whorl  are  present  (Fig. 
290),  as  in  Iris  (Fig.  279),  certain  Juncaceae  and  Cyperaceae  (Fig.  284),  but  in 
some,  such  as  the  Eice  and  certain  species  of  Bamboos,  all  6  are  found. 
Pariana  has  more  than  6.  Only  1  of  the  carpels  is  present,  namely,  th<>, 
anterior  (of  those  in  Fig.  284),  so  that  the  ventral  suture  and  the  place  of  at- 
tachment of  the  ovule  are  situated  at  the  back  of  the  ovary.  The  number  of 
styles  does  not  correspond  with  the  number  of  carpels,  and  the  styles  may 
therefore  be  supposed  to  arise  from  the  edges  of  the  leaf  to  the  right  and  left — a 
position  which  is  not  without  analogy.  In  addition,  a  stylar  projection  is  some- 
times found  on  the  anterior  side  and  in  the  median  line  (e.g.  in  Phragmites), 
and  the  solitary  style  in  Nardus  has  exactly  this  position  ;  a  similar  arrange- 
ment is  found  in  some  species  of  B  ambus  a  which  have  only  one  style  ;  other 
species  of  Barnbuna  have  three  styles.  A  tripartite  style  is  found  in  Pharus. 

[The  Grass-flower  may  be  reduced  to  the  ordinary  Monocotyledonous  type 
thus : — The  outer  pale  is  the  bract  of  the  flower  since  it  bears  in  its  axil  the 
floral  shoot ;  the  inner  pale  occupies  the  customary  position  of  the  bracteole. 
The  fact  that  it  is  binerved  can  be  explained  by  its  having  been  pressed  against 
tbe  main-axis  during  development.  Similar  binerved  bracteoles  are  found  in 
Iris  (Fig.  279).  These  bracteoles  in  both  Grass  and  Iris  arise  from  single  pri- 
mordia,  and  are  not  produced  by  the  coalescence  of  two  leaves.  The  lodicules 
are  the  only  parts  of  the  perianth  remaining,  the  outer  whorl  having  been  sup- 
pressed, and  also  the  posterior  leaf  of  the  inner  whorl ;  a  posterior  lodicule, 
however,  is  found  in  the  Eice  and  some  species  of  Bamboo.  The  outer  whorl  of 
stamens  is  usually  absent,  though  this  again  is  present  in  the  Eice  and  Bamboo. 
The  three  carpels  are  reduced  to  one  with  two  or  sometimes  three  stigmas.] 

THE  FLOWERING.  In  the  panicles  the  flowers  open  in  basipetal  order; 
the  flowers  in  the  spikes  situated  somewhat  above  the  middle,  commence  to 
open  first,  and  the  flowering  proceeds  upwards  and  downwards.  A  few  Grass- 
flowers  never  open  (cleistogamic) ;  Leersia  oryzoides,  Stipa-species,  and  e.g. 
Wheat  and  Eye  in  cold  damp  weather  ;  some  open  their  pales  so  wide  that  the 
anthers  and  stigmas  may  protrude  at  the  top  ;  most  frequently  the  lodicules 
expand  and  force  the  pales  suddenly  and  widely  apart.  The  filaments  elongate 
considerably,  so  that  the  anthers  are  pendulous  and  the  stigmas  unfold.  In 
some  Grasses,  e.g.  Wheat,  the  blooming  of  each  flower  only  lasts  a  short  time. 
POLLINATION  is  generally  effected  by  the  wind.  The  Eye  separates  the  pales 
very  widely  in  the  morning,  and  allows  the  anthers  and  stigmas  to  appear ; 
it  is  almost  entirely  sterile  when  self -pollinated.  The  Wheat  flowers  at  any 
time  of  the  day,  each  flower  lasting  only  a  quarter  of  an  hour.  The  pales  open 
suddenly,  but  only  half  way,  and  the  anthers  scatter  one-third  of  the  pollen  in 
their  own  flower  and  two-thirds  outside.  Self-pollination  is  effectual,  but 
crossing  gives  better  results.  In  Hordeum  vulgare  (all  flowers  £ )  tne  flowers 


292 


MONOCOTYLEDONES. 


of  the  4  outer  rows  behave  as  in  the  Wheat,  but  those  in  the  two  central  rows 
always  remain  closed.  The  £  -flowers  in  the  two  central  rows  of  H.  distichuni 
remain  closed  and  fertilise  themselves ;  they  open  exceptionally,  and  may  be 
pollinated  by  the  ^-flowers  in  the  4  lateral  rows.  H.  hexastichum  is  cleisto- 
gamic.  Gats  pollinate  themselves. 

The  ripe  Grass-fruit,  in  some  species  of  Bamboo,  is  a  berry  ;  in  some  other 
Grasses  a  nut  with  loosely  lying  seed,  in  some  even  a  capsule,  but  otherwise  a 
"caryopsis."  In  some  instances  it  is  loosely  enveloped  by  the  pales  (Oat),  in 
others  firmly  attached  to  these  (Barley),  and  finally,  in  others,  "naked,"  i.e.  it 
is  entirely  free  from  the  pales  (Wheat  and  Eye).  On  the  ventral  tide  there  is  a 


FIG,  292.— Barley  grain  :  A  sec- 
tion through  the  skin  (a-d)  and  the 
most  external  part  of  the  endo- 
sperm ;  Gl  the  "  aleurore  layer  "  ; 
st  starch-containing  cells;  B  starch 
grains. 


FIG.  293.  —  Wheat- 
grain  germinating:  g 
the  plumule;  b  the  first 
leaf  succeeding  the  co- 
tyledon ;  r1  the  primary 
root ;  r2  lateral  root. 


FIG.  294. -Older seed- 
ling of  the  Wheat :  s  the 
second  sheathing-leaf ; 
I  first  foliage-leaf. 


groove  (Fig.  288  F)  •  on  the  anterior  side  (dorsal  suture),  which  is  turned  towards 
the  inner  pale,  it  is  convex,  and  at  the  base  on  this  side,  inside  the  testa,  lies  the 
embryo  (Fig.  288  D).  The  apex  of  the  fruit  is  often  hairy  (Fig.  293).  The  xkin 
(Fig.  291,  a)  is  formed  by  the  pericarp  and  testa,  and  in  some  cases  (Barley)  the 
pales-  also  form  the  outer  portion.  The  endosperm  (b)  is  large,  and  formed  of 
parenchymatous,  starch-containing  cells  :  aleurone  (proteid)  grains  may  also  be 
found  among  them.  When  the  fctarch-grains  and  the  aleurone-grains  adhere 
together  the  endosperm  becomes  "  horny,"  but  is  "  floury  "  when  the  starch- 
grains  lie  loosely  with  air  between  them.  In  the  most  external  region,  just 
beneath  the  skin,  1-several  layers  of  nearly  cubical  cells  (filled  principally  with 
aleurone-grains  and  fat)  are  found,  the  aleurone-layer  (Fig.  292).  The  embryo 
(Fig.  291  c-d)  contains  large  quantities  of  fatty  oil ;  the  large  shield- like  structure, 
attached  to  the  embryo  and  turned  inwards  towards  the  endosperm  (c),  is  the 
cotyledon  (scutellum) ;  it  remains  enclosed  in  the  seed  during  germination,  and 


GLUMIFLOR.S:.  293 

dissolves  the  endosperm  by  means  of  the  peculiar  epithelial  cells  developed  on 
the  dorsal  surface.  The  radicle,  on  germination,  is  obliged  to  perforate  a  mass 
of  cells  derived  from  the  suspensor  and  which  form  the  "root-sheath" 
(coleorhiza,  Fig.  293)  round  its  base.  In  addition  to  the  tap-root,  lateral  roots 
are  frequently  developed  before  germination  ;  these  quickly  break  through, 
and  later  on  are  followed  by  others  which  appear  at  the  base  of  the  leaf  (Figs. 
293,  294). 

The  DISTRIBUTION  OF  THE  FRUIT  is  most  frequently  effected  by  the  wind.  The 
spirally-twisted  and  hygroscopic  awn  which  persists  on  the  fruits  of  some  species 
(Avena,  Stipa,  etc.)  assists  in  their  dissemination,  and  even  helps  to  bury  them 
in  the  ground. 

The  two  preceding  orders  are  more  closely  related  to  each  other  than  they 
are  10  the  Grammes. 

The  generic  differences  are  chiefly  founded  on  the  form  of  the  inflorescence, 
the  number  and  sex  of  the  flowers  in  the  spikelets,  the  shape  and  relative 
length  of  the  pales,  awns,  etc.  In  addition  to  these  the  structure  of  the  fruit 
and  seed  presents  a  great  many  differences  ;  some  have  compound  starch-grains, 
while  in  others  they  are  single  ;  some  have  1  layer  of  aleurone-cells,  others  have 
several  (Fig.  292),  etc. 

1.  BAMBUSE^E.     Tall  Grasses  with  woody,  very  siliceous  stems 
which  bear  many  branches  in  the  axils  of  the  leaves.     6  stamens. 
Bambusa  (Bamboo). 

2.  ORYZE.E.     Oryza  saliva  (Rice)  is  a  herbaceous  marsh-plant, 
with  panicle  and  small,  1-flowered  spikelets,  with  two  small  glumes 
and  two  large,  boat-shaped,  strongly  siliceous  pales.     6  stamens. 
— Leersia.     Lygeum.     Pliarus.     Zizania  aquatica. 

3.  MAYDILE.     Zea  mais  (Indian-corn,  Maize)  ;  the  spikelets  are 
unisexual ;  the  £  -spikelets  in  a  terminal  panicle  ;  the   ?  -spikelets 
closely  crowded  and  arranged  in  many  rows  in  a   thick,  axillary 
spike,  enclosed  by  large   sheathing-leaves.     The   ?  -spikelets  are 
l-(2-)  flowered ;  the  ovary  bears  one,  long,  filamentous  style,  with 
bifid  stigma. — Euchlsena ;   Coix. 

4.  ANDROPOGONEYE.     Saccharum  (Sugar-cane)  ;  the  spikelefcs  are 
exceptionally    small,    1-flowered,    and    borne    in    pairs   in    many- 
flowered,  long-haired  panicles.       Tall    grasses    with  solid,  sappy 
stem. — Andropogon. 

5.  FESIUCE^;.     Grasses  with  panicle  (or  spike-like  panicle)  and 
2-several-flowered  spikelets.     Glumes  small,  in  each  case  shorter 
than  the  spikelet. — Festaca    (Fescue)    and   Bromus   (Brome,   Fig. 
288)  have  the  awn  placed  at  the    apex  of  the  pale,  or  slightly 
below  it.     Festuca  has    perennial    species,    with    only  a  sparsely- 
branched  panicle  with  branches  solitary  or  in  pairs,  and  round 
spikelets ;    the    leaf-sheath    is    widely    open.       Bromus   has    the 


294  MONOCOTYLEDONES. 

branches  borne  in  half  whorls,  and  the  leaf-sheath  scarcely  half 
open.  ]3r  achy  podium  has  very  short-stalked  spikelets  in  a  raceme. 
— Poa  (Meadow-grass),  Briza  (Quaking-grass)  and  Glyceria  have 
awnless  spikelets  ;  these  in  Poa  are  ovoid,  compressed,  and  with 
sharply-keeled  glumes;  in  Briza  they  are  broad,  cordate  and 
drooping,  with  boat-shaped  glumes  ;  in  Glyceria  round,  long,  many- 
flowered,  linear  or  lanceolate  :  some  species  of  Glyceria  have  closed 
leaf-sheaths. — Dactylis  (Cock's-foot)  differs  from  all  others  in  the 
somewhat  crowded  and  unilateral  (subsecund)  spikelets,  which  are 
compressed  and  oblique  (i.e.  one  side  more  convex  than  the  other). 
— Phragmites  (P.  communis,  Reed);  the  lowermost  flowers  of  the 
spikelet  are  <§  ;  its  axis  is  covered  with  long,  silky  hairs ;  pales 
without  awns,  but  acuminate.  Perennial  marsh-plants. — Melica  ; 
panicle  small,  sparsely-branched  with  round,  awnless,  few-flowered, 
usually  drooping  spikelets.  The  upper  pales,  with  arrested  flowers, 
are  generally  united  into  a  club-like  mass. — Molinia,  Eragrost/s, 
Koeleria,  Catabrosa. — Cynosurus  (Dog's-tail)  has  a  small,  spicate 
panicle  with  unilateral  spikelets,  some  of  which  are  fertile,  some 
barren,  each  supported  by  a  pectinate  scale.  Arundo.  Sesleria. 
Gynerium.  Triodia. 

6.  AVENEJ;.  Panicles  with  2-many-flowered  spikelets  ;  at  least 
one  of  the  glumes  is  quite  as  long  as  the  entire  spikelet. — 
Avena  (Oat).  The  pale  is  boat-shaped,  often  bifid,  and  at  about 
the  middle  of  the  back  has  a  twisted,  bent  awn. — Aira  (Hair-grass) 
has  a  long-branched  panicle  with  small,  2-flowered  spikelets ;  the 
pale  has  a  dentate  apex  and  bears  an  awn  on  the  posterior  side 
close  to  the  base. —  Weing drtneria. — Holcus  (Yorkshire-fog)  ;  a  soft, 
hairy  Grass  with  an  open  panicle,  keeled  glumes  ;  2  flowers  in  the 
spikelet,  of  which  the  lower  one  is  $ ,  the  upper  <£  ;  the  pale 
which  supports  the  ^  -flower  has  no  awn,  but  that  which  supports 
the  ^-flower,  on  the  contrary,  is  awned. 

7-  AGROSTIDE.E.  Panicles  or  spike-like  panicles  with  1-flowered 
spikelets.  Generally  2  glumes  and  only  1  pale. — The  following1 
have  PANICLES  :  Milium  with  square  panicle- branches  and  round 
spikelets ;  Agrostis  (Fiorin),  with  compressed,  glabrous  spikelets, 
whose  glumes  are  longer  than  the  pales.  Calamagrostis  differs  in 
having  a  chaplet  of  long  hairs  at  the  base  of  the  pale. — Stipa 
(Feather-grass)  has  a  long,  twisted  awn. — The  following  have  SPIKE- 
LIKE  PANICLES  :  Phleum  (Cat's-tail,  Timothy- grass)  has  sharply 
pointed,  entirely  free  glumes,  which  are  much  longer  than  the 
awnless  pales.  Alopecurus  (Fox-tail) ;  glumes  united  below  ;  pale 


GLUMIFLOR^l.  295 

with  awn.  Ammophila  (Psammci).  A.  arundinacea ;  pales  hairy 
at  base;  perennial,  stiff-leaved,  glaucous  sand-grass  with  creeping 
rhizome.  Aristida.  Sporobolus. 

8.  PHALAEIDEJE.  Panicles  and  spike-like  panicles.  The  spike- 
let  has  in  the  upper  part  a  single  fertile  flower ;  below  it  are 
placed  4  pales,  of  which  the  upper  1-2  sometimes  support  <£- 
flowers.  On  the  whole,  6  floral-leaves  of  the  first  order  are 
present. — Phalaris  (P.  canariensis,  Canary-grass)  has  an  ovate, 
spike-like  panicle,  the  spikelets  are  compressed,  convex  on  the 
outer  side,  concave  on  the  inner.  The  large  glumes  are  winged 
on  the  back. — Digraphis  (D,  arundinacea)  is  closely  allied  to 
Phalaris,  but  the  keel  of  the  glumes  is  not  winged. — Anthoxanthum 
(A.  odoratum,  Sweet-vernal)  has  a  small,  lanceolate,  open,  spike-like 
panicle;  the  spikelets  have  below  2  barren  flowers,  and  a.bove 
these  an  ty  -flower  with  2  stamens.  The  upper  glume  is  longer 
than  the  flower. — Hierochloa. 

g.  CHLORIDES.  The  spikelets  are  arranged  in  the  form  of  a  spike  in  two 
rows  on  one  side  of  an  often  flatly-compressed  axis;  they  are  mostly  1-flowered. 
— Chloris  ;  Cteniuin ;  Cyuodon;  Eleusine ;  Microchloa. 

10.  PANICE/E.     The  spikelets  are  borne  in  panicles  or  spikes,  which  may  be 
arranged  like  fingers  or  in  a  raceme.      There  is  a  centrally-placed  £  -flower  ; 
below  it  is  sometimes  a  $  -flower. — Panicum  ;  Paxpulum;  Oplismenus ;  Setaria 
has  an  almost  cylindrical   spike-like  panicle  with  i-everal  barren  branchlets, 
which  project  as  stiff,  rough  bristles. — Cenchnts ;  Pennisetum. 

11.  HORDE M.    Spikes  compound  ;  spikelets  sessile  in  the  notches 
of  a  toothed  axis. 

A.  Spikelets  solitary. — Triticum  (Wheat,  Fig.  287)  has  in  each 
tooth  of  the  main  axis,  a  several-flowered  spikelet  which  turns  its 

flat  side  towards  the  central  axis.  The  cultivated  species  (true 
Wheat)  are  1-2- annual,  the  wild  ones  (T.  repens,  Couch,  also  as 
an  independent  genus,  Agropyrum)  are  perennial,  with  creeping 
rhizome  and  lanceolate  glumes. — Lolium  (Rye-grass)  has  in  each 
tooth  of  the  main  axis  a  many-flowered,  compressed  spikelet, 
which  is  placed  edgewise  towards  it  and  (with  the  exception  of 
L.  perenne)  has  only  one  outwardly-turned  glume  (L.  temulentum 
has  a  rudiment  of- the  inwardly-turned  lower  glume) ;  the  terminal 
spikelet  has  two  glumes. —Secale  (Rye).  A  two-flowered  spikelet 
in  each  tooth  ;  small,  lanceolate,  acuminate  glumes.  Nardus  and 
Lepturus  have  very  narrow  spikes,  the  former  with  unilateral 
spikelets. 

B.  In   each   notch    of  the   axis  2  or  more  spikelets  are  placed 


296  MONOCOTYLEDONES. 

close  together. — Hordeum  (Barley).  In  each  tooth  three  1 -flowered 
spikelets.  H.  liexasticlium  (6-rowed  Barley),  has  6  rows  of  fruits, 
since  all  the  spikelets  are  fertile,  and  H.  distichum  (2-rowed  Barley) 
2  rows,  since  the  lateral  spikelets  are  <^,  and  barren  (p.  292). — 
Elymus  has  2-6  many-flowered  spikelets  in  each  joint  of  the  main 
axis.  JEgilops  has  awns  upon  the  glumes. 

DISTRIBUTION.  315  genera  with  3,500  species.  The  order  is  distributed 
over  the  wbole  world,  and  as  regards  number  of  individuals  is  perhaps  the 
richest.  In  the  Tropics,  large,  broad-leaved,  tree-like  forms  are  found  (Bam- 
busece,  Olyrece,  Andropogonece,  etc. ;  in  S.  Europe,  Arundo  donax) ;  in  England, 
next  to  the  Composite,  it  is  the  order  most  rich  in  species  (about  134). — The 
origin  of  some  of  the  cultivated  Grasses  is  lost  in  obscurity.  The  Maize,  no 
doubt,  was  indigenous  to  America,  where  its  nearest  relatives  are  found,  and 
where  it  has  also  been  discovered  in  ancient  Indian  graves  ;  Durra  or  Guinea- 
corn,  Millet  and  Sugar-cane  are  South  Asiatic  plants,  and  our  own  cereals  no 
doubt  have  sprung  primarily  from  Western  Asia  and  South-Eastern  Europe 
(Barley  from  Armenia  and  Persia,  where  a  very  closely  related  wild  species 
is  found ;  Wheat  from  the  same  districts  ;  Kye  from  the  perennial  species 
S.  montanum).  Panicwn  altissimum  and  Rice  have  come  from  Africa. 

USES.  The  Grasses  play  a  very  important  part  as  cereals  and  fodder  plants. 
The  following  are  the  most  important  of  the  cultivated  ones:  Triticum  vulgare 
(common  Wheat),  turgidum,  amijleum,  polcnicum,  ppelta,  durum,  etc.  ;  Secale 
cere.ale  (Rj'e)  ;  Barley  (Hordeum-species,  see  under  the  genus) ;  Maize ;  0>its 
(Avena  sativa,  orientalis,  nuda)  ;  Millet  (Panicum  miliaceum) ;  Durra  (Turkish 
Millet,  or  Guinea  corn,  Sorghum  vulgare,  cernuum  and  saccharatuni) ;  Manna- 
grass  (Glyceria  fluitarw).  As  fodder-plants  especially :  Rye-grass  (Lolhun 
perenne) ;  Oat-grass  (Avf.na  elatior) ;  Timothy  (Phleum  pratense)  ;  Fox-tail 
(Alopccurus  pratensis) ;  Cock's  foot  (Dactylic  glomerata) ;  Dog's  tail  (Cyno&uru* 
cristatus) ;  Sweet-vernal  (Anthoxanthum  odoratum) ;  Soft  grass,  or  Yorkshire-fog 
(Holcus  lanatus  and  moll  is)  ;  Quaking-grass  (Briza  media)  ;  species  of  Meadow- 
grass  (Pod) ;  Fescue  (Festuca)  and  Brome  (Bromu*).— Several  cultivated  species 
of  Grass  are  also  used  in  the  preparation  of  fermented  liquors,  the  starch  in 
the  seeds  being  transformed  to  sugar  (beer  from  "  Malt,"  i.e.  the  germinated 
Barley  ;  arrack  from  Rice) ;  or  the  stem  becomes  specially  saccharine  before 
flowering:  the  Sugar-cane,  Sorghum  sacchai atuin. 

OFFICINAL.  The  rhizome  of  Triticum  repeiis.  Oat-grain,  flour  of  Barley,  and 
the  starch  of  Wheat,  also  sugar. 

The  seeds  of  Lolium  temulentum  are  considered  poisonous. — The  stems  of 
many  species  (including  our  common  grains)  are  used  in  the  manufacture 
of  paper,  especially  "  Esparto  grass "  (Stipa  tenacissima)  from  Spain  and  N. 
Africa,  and  the  sheathing-leaves  of  the  ?  -spike  of  Maize.  Sand  Lyme-grass  (Ely- 
mus arenarius),  and  especially  Psamma  arenaria,  are  important. — But  few  Grass- 
species  are  sweet-scented :  AnthoxantJium  odoratum  and  Hicrocliloa  odorata 
contain  coumarin  ;  Avdntyoyon-species  have  essential  oils  (''  Citronella  oil"). 
— OBNAMENTAL  PLANTS  are  :  the  "  Ribbon-grass"  (a  variety  of  Digraphis  arundi- 
nacea),  Stipa  pennata  (whose  awn  is  exceedingly  long  and  feathery),  Gynerium 
firgenteum  (Pampas-grass),  Lagurus  ovatus,  Hordeumjulatum,  Bromus  brizi- 
formis. 


SPADICIFLOE^:. 


297 


Family  3.     Spadiciflorse. 

The  primitive  form  resembles  that  of  the  preceding  family.  In 
it  we  find  the  typical,  perfectly  developed,  Monocotyledonous 
flower,  sometimes  even  with  free  carpels  and  with  a  dry  or  some- 
what fleshy,  but  never 
petal oid  perianth  ;  and  this 
passes  over  into  very  differ- 
ent forms  by  the  suppres- 
sion of  the  floral-leaves, 
perianth  and  sporophylls 
(unisexual  flowers  are 
common),  and  by  the  close 
aggregation  of  the  flowers 
in  the  inflorescence.  The 
flower  is  hypogynout  in 
every  case.  The  inflores- 
cence is  a  spike  which 

may  be    either    single    or 

-,      -.          j    -,  p,  FIG.  295. — Pia8savtt(Jttal6a/unir0ra). 

branched,  and  has  otten  a 

thick  and  fleshy  axis  (a  spadix).  In  Palms  and  Aracese  it  is  en- 
veloped, at  any  rate  prior  to  the  opening  of  the  flowers,  by  a  very 
large  floral-leaf,  the  spathe,  which  may  be  petaloid  (Figs.  297,  301). 

The  fruit  is  most  frequently  fleshy  (berry,  drupe}  or  a  nut,  never 
M.  capsule.  The  embryo  is  small,  with  large,  fleshy  endosperm 
(Fig.  299  C)  ;  very  rarely  the  endosperm  is  wanting. 

The  numerous  plants 
belonging  to  this  family 
are  large,  herbaceous  or 
tree-like,  and  the  leaves 
seldom  have  the  usual 
Monocotyledonous  form, 
i.e.  linear  with  parallel 
venation,  but  most  fre- 
quently have  pinnate  or 
palmate  venation. 

Order    1.         Palmse 
(Palms).      The   majority 
are     trees    with    an     un- 
branched,  cylindrical  stem,      pj&  296>_A portion of  the  stemof  Attalea fun1fm 
having      short     internodes  with  persistent  leaf-bases. 


298  MONOCOTYLEDONES.   ' 

and  covered  with  leaf-scars  or  the  bases  of  the  leaf-stalks  (Fig. 
296),  and  at  the  summit  a  rosette  of  large  leaves  closely  packed 
together  (Fig.  295).  An  exception  to  this  is  found  in  Calamus 
(Cane,  "Rotang"),  whose  thin,  creeping  or  climbing  stems  have 
long  internodes  ;  sparsely l  branched  is,  e.g.  the  African  Doum-palm 
(Ilyphcene).  Notwithstanding  their  often  enormous  stems  the 
Palms  have  fibrous  roots,  like  the  bulbous  Monocotyledons.  The 
leaves  are  pinnate  (Feather-palms,  Fig.  298)  or  palmate  (Fan- 
palms,  Fig.  295)  and  often  very  large;  they  have  a  well-developed 
petiole  with  an  amplexicaul  sheath,  which  is  often  more  or 
less  separated  into  a  large  number  of  fibres.  In  the  bud  the  blade 
is  entire  but  folded,  as  the  leaf  expands  the  lines  of  folding  are 
torn,  either  those  which  are  turned  upwards  (thus  V  V  V  V ,  e.g. 
Pritchardia,  Livistona,  Phoenix,  Chamcerops)  or  those  turned  down- 
wards (thus  A  A  A  A,  e.g.  Gocos,  Chamcedorea,  Calamus).  The 
inflorescence  is  usually  lateral ;  when,  as  in  Sago-palm  (Me- 
troxylon  rumphii)  or  Talipot  (Corypha  umbraculifera)  it  is  termi- 
nal, the  plant  is  monocarpic,  and  dies  after  flowering;  it  is 
often  a  very  large  and  branched  spadix  with  numerous  flowers 
either  borne  externally  or  embedded  in  it,  and  enclosed  either  in 
one  woody,  boat-shaped  spathe  (Fig.  2Q7)  or  several  spathes,  in 
the  latter  case  one  for  each  branch.  The  flowers  are  sessile  or 
even  embedded,  regular,  generally  unisexual  (monoecious  or 
dicecious)  with  the  usual  diagram  (Fig.  278) ;  the  perianth  is  in- 
conspicuous, green  or  yellow,  persistent,  and  more  or  less  leathery 
or  fleshy.  6,  rarely  3  or  many  stamens.  The  3  carpels  remain 
either  distinct  or  form  one,  generally  3-locular,  ovary.  The  style 
is  short.  There  is  one  ovule  in  each  carpel.  Often  during  ripening 
2  carpels  with  their  ovules  are  aborted.  The  fruit  is  a  berry, 
drupe  or  nut,  generally  one-seeded,  with  a  large  horny  or  bony 
endosperm  with  hard  thick-walled  cells  (e.g.  Date-palm).  In 
some  (e.g.  Cocoanut)  it  is  thin-walled,  soft,  and  oily;  in  several 
"  ruminate." 

When  germination  commenced  in  the  Cocoanut,  Date,  etc.,  the  apex  of  the 
cot  \ledon  remains  in  the  seed  and  developesinto  a  spongy  mass  to  withdraw  the 
endosperm ;  in  the  Cocoanut  it  attains  a  considerable  size  (Fig.  299  C)  and 

1  [Although  unbranched  stems  are  characteristic  of  the  Palms,  yet  branched 
specimens  are  recorded  from  some  eleven  genera.  The  branches  are  developed 
from  lateral  buds,  which  in  many  instances  only  develope  when  the  terminal 
bud  has  been  destroyed.  A  few  Palms  develope  axillary  branches  at  the  base 
of  the  stem ;  these  form  rhizomes,  and  give  rise  to  clusters  of  aerial  stems.] 


SPADICIFLOR.E. 


299 


assumes  the  form  of  the  fruit.  The  endosperm  in  the  Cocoanut  is  hollow 
and  the  interior  is  filled  with  "milk."  In  the  Date-palm  and  the  Vegetable- 
ivory  (Phytelephas)  the  cell-walls  of  the  hard  endosperm  serve  as  reserve 
material. 

I.     PH<ENICEJ:.     Phoenix   (Date-palm)  has  pinnate  leaves  with 
channeled  leaflets   and  dioecious  flowers  with  3  free  carpels,   of 


' 


FIG.  297.— Inflor- 
escence of  a  Palm 
with  spathe.  At  the 
top  <?-,  at  the  base 
? -flowers. 


FJG.  208. — Livistona  australis. 


wKich  usually  only  one  developes  into  a  berry  with  membranous 
endocarp  ;  the  large  seed  has  a  deep  furrow  on  the  inner  side,  and 
horny  endosperm. 

2.      SABALE.S;.     These  have  fan-like  leaves  with  channeled  seg- 


300 


MONOCOTYLEDONES. 


ments  ;  flowers  ^  or  polygamous,  rarely  dioecious,  with  3  separate 
or  only  slightly  united  carpels,  all  of  which  are  sometimes 
developed  into  fruits  (berry  or  drupe,  with  thin  stone). — 
Chamcerops,  the  Dwarf-palm.  The  pericarp  is  externally  fleshy, 
internally  more  fibrous,  and  provided  with  a  membranous  inner 
layer.  The  endosperm  is  ruminate  (that  is,  the  testa  is  several 
times  deeply  folded  into  the  endosperm).  —  Sdbal,  Copernicia, 
Livistona  (Fig.  298),  Thrinax,  Corypha,  Brahea,  and  others. 


^  IG.  299.— A  Longitudinal  section  of  a  Cocoannt  (diminished),  the  inner  layer  only  (the 
stone,)  not  being  divided  B  End  view  of  the  stone,  showing  the  sutures  for  the  3  carpels 
(a),  and  the  3  germ-pores ;  the  embryo  emerges  from  the  lowest  one  when  germination 
begins.  C  Germinating ;  inside  the  stone  is  seen  the  hollow  endosperm  and  the  enlarging 
cotyledon. 

3.  COCOIJSIE^E.  With  pinnate  leaves.  Monoecious  inflorescence. 
The  carpels  are  united  into  a  3-loo.ular  ovary.  The  fruit  is  most 
frequently  1-locular,  only  1  of  the  loculi  becoming  developed, 
rarely  3-locular ;  it  is  a  drupe  with  a  large,  fibrous,  external  layer 
(mesocarp)  and  most  frequently  a  very  hard  inner  layer  (endocarp, 
stone)  which  has  3  germ-pores,  the  2  of  .these,  however,  which 
correspond  to  the  suppressed  loculi  are  closed;  internal  to  the 
third  lies  the  small  embryo  (Fig.  299).  Endosperm  containing 


SPADICIFLOR^;.  301 

abundance    of   oil.      Cocas    (the    Cocoannt-palm),    Attalea,   Elceis, 
Acrocomia,  Bactris. 

4.  LEPIDOCARYIIO;.     The  floral-leaves  and  flowers  are  borne  in 
2  rows  on  the  spadix.     The  carpels  are  united  into  one  3-locular 
ovary ;  the  fruit  is  coated   by  a  layer  of  hard,  shining,  imbricate 
scales.      The    majority   of  the   species  are  thorny,  and  climb  by 
means  of  the  thorny  leaves.    Some  have  fan-like  (Mauritia),  others 
pinnate    leaves    (Raphia,    Calamus,    Eugeissonia,    Metroxylon;    the 
stems  of  the  latter  die  after  the  first  flowering). 

5.  BORASSIN^E.      Large  Fan-palms  without  thorns,  with  3-locular   ovary. 
Drupe   with   separate   stones.     I.atania   and   Lodoicea  have   many   stamens ; 
Hyplicene',  Borassus  (Palmyra-palm). 

6.  ARECINEJE.     The  most  numerous  group.    Feather-palms.    Berry.     Areca, 
Euterpe,  Oreodoxa,  Ceroxylon,  Chamcedorea,  Geonoma,  Caryota  with  bi-pinnate 
leaves. 

7.  PHYTE^EPHANTIN^E.     Flowers  with  rudimentary  perianth  united  in  close 
capitula.     Pliyteleplias  (Vegetable-ivory).     Nipa. 

DISTRIBUTION.  About  1,100  species  are  known.  In  Europe  only  the  Dwarf- 
palm  (Chamcerops  humilis)  is  wild  (Western  Mediterranean).  The  Date  palm 
(Phce.nix  dactylifera)  belongs  to  North  Africa  and  West  Asia.  Other  African 
genera  are  Hyphcene  (Doum-palm)  and  Elceis  (E.  guineensis,  Oil-palm).  A 
large  majority  of  the  genera  are  found  in  South  America  and  in  the  East 
Indies.  The  following  are  AMERICAN: — Mauritia,  Acrocomia,  Bactris,  Chamce- 
dorea,  Oreodoxa,  Euterpe,  Attalea,  etc.  ASIATIC  : — Metroxylon,  Calamus,  Areca, 
Borassus,  Lodoicea  ("Double -cocoanutp,"  Seychelles)  and  others.  The  Cocoa- 
nut-palm  has  perhaps  an  American  origin  ;  all  the  other  species  of  the  same 
genus  being  endemic  in  America  ;  it  is  the  only  Palm  found  on  the  coral  islands 
of  the  Pacific  Ocean,  and  is  also  the  ouly  one  which  is  common  to  both  hemi- 
spheres. 

USES.  Palms  belong  to  the  most  useful  plants  ;  they  contain  no  poison,  and 
are  of  little  medicinal  interest,  but  are  largely  employed  in  the  arts  and  manu- 
factures, the  hard  timber  being  adapted  for  many  purposes  on  account  of  the 
hard  tissue  in  which  the  vascular  bundles  are  embedded.  "  Cane  "  is  the 
stem  of  Calamus-species  (from  India).  SAGO  is  obtained  from  the  pith  of 
Metroxylon  rumphii  (Sago-palm,  Sunda-Is.,  Moluccas),  Mauritia  flexuosa,  etc. 
Sugar-containing  sap  ("  palm  wine  ")  is  obtained  from  the  American  Mauritia 
vinifera  and  flexuosa,  Borassus  fiabelliformis  (Asiatic  Palmyra-palm),  Arenga 
saccharifera,  etc.,  by  cutting  off  the  young  inflorescences,  or  by  perforating  the 
stem  before  the  flowering  (arrack  is  distilled  from  this).  Vascular  strands  for 
the  manufacture  of  mats  and  brushes,  etc.,  are  obtained  from  the  outer  cover- 
ing (mesocarp)  of  the  Cocoanut,  and  from  the  detached  leaf-sheaths  of  Attalea 
funifera  (Brazil)  (Fig.  296).  WAX  is  yielded  by  the  leaves  of  Copernicia  cerifera 
(caruaiiba-wax,  Amazon  region),  and  by  the  stem  of  Ceroxylon  andicola 
(palm-wax,  Andes) ;  East  Indian  Dragon's  blood  is  from  the  fruit  of  Calamus 
draco  ;  the  young  buds  of  many  species,  especially  Euterpe,  Cocos,  Attalea, 
64,0.,  are  used  as  "  cabbage."  Palm-oil  is  obtained  from  the  oily  mesocarp 


302  MONOCOTYLEDONES. 

of  the  plum-like  fruits  of  Elceis  guineensis  (W.  Africa),  and  from  the  seeds, 
when  it  is  largely  used  in  the  manufacture  of  soap.  EDIBLE  FRUITS  from 
tbe  Date-palm  (Phoenix  dactylifera,  Arabia,  Egypt,  W.  Africa),  and  the 
endosperm  of  the  Cocoa-nut  (Cocos  nucifera).  The  seeds  and  the  unripe  fruits 
of  the  Areca-palm  (Areca  cateclni)  are  chewed  with  the  leaves  of  the  Betel- 
peper,  principally  in  Asia.  VEGETABLE  IVORY  from  the  hard  endosperm  of 
Phytelephas  macrocarpa  (S.  America.) — Many  species  are  cultivated  in  the  tropics 
as  ornamental  plants,  but  in  this  country  only  Chamcerops  humilis,  Livistona 
australis  and  chinensis  are  generally  grown.  In  addition  to  the  few  just 
mentioned,  many  others  are  of  importance,  but  these  are  much  the  most 
useful. 

Order  2.  Cyclanthaceae.  This  is  a  small  order  related  to  the  Pahus  (44 
species  from  Tropical  America),  with  fan-like,  folded  leaves.  The  flowers  are 
unisexual  and  arranged  in  whorls  or  close  spirals  on  an  unbranched  spadix. 
Ovary  unilocular,  ovules  numerous.  To  this  belongs  Carludovica  palmata, 
whose  leaves  are  used  for  Panama  hats. 

Order  3.  Pandanaceae  (Screw-pines)  is  another  small  order,  forming  a 
transition  to  the  Araceae.  The  woody,  (apparently)  dichotomous  stem  is 
supported  by  a  large  number  of  aerial  roots,  which  sometimes  entirely  support 
it  when  the  lower  portion  of  the  stem  has  decayed.  The  leaves  are  closely 
crowded  together,  and  arranged  on  the  branches  in  three  rows,  which  are  often 
obliquely  displaced,  with  the  formation  of  three  spiral  lines ;  they  are,  as  in 
the  Bromeliaceae,  amplexicaul,  long,  linear,  the  edge  and  lower  midrib  often  pro- 
vided with  thorns.  The  $  -flowers  are  borne  in  branched,  the  ?  in  un- 
branched spadices  or  capitula,  which  resemble  those  of  Sparganium,  but  have 
no  floral-leaves.  Perianth  absent.  The  drupes  or  berries  unite  into  multiple 
fruits.— About  80  species  in  the  islands  of  the  Indian  Ocean. — Pandanus, 
Fieycinetia. — Fossils  perhaps  in  the  chalk  of  the  Harz. 

Order  4.  Typhacese.  The  flowers  are  unisexual,  monoecious^ 
and  borne  on  a  cylindrical  spike  or  globose  capitulura  ;  $ 
inflorescences  above,  the  $  below.  The  perianth  consists  of  a 
definite  number  of  scales  (^Sparganium),  or  in  its  place  numerous 
irregularly-arranged  hairs  are  found  (Typha)  ;  in  the  <$  -flower 
there  are  generally  three  stamens ;  the  gynceceum  is  formed  of  1-2" 
carpels  with  1  prolonged  style  ;  1  pendulous  ovule.  The  seeds  are 
furnished  with  a  seed-cover,  which  is  cast  off  on  germination. — 
The  few  species  (about  20)  which  belong  to  this  order  are  marsh 
plants  with  creeping  rhizome  (and  hence  grow  in  clusters)  ;  the 
leaves  on  the  aerial  shoots  are  borne  in  two  rows,  entire,  very 
long  and  linear. 

Sparganium  (Bur-reed).  The  flowers  are  borne  in  globose 
capitula ;  the  perianth  distinct,  generally  consisting  of  3  small 
scales  ;  pistil  bi-carpellate.  Drupe,  dry  and  woody.  The  stalk  of  the- 
lower  $  capitula  is  sometimes  united  with  the  main  axis,  and  consequently  the- 
capitula  are  situated  high  above  their  subtending-leaf. 


SPADICIFLOEJ;.  303 

Typha  (Bulrush,  Reed-mace)  has  a  long,  cylindrical,  brown 
spike,  the  lower  portion  bearing  ?  -flowers,  and  the  upper  £  -flowers, 
which  is  divided  into  joints  by  alternate  leaves.  The  ?  -flowers 
have  1  carpel.  The  perianth  is  wanting,  represented  by  a  number 
of  fine,  irregularly-placed  hairs  ;  pistil  unicarpellate.  Fruit  a  nut. 

The  two  genera,  according  to  some,  are  related  to  the  2nd  order  In  both 
genera  native  species  are  found.  The  pollination  is  effected  by  the  wind,  and 
consequently  the  anthers  project  considerably,  and  the  stigma  is  large  and 
hairy.  Typha  is  protandrous,  Sparganium  protogynous.  The  small,  fine 
hairs  surrounding  the  nut  of  Typha  assist  in  its  distribution  by  the  wind. — 
Fossil  Tijphas  in  the  Tertiary. 

Order  5.  Araceae  (Arums).  The  flowers  are  small,  and  always 
borne  without  bracts  or  bracteoles  on  an  unbranched,  often  very  fleshy 
spike,  which  is  enclosed  by  a  spathe,  often  petaloid  and  coloured 
(Fig.  301).  The  fruit  is  a  berry.  Outer  integument  of  the  seed 
fleshy. — The  leaves  have  generally  sheath,  stalk,  and  blade  with 
distinctly  reticulate  venation  ;  they  are  chiefly  cordate  or  sagittate- 
(Fig.  302),  seldom  long  with  parallel  venation  as  in  the  other 
Monocotyledons  (Acorus,  Fig.  300).  The  Aracese  are  quite 
glabrous,  generally  perennial  herbs  with  tubers  or  rhizomes.  Many 
have  latex. — For  the  rest  the  structure  of  these  plants  varies  ;  for 
example,  while  some  have  a  perianth,  in  others  it  is  wanting ;  in 
some  the  perianth-leaves  are  free,  in  others  united ;  some  have 
hermaphrodite  flowers,  but  the  majority  unisexual  (monoecious)  ; 
some  have  free,  others  united  stamens ;  the  ovules  are 
orthotropous,  anatropous,  or  campylotropous,  erect  or  pendulous  ; 
the  ovary  is  1-many-locular ;  some  have  seeds  with  endosperm, 
others  without.  In  habit  there  are  great  differences.  While  some,  e.g.  Colo- 
casia  (Fig.  302),  have  a  thick,  more  or  less  upright  stem,  with  leaf-scars,  but  not 
woody,  others  are  climbers,  epiphytic,  and  maintain  themselves  firmly  by  means 
.of  adventitious  roots,  on  the  stems  and  branches  of  trees,  or  even  on  steep  rocks, 
e.g.  PhUodendron  ;  the  cordate,  penninerved  leaf  is  the  most  common  (Fig. 
302),  but  various  branched  forms  appear  ;  the  pedate  leaves  of  Helicophyllum, 
Dracunculus,  etc.,  are  cymosely  branched  ;  the  leaves  of  Monstera  deliciosa, 
perforated  by  tearing,  should  be  noticed  (the  vascular  bundles  while  ia  the 
bud  grow  faster  than  the  tissue  between  them,  causing  the  latter  to  be  torn, 
and  the  leaf  perforated).  With  regard  to  the  anatomical  structure,  the 
presence  or  absence  of  latex,  raphides,  resin-passages,  groups  of  mucilage- 
cells  should  be  noted.  Engler  makes  use  of  these  anatomical  peculiarities  for 
a  scientific  arrangement  of  the  order. 

A.  ORONTIE&,  CALAMUS-GROUP.  $,  hypogynous  flowers  of  a 
completely  formed  monocotyledonons  type  (number  in  the  whorls 
2,  3,  or  4).  —  Acorus  (A.  calamus,  Sweet- flag)  has  a  regular, 

W.  B.  X 


304 


MONOCOTYLEDONES. 


3-merous,  pentacyclic  flower 
(Fig.  300  (7,  D).  They  are 
marsh-plants,  with  creeping 
rhizome,  triangular  stem, 
and  long,  sword-like  leaves 
(Fig.  300  A)  i  the  inflor- 
escence is  terminal,  appa- 
rently lateral,  being  pushed 
to  one  side  by  the  upright, 
sword-like  spathe  (Fig.  300 
.— Anthurium  (Pr2  +  2,  A2+2, 
G2);  Pothos;  Orontium  (unilocular 
ovary  with  one  ovule),  etc. 


'h      h 


FIG.  300. — Acorus  calamus:  A  habit  (much 
reduced) ;  B  inflorescence  ;  C  a  flower ;  D 
•diagram ;  E  longitudinal  section  of  an  ovary ; 
F  an  ovule. 


A 


FIG.  301. — Arum  maculatum.  The 
Bpnthe  (h)  in  B  is  longitudinally 
divided. 


SPADICIFLOR.3S. 


305 


B.  CALLED.      Flowers    hypogynous,  naked,     $ .  —  Calla    (C. 
palustris).     All  flowers  in  the  spike  are  fertile,  or  the  upper  ones 
are  £  ;  6-9  stamens  ;  ovary  unilocular  with  many  basal  ovules. 
Marsh-plants  with  creeping  rhizome  and  cordate  leaves. — Honstera, 
RJiaphidophora,  etc. 

C.  AKINEJ;.    Flowers  monoecious,  naked,  <$  -flowers  on  the  upper, 
?  on  the  lower  part  of  the  spadix. — Arum  (Fig.  301).     The  spadix 

terminates   in   a   naked,  club-like   portion  (&)  ;    below  this   is    a 
number  of  sessile  bodies  (rudimentary  flowers),  with  broad  bases 


FIG.  302.— Colocas-iu  Boryi. 

-and  prolonged,  pointed  tips  (6);  underneath  these  are  the  $- 
flowers  (m),each  consisting  only  of  3-4  short  stamens,  which  eject 
vermiform  pollen-masses  through  the  terminal  pores  ;  then  follow, 
last  of  all,  ?  -flowers  (/),  each  of  which  consists  of  one  unilocular 
ovary,  with  several  ovules.  Perennial  herbs,  tuberous,  with 
cordate  leaves. — Dracunculus  ;  Biarum ;  Arisarum ;  PinrlHa  (Athcrurus)  tcr- 
nata  with  leaves  bearing  1-2  buds.  Zantedeschia  cethiopica  (Eichardia,  Nile  lily) ; 
$ ,  2-3  stamens ;  f?  with  3  staminodes,  1-5-locular  ovary  (S.  Africa.) — In  some 
genera  sterile  flowers  are  present  between  the  $  and  ?  portions  of  the  spadix 
(e.g.  in  Philodendron) ;  in  Ambrosinia  a  lateral,  wing-like  broadening  of  the  axis 
of  the  spadix  divides  the -cavity  of  the  spatlie  into  two  chambers,  the  anterior 


306  MONOCOTYLEDONES. 

containing  one  ?  ,  nnd  the  posterior  8-10  $  -flowers  in  two  series  ;  in  some  the 
stamens  in  the  single  c? -flowers  unite  and  form  a  columnar  "  synaudrium  " 
(e.g.  in  Iricffenbachia,  Colocasia,  Alocasia,  Caladium,  Taccarnm,  ^ijngoninm). 
A  remarkable  spadix  is  found  in  Spathicarpa ;  it  is  united  for  its  entire  length, 
ou  one  side,  with  the  spathe,  and  the  flowers  are  arranged  upon  it  in  r-iws,  the 

9  to  the  outside,  and  the  $  in  the  middle  (Zostera  has  a  similar  one).— Pistia 
similarly  deviates  considerably,  it  is  a  floating  water-plant,  with  hairy,  round 
rosettes  of  leaves  ;  in  it  also  the  spathe  and  spadix  are  united  ;  at  the  base  a 

$  -flower  is  borne,  which  consists  of  one  unilocular  ovary,  and  above  several 

$  -flowers,  each  composed  of  two  united  stamens. 

BIOLOGY.  The  inflorescences  are  adapted  for  insect-pollination ;  they  are 
protogynous,  since  the  viscous,  almost  sessile  stigmas  come  to  maturity  and 
wither  before  the  pollen,  which  is  generally  dehisced  by  apical  pores,  is  shed  ; 
some  pollinate  themselves  freely  by  the  pollen  from  the  higher  $  -flowers 
falling  upon  the  $  -flowers  below  them,  and  in  some  it  is  conjectured  that  the 
pollination  is  effected  by  snails.  The  coloured  spathe,  and  the  naked  end  of 
the  5-padix  (often  coloured)  of  certain  genera  function  as  the  coloured  perianth 
in  other  orders  ;  during  flowering  a  very  powerful  smell  is  often  emitted.  Arum 
maculatum  is  worthy  .of  notice  ;  small  flies  and  midges  creep  down  into  the. 
spathe,  between  the  sterile  flowers  (Fig.  301  b),  which  are  situated  where  the 
spathe  is  constricted,  and  pointing  obliquely  downwards  prevent  the  escape- 
of  the  insects ;  in  the  meantime,  the  stigmas  are  in  a  condition  to  receive 
any  pollen  they  may  have  brought  with  them  ;  after  pollination  the  stigmas 
wither,  and  exude  small  drops  of  honey  as  a  compensation  to  the  flies  for  their 
imprisonment ;  after  this  the  anthers  (m)  open  and  shed  their  pollen,  the  sterile 
flowers  wither,  and  the  insects  are  then  able  to  escape,  and  enter  and  pollinate 
other  inflorescences. — In  many,  a  rise  of  temperature  and  evolution  of  carbonic 
acid  takes  place  during  flowering ;  a  spadix  may  be  raised  as  much  as  30°C. 
above  the  temperature  of  the  surrounding  air. — Again,  under  certain  conditions, 
many  species  absorb  such  large  quantities  of  water  by  their  roots  that  water  is 
forced  out  in  drops  from  the  tip  of  the  leaf ;  this  may  often  be  observed  in. 
Zantedescliia. 

About  900  species  in  100  genera.  Home,  the  Tropics,  especially  S.  America, 
India,  and  the  Indian  Islands,  preferably  in  shady,  damp  forests  growing 
as  epiphytes  upon  trees,  and  on  the  banks  of  streams.  Outside  the  Tropics 
few  are  found.  Acorns  calamus  was  introduced  into  Europe  from  Asia  about 
300  years  ago  ;  it,  however,  never  sets  any  fruit,  as  the  pollen  is  unfertile.  In 
England  Arum  maculatum  is  a  very  common  plant;  this  and  A.  italicum 
are  the  only  native  species.  Colocasia  antiquorum  comes  from  Polynesia 
and  the  Indian  Islands,  and  also  Alocasia  macrorrhiza.  Fossils  in  Cretaceous 
and  Tertiary. 

USES.  Many  species  have  pungent,  and  even  poisonous  properties  (e.g. 
Dieft'enbachia,  Lagenandra,  Arum),  which  are  easily  removed  by  boiling  or 
roasting;  the  rhizomes  of  many  species  of  Caladium,  Colocasia  (C.  antiquoi  urn, 
esculents  etc.),  are  very  rich  in  starch,  and  in  the  Tropics  form  an  important 
source  of  food.  An  uncommon  occurrence  in  the  order  is  the  highly  aromatic 
rhizome  of  Acorus  calamus;  this  contains  calamus-oil  and  acorin  which  are  used 
in  perfumery.  Many  are  ornamental  plants,  e.g.  Zantedescliia  ceilnopica  (South, 


SPADICIFLORJ).  307 

Africa),  generally  known  as  "  Calla,"  and   Monstera   deliciosa  •    many   other 
species  are  grown  in  greenhouses. 

Order  6.  Lemnacese  (Duck- weeds).  These  are  the  most 
reduced  form  of  the  Spadiciflorse.  They  are  very  small,  free- 
swimming-  water-plants.  The  vegetative  system  resembles  a  small, 
leaf-like  body  (Fig.  303/-/),  from  which  roots  hang  downwards; 
this  branches  by  producing  a  new,  similar  leaf-like  body,  which 
springs  from  a  pocket-like  hollow  (indicated  by  a  dotted  line  in 
the  figure)  on  each  side  of  the  older  one,  at  its  base  (or  only  on 
one  side).  The  branching  is  thus  dichasial  or  helicoid  (Fig.  303  A,  where 
/,/'./",/"  indicate  shoots  of  1st,  2nd,  3rd,  4th  generations  respectively).  The 
leaf-like  bodies  are,  according  to  Hegeluiaier,  leaf-like  stems,  and  thus  Lcnma 


FIG.  303. — T.imna,:  A  vegetative  system  ;    B  portion,  of  a  plant,  with  flowers ;   one  stamen 
and  tip  of  the  carpel  project ;  the  remaining  portions  being  indicated  by  the  dotted  line. 

has  no  other  leaves  than  the  spathe  and  the  sporophylls ;  according  to  the  in- 
vestigations of  Eugler  they  are  stems  whose  upper  portion  (above  the  "  pocket  ") 
is  a  leaf,  which  is  not  sharply  separated  from  the  underlying  stem-portion. 
The  inflorescence  is  a  very  much  reduced  Araceous-spadix,  consisting  in  Lemna 
of  1  or  2  stamens  of  unequal  length  (1-stamened  <J -flowers),  1  unilocular 
carpel  ( ?  -flower),  and  1  thin  spathe  (JB).  [The  same  is  found  in  Spirodela 
polyn-ldza,  etc.,  whose  daughter- shoots  begin  in  addition  with  1  basal-leaf. 
il'nlffia  anliza.  etc.,  have  no  roots.no  spathe,  and  only  1  <? -flower  in  the 
inflorescence  (Engler).] — On  the  germination  of  the  seed  a  portion  of  the  testa 
is  thrown  off  as  a  lid,  so  that  an  exit  is  opened  for  the  radicle. — 19  species.  In 
stagnant  fresh  water,  both  Temp,  and  Tropical. — In  Europe  the  species  are 
Lemna  minor,  trisulca,  gibba  ;  Spirodela  polyrrhiza,  and  Wolffia  arrhiza,  the 
smallest  Flowering-plant. 


308 


MONOCOTYLEDONES. 


Family  4.     Enantioblastse. 

The  flowers  in  this  family  are  hypogynous  and  have  in  part  the 
general  monocotyledonous  type  with  5  trimerous  whorls  completely 
developed  in  a  regular  hermaphrodite  flower,  and  in  part  the 
flowers  so  much  reduced  that  the  type  is  very  difficult  to  trace. 
On  the  one  hand  the  family  is  well  developed  and  has  capitate 
inflorescences  (Eriocaulacece)  and  on  the  other  hand  it  is  distinctly 
reduced  (Centrolepidacece) .  This  family  has  taken  its  name  from 
the  fact  that  the  ovule  is  not,  as  in  the  Liliifloras  and  nearly  all 
other  Monocotyledons,  anatropous,  but  orthotropous,  so  that  the 
embryo  (ftXa-a-rrf)  becomes  placed  at  the  end  of  the  seed  opposite 
(evavrtbs)  to  the  hilum.  Large,  mealy  endosperm. — The  orders 
belonging  to  this  family  are  by  certain  authors  grouped  with  the 
Bromeliacece  and  Pontederiacece,  etc.,  into  one  family,  FARINOSE, 
so  named  on  account  of  the  mealy  endosperm,  the  distinguishing 
character  of  the  Liliifloros  then  being  that  the  endosperm  is  fleshy 
and  horny. 

Order  1.  Commelinaceae.  The  complete  Liliaceous  structure  without  great 
reductions  in  the  number  of  whorls,  but  with  generally  few  ovules  in  each 
loculus  of  the  ovary,  is  found  in  the  CommelinaceEe,  an  almost  exclusively 
tropical  order  with  about  317  species  ;  herbs,  some  of  which  are  introduced  into 
our  gardens  and  greenhouses.  The  stems  are  nodose  ;  the  leaves  often  clasp- 
ing ;  the  flowers  are  arranged  in  unipared  scorpioid  cymes,  often  so  that  they 
form  a  zig-zag  series  falling  in  the  median  line  of  the  bracts,  and  after  flowering 
they  bend  regularly  to  the  right  or  left,  outwards  or  inwards.  They  are  more  or 
less  zt/gomorphic,  particularly  in  the  stamens,  which  in  the  same  flower  are  of 
different  forms  or  partially  suppressed.  The  outer  series  of  the  perianth  is 
sepaloid,  the  inner  petaloid,  generally  violet  or  blue  ;  the  filaments  are  some- 
times clothed  with  hairs  formed  of  rows  of  bead-like  cells  (well  known  for 
showing  protoplasmic  movements).  Fruit  a  trilocular  capsule  with  loculicidal 
dehiscence  (generally  few-seeded)  ;  in  some  a  nut.  The  radicle  is  covered  by 
an  external,  warty,  projecting  covering  which  is  cast  off  on  germination.— 
The  abundant  rapliides  lie  in  elongated  cells  whose  transverse  walls  they 
perforate.—  Commelina,  Tradtscantia,  Tinnantia,  Cyanotis,  Dichorisandra. 

Order  2.  Mayacaceae.  This  order  is  closely  allied  to  the  Commelinaceae. 
7  species.  American  marsh-  or  water-plants. 

In  many  of  the  following  orders  of  this  family  the  flowers  are  united  into 
compound  inflorescences,  with  which  is  accompanied  a  reduction  in  the  flower. 

Order  3.  Xyridaceae  (50  species).  Marsh-plants  with  radical,  often  equit- 
ant  leaves  arranged  in  2  rows,  and  short  spikes  on  long  (twisted)  stalks.  The 
flowers,  as  in  the  Commelinacere,  have  sepals  (which  however  are  more  chaffy) 
and  petals,  but  the  outer  series  of  stamens  is  wanting.  Capsule  (generally 
many-seeded). 

Order  4.     Rapateacese.     Marsh-plants  with  radical  leaves,  usually  in  two 


LILIIFLOR.E.  809 

rows,  and  several  spikelets  on  the  summit  of  the  main  axis,  clustered  into  a 
capitulum  or  unilateral  spike.  Each  spikelet  has  numerous  imbricate  floral- 
leaves  and  one  flower.  24  species.  South  America. 

'  Order  5.  Eriocaulaceae.  The  "  Composite  among  Monocotyledons,"  a 
tropical  order.  The  flowers  are  borne  in  a  capitulum  surrounded  by  an  involucre, 
very  similar  to  that  of  the  Compositae.  The  flowers  are  very  small,  unisexual, 
$  and  £  often  mixed  indiscriminately  in  the  same  capitulum ;  they  have  the 
usual  pentacyclic  structure  ;  the  leaves  of  the  inner  perianth  are  often  connate 
and  more  membranous  than  the  outer  ;  in  some  the  outer  series  of  stamens 
are  suppressed ;  in  each  of  the  3  loculi  is  one  pendulous  ovule.  Capsule.  The 
leaves  are  generally  radical  and  grass-like.— 335  species ;  Eriocaulon,  Paepa- 
lanthus,  etc.,  E.  septangulare  on  the  west  coast  of  Scotland,  and  Ireland,  and  in 
North  America. 

Order  6.  Restiaceae.  A  small,  especially  S.  African  and  S.  Australian, 
xerophilous  order  (about  235  species),  which  is  quite  similar  in  habit  to  the 
Juncacese  and  Cyperaceae.  The  leaves  are  often  reduced  to  sheaths.  The 
flowers  are  dioecious,  the  perianth  as  in  Juncus,  but  the  outer  series  of  stamens 
suppressed.  The  ovary  and  fruit  as  in  Eriocaulaceaa  ;  the  ovary,  however,  may 
be  unilocular,  and  the  fruit  a  nut.  Restio,  etc. 

Order  7.  Centrolepidacese.  These  are  the  most  reduced  plants  in  the 
family ;  small  grass-  or  rush-like  herbs.  The  flowers  are  very  small,  naked. 
Stamens  1-2,  carpels  1-  .  32  species.  Australia. — Centrolepis  (flowers 
generally  $  with  1  stamen  and  2-oo  carpels). 

Family  5.     Liliiflorse. 

The  flower  is  constructed  on  the  general  monocotyledonous  typer 
with  5  alternating,  3-merous  whorls  (Fig.  278),  bat  exceptions  are 
found  as  in  the  Iridaceae  (Fig.  279)  by  the  suppression  of  the  inner 
whorl  of  stamens  ;  in  a  few  the  position  in  relation  to  the  bract 
differs  from  that  represented  in  Fig.  278,  and  in  some  instead  of  the 
trimerous,  di- ortetra-merous  flowers  are  found  (e.g.  Majanthemurti^ 
Paris}.  Flowers  generally  regular,  hermaphrodite,  with  simple, 
petaloid,  coloured  perianth  (except,  for  example,  Bromeliaceas) ; 
ovary  trilocular,  generally  with  2  ovules  or  2  rows  of  ovules  in  the 
inner  angle  of  each  loculus  (Fig.  304  0,  D).  Endosperm  always 
present. — A  very  natural  family,  of  which  some  divisions  in  part 
overlap  each  other.  The  habit  varies  ;  the  leaves  are  however  long, 
entire,  with  parallel  venation,  except  in  Dioscoreaceae  (Fig.  313). 

In  the  first  orders  of  this  family  the  flowers  are  hypogynous,  and  in  the 
first  of  all  the  styles  are  free,  and  the  capsule  dehisces  septicidally ;  in  the 
following  the  flowers  are  epigynous  and  in  some  reduced  in  number  or  uni- 
sexual ;  capsule  with  loculicidal  dehiscence,  or  a  berry. 

HYPOGYNOUS  flowers  :  Colchicaceaa,  Liliaceaa,  Convallariaceas,  Bromeliaceas  (in 
part). 


310 


MONOCOTYLEDONS. 


EPIGYNOUS  flowers:  Amaryllidacese,  Iridaceee,  BromeliacesB  (in  part),  Dios- 
coreaceae. 

Order  1.  Colchicacese.  The  flower  (Fig.  304.4)  is  g,  regular, 
hypogynous,  trimerous  in  all  five  whorls  (6  stamens) ;  anthers 
usually  extrorse.  Gynceceum  with  3  free  styles  (yl,  D) ;  fruit  a 
capsule  with  septicidal  dehiscence  (E)  ;  embryo  very  small  (F).  The 
underground  stem  is  generally  a  corm  or  rhizome,  seldom  a  bulb. 

A.  VERATREJ;. — Veratrum ;    perennial    herbs,    stem    tall    with 
long   internodes    and    broad,   folded   leaves  ;  the    flowers    andro- 
moncecious,  with  free,  widely  opening  perianth-leaves  (Fig.  304  A}, 
and    globular     anthers;      inflorescence    a    panicle. —  Zygadenus, 
Melanthium,  Schcenocaulon,  Uvularia,  Tricyrtis. 

B.  TOFIELDIEJ;. — Narthecium   and    Tojieldia   have   leaves  alter- 
nate (arranged  in  two  rows),   sword-like  and  borne  in  rosettes  ; 
racemes  or  spikes.    Narthecium  forms  an  exception  to  the  order  by 
having   a   simple    style    and  fruit   with    loculicidal    dehiscence ; 


FIG.  304.— Veratrum  :  A  flower;  B  stamen  ;  C  transverse  sect  ion  of  ovary  ;  D  gynreceum, 
with  one  carpel  bisected  longitudinally,  and  the  third  removed  ;  E  fruit  after  dehiscence  ; 
f  longitudinal  section  of  a  seed. 

Tojieldia  by  the  introrse  anthers.  In  this  they  are  related  to  the 
Liliacese.  Narthecium  has  poisonous  properties,  like  many  other 
Colchicaceae. 

C.  CoLCHiCE^:. — Colchicum  (Autumn  Crocus)  ;  perennial  herbs, 
with  a  long,  funnel-shaped,  gamophyllous  perianth,  and  introrse 
anthers.  The  flowers  of  C.  autumnale  spring  up  immediately 
from  the  underground  stem,  which  is  in  reality  a  corm  formed  of 
one  internode.  Colchicum  autumnale  flowers  in  autumn  without  leaves  ;  in 
spring  the  radical  foliage-leaves  appear  simultaneously  with  the  fruit.  The 
flower  is  protogynous,  and  is  pollinated  by  insects  (humble-bees,  etc.)  which 
seek  the  honey  secreted  by  the  free  part  of  the  stamen  a  little  way  down  the 
tube.  The  length  of  the  tube  protects  the  fruit,  and  not,  as  in  other  cases,  the 
nectary. — Tiulbocodium  and  Merendera  have  unguiculate  perianth  leaves,  free, 
but  closing  together  like  a  tube. 

175   species ;    chiefly  in  North  America  and  South  Africa.     Tofieldia  is  an 


LILIIFLORJ3. 


311 


Arctic  plant.  The  order  is  rich  in  pungent,  poisonous  alkaloids  (veratrin, 
colchicin,  etc.).  OFFICINAL;  the  seeds  of  Colchicum  autumnale  (Europe)  and 
Scluznocaulon  officinale  (Mexico),  and  the  rhizome  of  Veratrum  album  (mountains 
of  Central  Europe). 

Order  2.  Liliaceae  (Lilies).  Flowers  as  in  the  Colchicaceee 
but  with  introrse  anthers ;  ovary  free,  3-locular,  with  single  style ; 
capsule  3-locular  with  loculicidal  dehisceiice. — The  majority  are 

A 


FIG-.  305. — Colchicum  autumnale.  A  Conn  seen  from  the  front :  7;  corm  •  s's"  scale-leaves 
embracing  the  flower- stalk ;  trTi  base  of  flower-stalk  with  roots  (tc).  B  Longitudinal  section 
of  corm  and  flower-stalk :  hh  brown  membrane  surrounding  the  underground  portion  of 
the  plant ;  st  flower-  and  leaf-stalk  of  previous  year,  the  swollen  basal  portion  forming  the 
reservoir  of  reserve  material.  The  new  plant  is  a  lateral  shoot  from  the  base  of  the  corm 
(Jc)  and  has  the  following  parts  :  the  base  bearing  the  roots  (to),  the  central  part  (fc')  which 
becomes  the  corm  in  the  next  yenr,  the  axis  bearing  the  scale-leaves  (s',  s"),  the  foliage- 
leaves  (I,  I'"),  and  the  flowers  (b,  b')  which  are  borne  in  the  axils  of  the  uppermost  foliage- 
leaves. 


312  MONOCOTYLEDONES. 

herbs  with  bulbs ;  the  inflorescence  is  terminal.  In  many  species  repro- 
duction takes  place  by  means  of  bulbils  (small  bulbs)  formed  in  the  axils  of 
the  foliage-leaves  (e.g.  Lilium  bulbiferum,  Jancifolium,  etc.,  Gagea  lancij'olia, 
etc.),  or  in  the  bracts  of  the  inflorescence  (many  species  of  Alliuni) ;  in  many 
species  several  buds  are  developed  as  bulbs  in  the  axils  of  the  bulb-scales  them- 
selves (accessory  buds  arising  close  together),  and  in  some  the  formation  of 
buds  is  common  on  the  leaves. 

A.  TULIPE.E,  TULIP  GROUP.    Bulbs.     The  aerial,  elongated  stem 
bears  the  foliage-leaves.     Flowers  few  but  generally   large,  with 
free   perianth-leaves.      Tulipa ;    style    absent,  no   honey ;    flowers 
generally  solitary,  erect. — Fritillaria   perianth  campanulate  with 
a  round    or  oblong  nectary  at  the  base  of  each  perianth-leaf. — 
Lilium;    perianth    widely   open,    generally    turned    back   with    a 
covered  nectary-groove  in  the  centre  of  each  segment.     Anthers 
versatile. — Lloydia ;  Erythronium. 

B.  HYACINTHEJE,  HYACINTH  GROUP.    Bulbs.    Leaves  radical ;  aerial 
stem  leafless  with  raceme  or  spike.    In  some  the  perianth-segments 
are  free,  in  others  united.     Honey  is  produced  often  in  glands  or 
in  the  septa  of  the  ovary    (septal  glands). — Ornithogalum  has  a 
leafy  stem ;  Scilla  ;  Eucomis  has  a  tuft  of  floral-leaves  above  the 
raceme;  A gr aphis;  Hyacinthus;  PuscTikinia  ;  CMonodoxa;  Muscari; 
VeWieimia ;   Urginea. 

C.  ALLIED,  ONION    GROUP.     Generally  bulbs.     Leaves  radical. 
Stem  leafless  with  a  compound  umbellate  or  capitate  inflorescence 
of   uni pared    helicoid    cymes,  which   before    flowering    are   sur- 
rounded by  two  broad  involucral    leaves. — Allium.    Filaments  often 
petaloid  and  bidentate  ;  in  many  species  bulbils  are  found  in  the  inflorescence. 
— Some  species  have  flat  leaves  :  A.  sativum,  Garlic ;   A.  porrum,  Leek ;  A. 
ursinum;  others  have   round,  hollow  leaves:  A.  cepa,   Onion;  A.  jistulosum, 
Winter  Onion  ;  A.  ascalonicum,  Eschalot ;  A.  schcenoprasum,  Chive. — Gagea  ; 
honey  is  secreted  at  the  base  of  the  perianth,  no  special  nectary; 
inflorescence  few-flowered — AgapantJius ;   Triteleia. 

D.  ANTHERICE^E.      Khizome;    raceme;   the   leaves  not  fleshy  and  thick. — 
AntJtericum ;  Asphodclus  ;  Bulbine  ;   Cldorophytum  ;  Bowiea  has  an  almost  leaf- 
less stem  frith  curved,  climbing  branches. 

E.  ALOINEJE,  ALOES.    Stem  generally  aerial  and  tree-like,  bearing  on  its  sum- 
mit thick,  fleshy  leaves,  often  with  a  thorny  edge  (Fig.  306).     Eaceme  branched 
or  unbranched. — Aloe  ;  Gaateria  ;   Yucca  (has  secondary  thickening,  p.  274). 

F.  HEMEROCALLIDE^E.    Phormium,  (Ph.  tenax.    New  Zealand  Flax) ;  Funckia 
(Hosta') ;  Hemcrocallis. 

At  this  point  the  following  are  best  placed :  Aphyllanthes  (A.  monspeliensis) ; 
Xanthorrhcca  (Black-boy) ;  Xerotes  ;  Lomandra ;  Kingia  ;  the  very  membranous, 
dry  perianth  of  the  last  resembles  that  of  the  Juncacea*,  and  also  there  are  only 
1-few  ovules  in  the  loculi. 


313 


POLLINATION  by  insects.  Honey  in  some  is  produced  on  the  perianth  (see 
Tulipeae),  in  others  by  glands  on  the  carpels  (in  the  septa  and  parietal  placentas, 
septal  glands) :  Hyacinthus,  Allium,  Anthericum,  Asphodelus,  Yucca,  Funckia, 
Hemerocallis,  etc.  Some  Allium- species  are  protandrous.  Fritillaria  is  visited 
by  bees,  Lilium  martagon  by  moths,  L.  bulbiferum  by  butterflies,  Phormium 
(New  Zealand)  by  honey-birds. 


FIG.  308.— Aloe. 

About  1,580  species ;  rare  in  cold  climates ;  their  home  is  in  sunny  plains 
with  firm,  hard  soil,  and  warm  or  mild  climate,  particularly  in  the  Old  World 
(S.  Africa;  As.  Steppes;  Mediterranean)  ;  at  the  commencement  of  spring  the 
flowers  appear  in  great  profusion,  and  after  the  course  of  a  few  weeks  disappear  ; 
during  the  hot  season  their  life  lies  dormant  in  the  bulb,  hidden  underground. 


314  MONOCOTYLEDOXES. 

The  woody  species  are  tropical. — The  majority  of  the  introduced  Liliaceae 
(b'ritlUaria  imperialis,  Crown-imperial;  Lilium  candidum;  Tulipa  gesneriana; 
Hyacinth  ;  J/uscari-species ;  Scilla-species ;  Ornithogalum  nutans  ;  Hemerocallis 
Julva  an&flava  ;  Asphodelus  luteus  and  albus)  come  from  the  Mediterranean  and 
W.  Asia ;  Funckia  from  China  and  Japan  ;  several  Lilies  from  Japan  and  the 
Himalayas  ;  Agapanthun  from  the  Cape  ;  Allium  sativum  is  a  native  of  the 
Kerghis -Steppes ;  A.  cepa  from  Persia  (?) ;  A.  ascalonicum  is  not  known 
•wild  (according  to  others  a  native  of  Asia  Minor),  perhaps  a  form  of  A.  cepa  ; 
A.  schcenoprasum  from  the  N.  temp,  region. 

Many  bulbs  have  pungent  properties  ;  many  Onions  are  used  as  culinary  plants 
The  bast  fibres  of  Phormium  tenax  (New  Zealand  Flax)  are  used  technically. 
Dyes  are  obtained  from  the  Aloe  ;  gum  for  varnish  from  the  stem  of  Xanthorrlicea 
li'ixtile  and  australe.  OFFICINAL  ;  "Aloes,"  the  dried  sap  of  S.  African  species 
of  Aloe  (A.  Africana,  A.ferox,  etc.) ;  the  bulb  known  as  "Squills"  from  Urginea 
(Scilla)  maritima  (Mediterranean). 

Order  3.  Convallariaceae.  This  order  differs  from  the 
Liliaceae  in  having  the  fruit  a  berry  (Fig.  308)  and  in  never  being 
bulbous  ;  the  seeds  are  less  numerous. 

A.  CONVALLARIEJE,  LlLY   OF  THE   VALLEY   GROUP.       Rhizome  (Fig. 

507)  and  normal  foliage-leaves. — Polygonatum  :  rhizome  creeping ; 

aerial  shoot  leafy,  bearing  the 
flowers  in  racemes  in  the  axils  of 
the  foliage-leaves ;  perianth  tubu- 
lar. P.  multfylornm  (Solomon's 
seal),  P.  officinale,  etc. — Majanthe- 
mum :  flower  2-merous  ;  perianth 
almost  polyphyllous,  spreading. 
Smilacina,  Streptopus  (S.  amplexi- 
FIG.  307.— Rhizome-  of  Polygonatum  folius ;  the  flowers  or  inflorescence 

muitiflorum :  a  bud ;  b  shoot ;  c  d  scars      unite    with  the   entire  succeeding 

left  by  shoots  of  previous  years. 

internode). — Convallaria  (1  species 

C.  majalis,  Lily  of  the  valley)  ;  flowers  in  terminal  racemes  ;  2  basal 
foliage-leaves  ;  perianth  globose,  bell-shaped.  lleineckea  carnea 
(Japan,  China)  in  gardens. — Paris  (P.  quadrifolia,  Herb- Paris)  ; 
flowers  solitary,  terminal,  4-merous,  polyphyllous ;  styles  4,  free 
(approaching  the  Colchicacese ;  it  is  also  poisonous)  ;  a  whorl  of 
4  (-more)  3-nerved,  reticulate  leaves  on  each  shoot. — Ornamental 
plants :  species  of  Trillium,  Aspidistra  elatior  (Japan). 

B.  ASPARAGEJ;,  ASPARAGUS  GROUP.     Scale-like  leaves  and  green 
assimilating  branches. — Asparagus  :  horizontal  rhizome.     The  aerial 
shoots  are  very  richly  branched ;  the  numerous  needle-like  bodies 
upon  the  plant  are  leafless  shoots,  which  are  crowded  together  in 
double  scorpioid  cymes  in  the  axils  of  the  scale-leaves  ;  the  two  first 


315 


lateral  axes,  placed  outside  to  the  left  and  right,  generally  bear 


FIG.  308. — Smilax  pseudosiiphilUica :  A  shoot  of  male  plant  ;  C  3  -flower  ;  D  berry,  almost 
ripe;  E  the  same  in  longitudinal  section.  B  Smilax  syphilitica :  portion  of  branch  with  base 
of  leaf  and  tendrils. 


316  MONOCOTYLEDOXES. 

flowers.  Polygamous. — P..USCUS  (Butcher's  broom)  is  a  S.  European  shrub 
with  leaf-like,  ovoid  or  elliptical  shoots  (phylloclades)  which  are  borne  in  the 
axils  of  scale-like  leaves,  and  bear  flowers  on  the  central  line.  Dioecious. 
Stamens  3,  united,  anthers  extrorse.  Semele  androgyna  bears  its  flowers  on 
the  edge  of  the  flat  shoot. 

C.  SMILACE.ZE.      Smilax    (Sarsaparilla)     (Fig.    308) ;    climbing 
shrubs  with  the  leaf-sheath    produced  into  tendrils.     The  leaves 
have  3-5  strong  nerves  proceeding  from  the  base,  and  are  reticulate. 
Orthotropous  or  semi-anatropous  ovules.    Dioecious  (Fig.  308  C,E). 

D.  DKACJENE.E.    Fruit  in  some  a  berry,  in  others  a  capsule.     The  stem  of 
DRACJENA,  when  old,  has  the  appearance  of  being  dichotomously  branched ;  it 
has  the  power  of  increase  in  thickness,  and  may  become  enormously  thick.    The 
Dragon-tree  of  Teneriffe,  measured  by  Humboldt,  attained  a  circumference  of 
14  m.  and  a  height  of  22  m. ;  the  leaves  are  large,  linear  or  linear-lanceolate. — 
Cordyline  (East  Asia),  various  species  in  gardens  and  greenhouses  (Yucca  is 
closely  allied).     Astelia. 

POLLINATION.  Paris  quadrifolia  and  Convallaria  majalis  have  no  honey, 
and  are  chiefly  visited  by  pollen-collecting  bees  (in  the  absence  of  insect  visits 
self-pollination  takes  place) ;  Polygonatum  multiflomm  has  honey  secreted  by 
septal  glands  and  protected  by  the  base  of  the  tubular  perianth  ;  it  is  pollinated 
by  humble-bees,  etc.  Asparagus  officinalis  has  small,  polygamous,  greenish, 
honey-bearing  flowers ;  the  $  -flower  is  almost  twice  as  large  as  the  9  ;  both 
have  rudiments  of  the  opposite  sex. 

About  555  species  ;  especially  from  N.  America,  Europe,  and  Central  Asia. 

OFFICINAL  :  "  Dragons' -blood,"  a  red  resinous  juice  from  the  stem  of 
Draccena,  and  the  roots  of  some  Central  American  species  of  Smilax.  The 
tuberous  stems  of  the  Eastern  Asiatic  Smilax  glabra  are  officinal.  The  flowers 
of  Conrallaria  majalis  have  been  lately  used  as  a  substitute  for  Digitalis. 
Pungent,  poisonous  properties  are  possessed  by  Paris.  None  of  the  species  are 
used  as  food,  except  the  young  annual  shoots  of  Asparagus  officinalis,  a  shore- 
plant  which  is  used  as  a  vegetable. 

Order  4.  Pontederiaceae.  Flowers  generally  zygomorphic,  hypogynous,  $  , 
with  handsome,  white  or  violet,  petaloid  perianth  which  forms  a  tube  at  its  base. 
The  stamens  are  inserted  at  different  heights  in  the  perianth-tube,  and  are  re- 
duced to  three  (in  Heteranthera  seldom  to  one).  In  some  the  ovary  is  trilocular 
with  oo  ovules  (Eichhornia),  in  others  reduced  to  one  loculus  with  one  ovule 
(Pontederia).  Fruit  a  capsule  or  nut.  Embryo  as  long  as  the  abundant,  mealy 
endosperm. — Tropical  water-plants  (22  species)  with  peculiar  sympodial  branch- 
ing, nearly  the  same  as  in  Zostera.  Spikes  without  floral-leaves.  Many  inter- 
cellular spaces  in  the  stem  and  leaf. — In  greenhouses:  Eichhornia  azurea,  E. 
crassipes  (both  from  tropical  and  sub-tropical  S.  America)  ;  the  latter  has 
swollen  petioles  which  serve  as  floats  and  enable  it  to  float  freely  on  the  water, 
sending  down  its  roots  into  the  mud.  Heteranthera  reniformis,  H.  zosterifolia. 
Pontederia  cordata. 

Order  5.  Amaryllidacese  (Narcissi).  The  flower  is  epi- 
gynous,  otherwise  exactly  the  same  as  in  the  Liliacea?  (6  stamens). 


LIL1IFLOBJE. 


317 


The  majority,  like  these,  are  also  perennial  herbs  with  bulbs  and 
scapes.  The  fruit  and  the  other  characters  as  in  the  Liliaceae. 
The  external  appearance  is,  however,  very  different. 

A.  AMAEYLLEJJ  have  bulbs  and  the  leaves  generally  arranged 
in  two  rows  ;  the  flowers  are  borne  singly  or  in  umbel-like  in- 
florescences on  lateral  scapes,  while  the  main  axis  of  the  bulb  is 
unlimited.  Beneath  the  inflorescence  is  an  involucre  (Fig.  309). — 
Galanthus,  Snowdrop,  has  a  polyphyllous  perianth  without  corona ; 
the  three  inner  'perianth-leaves  are  emarginate  and  shorter  than 


FIG.  309.— Pancratium  caribceum. 

the  outer ;  the  anthers  dehisce  apically.  Leucojum  differs  in  having 
the  perianth-leaves  equal  in  length.—  Amaryllis  has  a  funnel-shaped  perianth, 
entirely  or  nearly  polyphyllous,  but  somewhat  zygomorphic.  Crinum ; 
Heemanthus  ;  Clioia. — Narcissus  has  a  tubular  corona,  a  ligular  .struc- 
ture arising  from  the  perianth-tube  exterior  to  the  outer  stamens. 
In  Pancratium  (Fig.  309)  the  corona  is  united  with  the  filaments  which  appear 
to  spring  from  its  edge.  Eucharis  amazonica. 

B.  H-JPOXIDEJE.  The  leaves,  which  are  grass-like,  dry,  folded,  and  in  some 
hairy,  spring  from  a  rhizome,  generally  with  a  divergence  of  I/s.  Flowers  small, 
perianth  polyphyllous,  persistent,  on  which  account  perhaps  the  Hypoxideas  may 


318  .  MONOCOTTLEDONES. 

be  considered  as  the  least  altered  type.  The  chief  characteristic  is  that  the 
embryo  is  separated  from  the  hilum.  Hypoxis-  Curculigo  (C.  recurvata,  a. 
favourite  ornamental  plant;  S.E.  Asia). 

C.  ALSTRffiMERiEa;.  (Alstrcemeria,  Bomarea) ;  stems  long,  leafy,  often  climb- 
ing. 

D.  VELLOSIE^;  (Vellosia,  Barbacenia) ;  stem  woody,  usually  riichotomously 
branched,  with  terminal,  single  flowers ;  it  bears  numerous  aerial  roots  which 
pierce  the  leaves  and  surround  the  stem.     Stamens  often  (by  splitting)  6-18. 
High  table-lands  of  S.  America  and  S.  Africa. 

E.  AGAVES.     Very  similar  to  the  Bromeliacese  both  in  their   distribution 
(nearly  all  American)  and  in  external  appearance.  They  appear  as  gigantic  bul- 
bous plants  with  perennial,  aerial,  generally  short  stem,  and  perennial,  large, 
lanceolate  or  linear,  stiff,  thick,  and  often  thorny  leaves,  which  form  a  large 
rosette  ;  after  the  course  of  several  (8-20)  years  the  terminal  inflorescence  is 
developed,  which  is  10-12  m.  high,  paniculate,  and  freely  branched.     Before 
the  inflorescence  expands,  a  large  quantity  of  sugar-containing  sap  is  collected 
from  A.  americana  by  removing  the  terminal  bud ;  this  on  distillation  yields 
"  pulque,"  the  national  drink  of  Mexico.    After  flowering  the  entire  shoot  dies, 
but  the  subterranean  lateral  shoots  survive  and  reproduce  the  plant. — Agave 
americana,  etc. ;  Fourcroya  ;  Polianthes  tuber osa  (Tuberose  ;  Central  America). 

DISTRIBUTION.  The  650  species  are  chiefly  natives  of  S.  Africa  and  S. 
America.  Clivia,  Hcemanthus,  Amaryllis  are  from  the  Cape ;  Narcissus  from  S. 
Europe,  whence  many  species  have  been  introduced ;  Galanthus  and  Leucojum 
are  especially  from  S.  and  Central  Europe,  and  from  the  Caucasus. 

USES,  faw,  except  as  ornamental  plants  :  Galanthus  nivalis ;  Leucojum  ;  Nar- 
cissus pseudonarcissus,  N.  poeticus,  N.  jonquilla,  N.  tazetta,  etc.  ;  Amaryllis, 
Alstroemeria,  Eucharis,  Crinum,Vallotat  etc.  The  vascular  bundles  of  the  various 
species  of  Agave  (Agave  rigida,  var.  sisalana,  sisal  hemp,)  are  used  for  cordage,  etc. 

Order  6.  Bromeliaceae.  The  flowers  are  hypogynousr 
epigynous  or  semi-epigynous  ;  the  perianth  is  divided  into  calyx 
and  corolla;  stamens  6.  The  fruit  is  a  capsule  or  berry  with 
many  seeds.  Endosperm  mealy,  embryo  small,  at  the  edge  of  the 
endosperm,  but  not  enclosed  by  it. 

Perennial  herbs  with  a  very  characteristic  appearance  (Fig.  310)  ; 
the  stem  is  most  often  short,  thick,  arid  crowned  by  a  rosette  of 
many  leaves,  which  are  long,  often  very  narrow,  leathery,  stiff, 
and  with  a  spiny  edge;  they  are  usually  channeled,  completely 
closing  round  each  other,  with  their  edges  forming  a  tightly  closed 
hollow,  in  which  generally  water  is  collected  (this  among  other 
things  insulates  the  inflorescence  and  thus  prevents  the  access  of 
creeping  insects,  such  as  ants).  The  presence  of  numerous  stellate, 
water-containing  hairs  often  gives  the  leaves  a  grey  appearance, 
and  the  layers  of  cells  beneath  the  upper  epidermis  of  the 
lamina  form  an  "  aqueous  tissue,"  which  serves  as  a  protection 
against  the  rays  of  the  sun  and  regulates  the  evaporation.  The 


LILI1FLOR-E. 


FIG.  310. — Aechmca  miniata. 


storaata  are  often  situated  in 
furrows  on  the  under-side  of 
the  leaf,  and  hence  cause  a 
striped  appearance.  They  are 
all  American  (525  species),  es- 
pecially from  S.  America,  where 
they  live  partly  as  epiphytes  on 
trees,  partly  in  the  clefts  of  rocks, 
often  on  the  steepest  slopes,  to 
which  they  firmly  attach  them- 
selves by  aerial  roots ;  some  are 
terrestrial.  The  stem  is  seldom 
tree-like  or  many  metres  in 
height  (Puya,  in  Chili ;  Hechtia, 
in  Mexico).  The  inflorescence  is 
a  terminal  spike,  raceme,  or 
panicle,  often  with  large  and 
brightly-coloured  floral-leaves. 

The  flowers  are  without  scent, 
w.  B. 


FIG.  3L1.— Mnltiple-fruit  of  Ananassa  sativa, 
Y 


320  MONOCOTYLEDON!:  S. 

The  seeds,  in  the  species  whose  fruit  is  a  capsule,  are  often  pro- 
vided with  wings  (hairs,  expansions,  etc). — Ananassa  sativa,  Pine- 
apple (W.  Indies,  Central  America)  is  cultivated  for  the  sake  of 
its  juicy,  aromatic  fruits,  which  coalesce  with  their  fleshy  bracts 
and  form  a  large  spike-like  fruit-cluster  (multiple-fruits,1  Fig.  311) 
bearing  on  its  apex  a  leafy  shoot,  which  may  be  used  as  a  cutting. 
Seeds  very  rarely  developed. — Tillandsia  (T.  usneoides  is  a  fila- 
mentous, richly  branched,  rootless  epiphyte  hanging  in  masses 
from  trees;  Trop.  Am.),  Aechmea,  Billbergia,  Pitcairnia,  etc. 

USES.  The  leaves  of  the  Pine-apple,  in  its  native  country,  are  used  for  the 
manufacture  of  cloth. 

Order  7.  Haemodoraceae.  120  species  ;  in  all  parts  of  the  world  except 
Europe  ;  perennial,  often  tomentose  and  resembling  the  Bromeliaceae,  Iridaceae 
and  Amaryllidacese.  Hcemodorum  (Australia). — To  this  order  belong  Ophio- 
pogon,  Peliosanthes,  Sanseviera,  and  others. 

Order  8.  The  Iridaceae  have  epigynous,  hermaphrodite  flowers 
with  petaloid  perianth  as  in  the  Amaryllidaceoe,  but  the  interior  whorl 
of  stamens  is  entirely  suppressed,  and  the  3  developed  outer  stamens 
have  extrorse  anthers  (Fig.  279)  ;  there  is  1  style  with  3  large,  gene- 
rally more  or  less  leaf-like  branches  bearing  the  stigmas.  Ovary  and 
capsule  as  in  the  Amaryllidaceae  and  Liliacea3. — Perennial  herbs  ; 
bulbs  are  rarely  found,  but  horizontal  rhizomes,  corms,  etc.,  take 
their  place.  The  leaves  are  (except  Crocus}  as  in  the  Iris,  two- 
rowed,  equitant  and  sword-like.  Flowers  or  inflorescences  terminal. 

The  Iris  (Flag)  has  a  horizontal  rhizome.  The  flowers  are  borne 
in  the  leaf-axils  in  fan-like  inflorescences  (rhipidium).  The 
branches  of  the  style  are  large  and  petaloid ;  on  their  under  surface 
may  be  seen  a  small  projecting  shelf  (Fig.  312  a)  having  on  its 
upper  surface  the  stigmatic  hairs.  Beneath  the  branches  of  the 
style  are  3  well  protected  stamens,  and  immediately  outside  these 
the  external  perianth-leaves.  The  honey  is  secreted  in  the  perianth-tube, 
and  the  insects,  endeavouring  to  obtain  it  through  the  narrow  passages  at  the 
base  of  the  stamens,  settle  upon  the  outer  perianth-leaves,  which  are  bent  back- 
wards and  often  very  hairy  along  their  central  line.  The  insects  then  rub 
their  backs  on  the  anthers  just  above  them,  beneath  the  branches  of  the  style ; 
they  readily  deposit  the  pollen  on  the  stigma  of  another  flower  as  they  enter  it, 
but  cannot  do  so  in  withdrawing,  since  the  stigma  is  pushed  back,  and  self- 
fertilisation  is  thus  avoided.  The  stylar  branches  lie  close  to  the  outer 
perianth-leaves,  which  are  just  beneath  them,  or  separated  by  a  distance  of  only 
6-10  mm. ;  the  first  form  of  flower  is  adapted  for  Rhingia  rostrata,  the 
latter  for  bees.— Crocus  has  vertical,  tuberous,  underground  stems 

1  The  aggregation  of  the.  fruits  of  several  distinct  flowers  into  one  mass. 


L1LIIFI/ORJ5. 


321 


surrounded  by  the  leaf-sheaths  (corms),  and  terminal  flowers;  the 
linear  leaves  are  not  equitant,  but  have  two  longitudinal  furrows 
on  the  under  side.  The  perianth  is  gamophyllous  and  funnel- 
shaped.  The  stylar  branches  (stigmas)  are  fleshy,  rolled  together 
in  the  shape  of  a  horn,  and  split  along  the  edge. — Gladiolus  has 
corms  like  the  Crocus ;  spikes  with  slightly  zygomorphic,  almost 
bilabiate  flowers,  most  frequently  turning  to  one  side.  Position 
of  the  leaves  as  in  the  Iris.— Diplarrhena  has  2  fertile  and  1  barren 
stamen;  Hermodactylus  has  a  unilocular  ovary  with  3  parietal  placenta?. 
Cypella  and  Tigridia  have  bulbs. 


FIG.  312. — Iris  pseudacorus.  One  external  and  two  internal  perianth-leaves,  and  one 
of  the  stylar-branches  have  been  removed,  y  The  outer,  i  the  innac  perianth-leaves  ; 
g  stylar-branch ;  a  stigma ;  s  anther.  The  ovary  is  seen  in  longitudinal  section. 

770  species ;  chiefly  in  the  countries  round  the  Mediterranean,  and  in  Africa, 
especially  the  Cape  (Gladiolus,  Ferraria,  Morcea,  Galaxia,  Sparaxis,  Antholyza, 
Tritonia,  Ixia,  etc.),  Australia  and  Tropical  America  (Sisyrinchium,  Tigridia, 
Cipura,  Cypella,  etc).  A  great  number  are  ornamental  plants  :  the  cultivated 
Croczts-species  are  from  the  South  of  Europe  and  Asia  ;  Gladiolus  communis 
from  S.  Europe ;  the  other  species  principally  from  S.  Africa.  The  native 
species  of  Iris  are  I.  pseudacorus  (yellow)  and  I.  fcetidissima. 

OFFICINAL  :  the  stigmas  of  Crocus  sativus  (Oriental,  cultivated  in  France, 
Spain,  Italy,  and  Austria),  used  as  a  colouring  matter,  saffron ;  the  rhizomes 
of  the  S.  European  Iris  Jlorentina,  pallida,  and  germanica  ("  Orris-root "). 


322 


MONOCOTYLEDONES. 


Order  9.  Dioscoreaceae.  Perennial  herbs  with  fleshy,  often 
very  large  tuberous  rhizomes  (or  roots)  ;  twining  stems  ;  leaves 
stalked,  often  arrow-  or  heart-shaped,  lobed,  palminerved  and  finely 
reticulate  as  in  .  the  Dicotyledons  (Fig.  313).  The  flower  is' 
diclinous  (most  frequently  dioecious),  regular,  epigynous,  small,  and 


of  a  greenish  colour,  but  otherwise  typical  (Pr3  +  3,  and  A3 +  3, 
or  G-3) ;  in  most  instances  2  ovules  are  placed  one  above  the 
other  in  each  loculus.  The  inflorescence  is  a  spike  or  raceme,  some- 
times richly  branched  and  paniculate. — The  order  approaches  most 
nearly  to  the  Amaryllidacese. 


SCITAMINEJ:.  323 

Tamus  (Bryony)  has  a  berry,  Dioscorea  (Yam)  a  thin-walled, 
3-edged  or  3- winged  capsule  (Fig.  313).  Both  have  subterranean 
or  aerial  tubers  ;  the  Yam  very  often  also  developes  tubers  in  the 
axils  of  the  foliage-leaves  ;  tuberous  roots  are  said  to  occur  in 
D.  batatas.  The  tubers  of  many  species  of  Yams  (D.  batatas  from 
China  and  Japan,  D.  alata,  South  Sea  Islands  and  India,  D. 
bulbifera)  are  a  very  important  source  of  food  in  the  Tropics, 
especially  the  first-named. — Testudinaria  ;  Bajania. — The  tuberous 
stem  of  Tamus  communis  and  Testudinaria  elephantipes,  and  some  species  of 
Dioscorea  is  formed  from  one  single  internode  (epicotyl),  and  the  aerial  shoots 
are  developed  from  adventitious  buds ;  in  T.  elephantipes  the  stem  is  aerial,  and 
covered  with  thick  scales  of  cork,  regularly  arranged,  and  separated  by  grooves. 
Tropical  order  (167  species) ;  2  species  (Tamus  communis  and  Borderea 
pyrenaica)  in  Europe. 

Family  6.     Scitaminese. 

The  flowers  belong  to  the  ordinary  monocotyledonous  type. 
They  are  hermaphrodite,  epigynous,  and  have  either  a  petaloid 
perianth,  or  calyx  and  corolla ;  they  are,  however,  zygomorphic 
or  unsymmetrical,  and  of  the  stamens  most  frequently  only  one 
is  completely  developed,  the  others  being  generally  represented  by 
petaloid  staminodes.  The  ovary  has  3  loculi,  more  rarely  it  is 
unilocular  with  the  suppression  of  2  loculi.  Endosperm  is 
absent  (except  Zingiberacece)  ;  but,  on  the  other  hand,  there  is  a 
large  perisperm.  To  this  family  belong  large,  glabrous,  especially 
perennial  herbs  with  rhizomes ;  leaves  large,  distinctly  divided 
into  sheath,  stalk,  and  blade,  the  latter  being  more  or  less 
elliptical  or  lanceolate,  entire,  with  pinnate  venation,  and  always 
with  a  very  well-pronounced  midrib,  gradually  tapering  towards 
the  apex,  and  giving  off  numerous  branches,  which  run  outwards, 
towards  the  margin,  at  a  larger  or  smaller  angle;  these  lateral 
veins  are  closely  packed,  and  parallel,  but  with  only  weak,  con- 
necting branches  between  them ;  the  leaves,  therefore,  are  easily 
torn  pinnately  (Figs.  314,  317).  The  leaf -sheaths  close  tightly 
round  each  other  and  form  a  false  stem. 

This  very  natural  family  comprises  orders  closely  connected 
with  each  other,  but  is  not  itself  nearly  allied  to  any  other  family. 
First  in  the  series  stands  :  — 

Order  1.  Musaceae.  The  petaloid  perianth  is  strongly 
zygomorphic,  the  anterior  leaf  being  very  large  (a  kind  of 
"  labellum  "),  the  posterior  one  small ;  only  the  posterior  stamen 
is  wanting,  or  is  rudimentary,  the  other -five  are  developed,  and 


324 


MONOCOTYLEDONES. 


have  quadrilocular  anthers ;  ovary,  3-locular.     Seed  with  straight 
embryo  in  mealy  perisperm. 

The  best-known  genus  is  Musa,  the  Banana  (Fig.  314).  From 
the  short  rhizome  arise  enormously  large,  spirally-placed  leaves, 
whose  sheaths  envelope  one  another,  and  form  an  apparentlv  aerial 


FIG.  314.— Two  Mttsa-species. 

stem,  several  metres  in  height.  The  inflorescence  is  a  terminal 
spike  with  floral-leaves  placed  spirally,  and  sometimes  magnifi- 
cently coloured ;  in  the  axils  of  each  of  these  several  flowers  are 
situated  in  two  transverse  rows  (accessory  buds)  ;  the  lowest 
flowers  in  the  inflorescence  are  $  ,  the  central  ones  ty  ,  the  upper 


SCITAMINEJ1. 


325 


ones  (£,  so  that  fruits  are  only  found  in  the  lower  region  of  the 
inflorescence,  the  remaining  portion  persisting  as  a  naked  axis 
after  the  floral-leaves  and  flowers  have  fallen  off ;  the  inflorescence 
terminates  in  an  ovoid  bud  formed  by  the  flowers  which  have  not 
yet  opened  (Fig.  314,  the  left-hand  figure).  The  perianth-leaves 
are  united  (except  the  posterior  one).  The  fruit  (known  as 
a  "  Banana  ")  is  a  berry,  having  the  form  of  a  smooth,  short,  three- 
cornered  Cucumber  (as  much  as  30  cm.  in  length)  ;  inside  the 
tough  skin  is  found  a  farinaceous,  aromatic  pulp.  No  seed  is 
developed  in  the  cultivated  species. — Several  i)/?tsa-species  are  culti- 
vated in  the  Tropics  for  the  sake  of  the  fruit  (M.  paradisiaca,  M. 
sapientum) ;  for  the  fibrovascular  bundles,  M.  textilis  (Manilla  Hemp). — Their 
home  is,  no  doubt,  the  Tropics  of  the  Old  World  ;  they  were  introduced  into 
America  before  the  arrival  of  Europeans.  Musa  ensete  has  dry,  leathery  fruits ; 
an  ornamental  plant. 

In  Musa  the  barren,  posterior  stamen  belongs  to  the  inner  whorl ;  and  also  in 
Strelitzia  and  Ravenala  ;  the  latter  may  have  all  6  stamens  developed.  la 
Heliconia,  on  the  contrary,  it  belongs  to  the  outer  whorl ;  in  Heliconia  the 
perianth-leaves  are  differently  arranged,  and  there  is  only  one  ovule  in  each 
loculus.  The  three  latter  genera  have  dry  fruits  and  leaves  arranged  in  two 
rows.  In  the  "Travellers'  Palm"  (Ravenala  madagascariensis)  the  foliage - 
leaves  form  an  enormous  fan. — Tropical ;  about  50  species. 

The  order  may  be  divided  as  follows: — 1.  Musese:  Musa, 
Ravenala,  Strelitzia  in  the  Old  World.  2.  Heliconiee  :  Heliconia  in 
the  New  World. 

Order  2.  Zingiberacese.  Perianth  most  frequently  divided 
into  calyx  and  corolla.  Calyx 
gamosepalous.  Only  1  fertile 
stamen  (the  posterior,  Fig.  315, 
belonging  to  the  inner  whorl) 
with  quadrilocular  anther, 
which  encloses  the  style  in  a 
furrow  ;  the  2  stamens  in  the 
outer  whorl  are  staminodes, 
the  median  one  (the  anterior) 
is  wanting.  The  2  lateral 
staminodes  of  the  inner  whorl 
form  the  "  labellum "  (Fig. 
315  lab),  which  usually  is  the 
largest  segment  of  the  flower, 
and  is  often  bilobed.  Ovules 
many.  The  fruit  in  some  is 
a  leathery,  3-valved  capsule, 


FIG.  315.  —  Diagram  of  a  Zingiberaceou* 
flower  (Kampferia  ovalifolia):  b  bract;  v  brac- 
teole;  fc  calyx;  p1,  p2,  p3  the  petals;  sst, 
lateral  staminodes  ("wings");  lab  labellum 
(formed  of  two  staminodes);  st  the  fertile 
stamen;  *  position  of  suppressed  stamen. 
The  ovary  is  in  the  centre  of  the  diagram. 


326 


MONOCOTYLEDONES. 


with  loculicidal  dehiscence ;  in  others  it  is  more  or  less  berry-like 
and  indehiscent,  or  irregularly  dehiscent.  Straight  embryo. — The 
aerial  stem  is  seldom  developed  to  any  extent,  and  the  inflor- 
escences, which  are  (compound)  spikes  or  racemes,  often  with 
coloured  floral-leaves,  spring  in  some  (e-g-  Zingiber  officinale) 
directly  from  the  rhizome.  The  leaves  are  arranged  in  two  rows, 
— The  ovary  in  a  few  instances  {Globba  and  others)  is  unilocular, 
with  3  parietal  placentae. 

They  are  perennial  herbs  with  fleshy  and  tuberous  rhizomes,  which  are  used 
as  condiments  and  in  medicine  on  account  of  their  pungent  and  aromatic 
properties  and  also  for  starch,  dyes,  etc.  OFFICINAL  :  rhizomes  of  Zingiber 
officinale  (Ginger,  unknown  wild,  but  cultivated  generally  in  the  Tropics),  of 
Curcuma  longa  (Turmeric,  a  dye,  E.  India)  and  C.  zedoaria,  of  C.  angustifolia 
and  others  (as  E.  India  Arrowroot),  of  Alpinia  officinarum,  China  (galangal). 
"  Preserved  Ginger  "  from  Atyinia  galanga.  Similar  aromatic  materials  (vola- 
tile oils)  are  present  also,  for  example,  in  the  fruits  ;  Cardamom  fruits  and  seeds 
(from  Elettaria  cardamomum,  China,  seldom  from  E.  major). 

315  species  ;  Tropics,  preponderating  in  the  Eastern  Hemisphere,  India,  and 
especially  S.  Asia,  whence  all  the  aromatic  species  originate  ;  they  are  now 
commonly  cultivated  in  the  Tropics.  Some  are  ornamental  plants  in  green- 
houses, e.g.  Hedychium,  Costus,  etc.  Globba  (with  axillary  buds  in  the  in- 
florescence, as  in  Ficaria),  Renealmia,  Kiimpferia. 


Order  3.    Cannacese. 


FIG.  316. —  Flower  of  Canna  :  f 
ovary;  pa  calyx;  pi  corolla;  I  la- 
helium ;  st  stamens  ;  an  anther  ; 
g  stigma;  a  and  £  staminodes. 


American  herbs  without  aromatic  pro- 
perties. Flowers  asymmetric  (Fig. 
316).  Calyx  polysepalous.  The  sta- 
mens are  petaloid  (Fig.  316  st)  and 
barren  with  the  exception  of  one  (the 
posterior),  which  bears  on  one  of  its 
edges  a  bilocular  anther;  another, 
which  is  especially  large  and  coloured, 
is  termed  the  labellum,.  The  style  is 
compressed  and  leaf-like,  with  a 
small  stigma  at  the  apex.  Ovules 
numerous  in  the  3  loculi.  The  cap- 
sule is  furnished  with  warts  or  soft 
prickles.  Embryo  straight. 

Canna  (30  species;  Trop.  Am.). 
The  inflorescence  is  a  terminal  spike 
with  2-flowered  unipared  scorpioid 
cymes  in  the  axils  of  the  floral-leaves. 
Ornamental  plants :  Canna  indica,  etc. 

The  diagram  of  the  androecium   of 


8CITAM1NBJB. 


327 


the  Cannaceae  and  Marantaceee  may  be  represented  in  the  follow- 
ing manner  (calyx,  corolla  and  gynoeceum  being  omitted)  : — 


CANNACBJB. 

I'J  W 

st 
w  lab 


MARANTACE.ZE. 
w  w 


st 


IVl 


w  The  lateral  staminodes,  "  wings ;  "  st  fertile  stamen ;  *  the  suppressed  stamen ;  lab 
labellum ;  c  hood ;  wi  inner-wing. 

The  labellurn  of  the  Cannacese  corresponds  with  the  hood  of  the  Marantacese  and  not 
with  the  labellum  of  the  Zingiberacese. 

Order  4.  Marantacese.  The  flower  is  asymmetrical.  Only 
1  or  2  of  the  3  stamens  in  the  outer  whorl  are  present  as  stami- 
nodes ;  in  the  inner  whorl  2  are  petaloid  and  of  the  sixth  stamen 


FIG.  317.— Cilatliea  zebrina. 

one-half  is  developed  as  a  staminode  and  the  other  half  bears 
a  bilocular  anther.  One  ovule  only  in  each  loculus.  The  style 
is  strongly  curved  and  at  first  enclosed  in  one  of  the  staminodes 
(hood)  of  the  inner  whorl ;  later  on  it  springs  elastically  forward 
towards  the  other  staminode  (inner- wing)  of  the  same  whorl. 
The  stigma  is  very  oblique  or  2-lipped.  Two  of  the  three  loculi 
of  the  ovary,  in  some  (Haranta,  Thalia)  become  small  and  empty. 
Embryo  curved.  Leaves  in  two  rows,  with  sheath,  stalk,  and  blade 
(Fig.  317);  at  the  base  of  the  last  is  a  characteristic  swelling 
(articulus). — Phrynium,  Calathea,  Strornanthe,  Ctenanthe,  Saranthe,  etc. 
About  150  species ;  tropical,  especially  America.  The  starch  of  the  rhizome 
of  Maranta  arundinacea  is  OFFICINAL,  "  West  Indian  Arrowroot." 


323 


MONOCOTYLEDON  ES. 


Family  7.     Gynandrae. 

The  flowers  are  hermaphrodite  and  constructed  on  the  ordinary 
3-merous,  pentacyclic  type  with  petaloid,  epigynous^  strongly  zygo- 
morpliic  perianth,  and  generally  one-stamened  by  the  suppression 
of  the  other  5  stamens.  The  family  has  derived  its  name  from 
the  fact  that  the  stamen  is  united  with  the  style  into  a  "  stylar 
column  "  (except  Burmanniacece) .  All  are  herbs;  many  grow  as 
epiphytes  on  other  plants. 

This  family  and  the  Scitamine®  occupy  correspondingly  high  positions 
among  the  Monocotyledons  ;  these  two  families  may  therefore  be  placed  close 
together,  although  one  cannot  be  derived  from  the  other.  The  first  of  the  two 
orders  is  very  small,  but  the  second  is  very  rich  in  species.  The  Apostasieae 
are  best  classed  with  the  Orchidacese  and  have  no  independent  place. 

Order  1.  Burmanniacese.  This  order  forms  a  transitional 
link  between  the  Grynandree  and  the  epigynous  Liliifloree  (Ama- 
rylUdacece),  in  having  a  6-leaved  perianth,  and  6-3  stamens  ;  but 
some  have  a  labiate  perianth  (the  median  perianth-leaf  of  the  outer 
whorl  being  very  large).  The  ovary  is  most  frequently  unilocular 
with  three  parietal  placentae ;  but  in  some  it  is  3-locular  with  axile 
placentation.  Capsule.  Seeds  oo,  small,  with  endosperm.  The 
relationship  to  the  Orchidacese  is  shown  especially  in  the  very 
imperfectly  developed  embryo  and  in  the  ovary.  Small,  tropical 
herbs  (59  species)  ;  some  are  saprophytes. 

Order  2.     Orchidaceae.     The  epigynous,  petaloid  perianth  is 

strongly  zygomorphic  in 
having  the  posterior  leaf 
of  the  interior  whorl,  the 
lip  (labellum),  differing 
from  all  the  other  leaves 
in  form,  size,  and  colour 
(except  Apostasiece} ;  the 
position  of  the  labellum  is 
very  frequently  reversed, 
being  turned  forwards  and 
downwards  by  the  twisting 
of  the  ovary  (Fig.  3184). 
Only  1  of  the  stamens — 
the  anterior  of  the  external 

FIG.  318.— A  Diagram  of  an  Orchid-flower.      B,  whorl is     developed    and 

Cephalanthera.    Stylar-column:  a  anther  j  »  stigma;  bearg    &n    anther    (by    the 
at  the  foot  are  seen  scars  indicating  the  position  of  .  . 

the  parts  which  have  been  removed.  twisting  of  the  Ovary  it  IS 


GYNANDR&.  329 

turned  posteriorly  and  upwards)  ;  the  others  are  entirely  wanting 
(indicated  by  *  in  Fig.  318  A)  or  present  as  staniinodes  (Fig. 
318 .4,  (TO-)  (except  Apostasiece,  Cypripedileai);  the  filaments  are  united 
with  the  style  to  form  a  column  (Fig.  318  B),  the  stylar-cohimn*- 
(gynostemium),  and  the  anther  (a)  is  thus  placed  on  its  apex  and 
exactly  behind  or  over  the  stigma  (s).  The  anther  is  4-Iocular  ; 
the  pollen-grains  do  not  separate  (except  Apostasies,  Cypripedilect;} 
but  remain  united  either  in  tetrads  or  in  masses,  which  correspond 
to  a  pollen-mother-cell  (Fig.  320  C,  D,  E) ;  or  the  pollen-grains, 
formed  in  each  of  the  two  anther-halves,  remain  united  and  form 
one  or  a  few  wax-like  masses  (pollen-masses,  pollinia).  The  3 
carpels  form  a  unilocular  ovary  with  3  parietal,  deeply  bifid  placentre 
(except  Apostasiece,  Selenipedilum) .  Only  the  two  lateral  carpels 
are  prolonged  and  developed  into  the  stigma  (Fig.  318  J?,  s), 
while  the  one  lying  in  the  median  line,  which  is  situated  just 
within  the  anther  (Fig.  318  A),  becomes  either  rudimentary  or 
developed  into  the  "  rostellum  "  ("  a  small  beak  "),  on  which  the 
sticky  bodies  (glanduke)  arise  ;  by  aid  of  these  the  heavy,  connected 
pollen-masses  may  be  glued  to  the  insects  which  visit  the  flower, 
and  pollination  is  thus  secured  (in  Apostasiece  and  Cypripedilece  the 
3  carpels  each  contribute  to  the  formation  of  the  stigma).  The 
fruit  is  a  capsule  which  most  often  dehisces  by  6  valves,  3  of  which 
are  broader  and  bear  the  placenta?,  and  3  alternating  with  them 
are  narrower  and  barren  (except  Vanilla).  The  very  numerous 
and  exceedingly  small  seeds  have  no  endosperm,  and  have  a  some- 
what spherical  embryo  without  any  trace  of  external  organs.  The 
testa  is  membranous  and  loose. 

The  Orchids  are  all  perennial  herbs  with  diverse  habits  and 
varying  morphological  structure  (see  the  genera)  ;  the  leaves  are 
scattered,  of  the  usual  Liliaceous  form,  and  the  inflorescences  in 
all  cases  are  racemes  or  spikes  (sometimes  branched),  with  subtend- 
ing bracts,  but  without  bracteoles. 

The  forms  which  are  the  least  modified  are  described  first. 

I.  APOSTASIES.  The  perianth-leaves  are  almost  alike  and  free. 
The  column  is  straight,  with  3  equally-developed  stigmas.  Neu- 
wiedia  has  3  perfect  stamens  (1  median  of  the  outer  whorl,  and 
2  lateral  of  the  inner  whorl)  ;  Apostasia  has  only  2  perfect  (inner 
lateral)  and  one  barren  (the  median  of  the  outer  whorl),  which  how- 

1  According  to  Pfitzer,  the  column  is  the  prolongation  of  the  floral-axis 
beyond  the  insertion  of  the  perianth,  and  is  not  formed  by  the  coalescence  of 
sporophylls  (filament  and  style). 


-330 


MONOCOTYLEDONES. 


ever  may  be  entirely  wanting.  The  3  posterior  stamens  are  entirely 
suppressed.  The  pollen  is  powdery.  The  ovary  is  3-locular  with 
axile  placenta.  7  species  (Tropical  East  India,  Australia). 

2.  CYPRiPEDiLE^E.1  The  flower  is  strongly  zygomorphic  with  a 
large  boat-shaped  labellum.  There  are  two  perfect  stamens  be- 
longing to  the  inner  whorl,  and  the  median  anterior  (later  on  the 
posterior)  stamen  of  the  outer  whorl  is  transformed  into  a  large, 
barren,  shield-shaped  body  (Fig.  319) .  Selenipedilum  has  a  3-locular 
ovary,  but  Cypripedilum  (Ladies'-slipper)  has  a  unilocular  ovary 
with  3  parietal  placenta — the  typical  structure  for  the  Orchids. 


xt 


FIG.  319. — Cijpripcdilum  calceolus:  I  front  view  of  the  flower;  3  lateral  view,  after  the 
removal  of  all  the  perianth-leaves  with  the  exception  of  the  labellum,  which  has  been 
divided  longitudinally  ;  3  the  stylar-column  ;  ou  ovary;  s-s  exterior,  p  interior  perianth ; 
p'  the  labellum ;  a  the  two  fertile  stamens ;  a'  the  staminode  ;  st  the  stigma  ;  i  entrance  for 
the  insects ;  ex  exit. 

The  pollen-grains  are  separate  (not  in  tetrads)  and  all  the  3  lobes 
of  the  stigma  are  constructed  to  receive  them.  This  group  is  there- 
fore, next  to  the  Apostasiese,  the  least  modified  among  the  Orchids  ; 
in  all  the  following  groups,  one  of  the  lobes  of  the  stigma  is  differ- 
ently developed  from  the  others,  and  there  is  only  one  stamen. — 
Terrestial  Orchids — The  pollination  of  C.  calceolus  is  effected  by  the 
forcible  entrance  of  insects  into  the  boat-shaped  labellum  (Fig.  319  p')  at  i,  and 
their  escape  at  ex  (in  2)  where  the  anthers  are  situated ;  in  this  way  the 
stigmas  will  first  be  touched  and  then  the  anthers.  The  pollen- grains  are  sur- 
rounded by  a  sticky  mass  in  order  that  they  may  adhere  to  the  insects. 

1  Cypripedilum  =  Cypripedium . 


GYNANDR^!. 


331 


3.  NEOTTIE.®.  The  majority  are  terrestrial  Orchids  with  creep- 
ing, sympodial  rhizomes  ;  the  blades  of  the  leaves  are  not  de-. 
tached  from  the  stem  at  joints,  and  have  convolute  vernation. 
The  anthers  do  not  drop  off,  but  persist  in  the  withered  con- 
dition ;  their  apex,  is  brought  in  contact  with  the  rostellum  (acro- 
tonous  Orchids).  The  pollen-grains  are  united  in  tetrads,  which, 
however,  often  hang  loosely  together  in  pollinia,  attached  to  a. 
sticky  part  of  the  rostellum  ("  adhesive  disc "),  so  that  they 
adhere  to  the  insects,  and  are  by  them  transferred  to  the  stigmas. 
Spiranthes.  Listera  ;  Neottia.  N.  nidus-avis  (Bird's-nest)  is  brown  (it  has 
little  chlorophyll)  in  colour,  has  no  foliage-leaves,  and  lives  mainly  as  a  sapro- 
phyte ;  the  rhizome  is  studded  with  unbranched,  fleshy  roots  which  may  form 
buds  at  their  extremities.—  Vanilla  climbs  by  aerial  roots.  The  fruit 


FIG.  320.— A  Flower* of  OrcTu's  maculata  (front  view):  a  stamen;  b  the  cup;  n  the 
stigmas;  x  staminodes;  sp  the  spur;  spe  the  entrance  to  it;  sm--sl-sl  exterior  perianth- 
leaves;  pm  the  labellum,  and  pl-pl  the  other  2  interior  perianth-leaves.  B-E  Orchismascula : 
B  lateral  view  of  the  column ;  C  a  pollinium  with  massula?  (p),  caudicle  (c)  and  adhesive 
disc  (d)  ;  D  caudicles  with  the  cup  (V),  front  view;  the  latter  is  depressed  so  that  the 
adhesive  disc  is  seen  lying  inside  it ;  JB  a  pollinium,  more  highly  magnified  ;  some  massuise 
are  removed.  F  Ophrys  aranifera:  rostellum  and  the  base  of  the  anther-loculus ;  an 
adhesive  disc  is  seen  on  the  right. 

is  fleshy  and  hardly  opens,  or  does  so  irregularly,-— Epipactis, 
Cephalanthera. — Epipogon  and  Limodorum  are  saprophytes  without 
chlorophyll. 

4.  OPHRYDE^E.  Anthers  2-locular,  not  falling  off,,  on  a  very 
short  column.  The  anther  is  united  at  its  base  with  the  rostellum 
(basitonous  Orchids,  Fig.  320  A,  J5),  while  in  all  other  Orchids  it  is 
connected  at  the  apex  (acrotonous  Orchids).  The  pollen-grains  in 
each  loculus  are  united  into  small  "  masses  "  (massulae),  each  ofr 


332  MONOCOTYLEDONES. 

which  corresponds  to  a  pollen-mother-cell  in  the  anther,  and 
which  hang  together  by  elastic  threads  (Fig.  320  C,  E).  Each 
pollinium  is  attached  at  the  base  by  a  stalk  (caudicle)  to  an 
adhesive  disc,  formed  by  the  modified  stigma  (rostellum),  and  is 
easily  liberated  from  it  (Fig.  320  (7,  D,  F).  The  pollinium,  which 
is  formed  in  an  anther-loculus,  together  with  its  caudicle  and 
adhesive  disc,  is  termed  "  pollinarium "  (Fig.  320  0). —  All 
Ophrydese  are.  terrestrial  with  tuberous  roots,  two  of  which  are 
present  in  the  flowering  period,  an  older  one  (from  the  preceding 
year)  containing  the  nourishment  for  the  flowering-shoot  of  the 
year,  and  a  young  one  which;  is  intended  to  contain  the  reserve 
material  for  the  following  year.  Inflorescence  terminal. 

Orchis.  The  lip  has  a  spur ;  each  of  the  club-like  pollinia  is 
attached  to  its  own  adhesive  disc,  the  discs  being  enclosed  in  a 
•common  pouch  formed  by  the  rostellum  (Fig.  320  0,  D).  Tubers 
ovate,  undivided  :  0.  morio,  ma  scuta  ;  tubers  palmate  :  0.  incarnata,  maculata, 
majalis.—Ophrys;  no  spur,  the  two  adhesive  discs  are  each  enclosed  in 
a  separate  pouch  (Fig.  320  F~). — Anacamptis  and  Serapias  have  one 
adhesive  disc. — Habenaria,  Gymnadenia,  Platanthera,  Herminium, 
Nigritella,  Gceloglossum,etc.,  have  naked  adhesive  discs  (no  rostellum). 

5.  EPIDENDRE^:.      Acrotonous     Orchids    with     deciduous     anthers    (except 
Malaxis) ;    2-8   wax-like  pollinia,   with   or    without   caudicles ;    generally   no 
adhesive  discs.    Malaxis  (the flower  is  twisted  through  a  complete  circle,  causing 
the  labellum  to  be  turned  upwards),  Sturmia  and  Corallorhiza  *  (Coral-root) ; 
the  latter  has  a  creeping,  coral-like  rhizome  ivithout  roots,  and  is  destitute   of 
chlorophyll  except  in  the   ovary.      The    other  two   somewhat  resemble   the 
tropical  Orchids  in  having  the  lower  internodes  of  the  axis  of  the  inflorescence 
tuberous.     Liparis ;   Calypso.     Most  of  the  genera  are  tropical  epiphytes  and 
many  have  aerial,  green  tubers  formed  from  one  or  mor*e  stem-internodes  ; 
Dendrobium,  Eria,  Phaius,  Bletia,  Epidendrum,  Cattleya,  Lcelia,  Pleurothallis, 
Restrepia,  Masdevallia,  Bulbophyllum,  etc. 

6.  YANDEX      These  resemble  the  preceding  but  have  only  2  wax-like  pollinia 
in  each  anther,  which  are  attached  by  a  caudicle  to  the  adhesive  disc  of  the 
rostellum.    Nearly  all  are  tropical  epiphytes.    Stanhopea,  Catasetum,  Maxillaria, 
Oncidium,  Vanda,  Polystachya,  etc. 

6,000  (105000?)  species.  The  majority  live  in  the  Tropics  and  occur,  es- 
pecially, as  epiphytes  on  trees  or  in  the  crevices  of  rocks,  to  which  they  are 
attached  by  aerial  roots.  These  aerial  roots,  like  those  of  Araceae,  are  covered 
by  several  layers  of  spirally-thickened  cells  (tracheides)  which  contain  air 
and  form  the  velamen — an  apparatus  to  absorb  moisture  from  the  air.  The 
roots  have  a  white  appearance  when  the  cells  are  filled  with  air,  which 
changes  to  a  greenish  hue  when  they  are  filled  with  water,  the  chlorophyll 
then  shining  through.  They  generally  have  horizontal  rhizomes;  the 

1  Corallorhiza  =  CoralUorrhiza. 


GYNANDRJ1.  333 

ascending  shoots,  which  bear  the  foliage-leaves,  may  vary,  but  they  very  often 
swell  and  assume  the  form  of  a  tuber,  which  persists  for  several  years  fresh 
and  green  after  the  leaves  have  fallen  off  (Fig.  321).  Vanilla  is  an  exception 
(see  above).  Our  Orchids  are  all  terrestrial  (or  marsh-plants) ;  the  largest 
number  of  species  is  found  in  calcareous  soils. 

POLLINATION  takes  place  principally  by  means  of  insects,  but  self-pollina- 
tion occurs  in  some.  The  lip  serves  as  a  landing-stage  for  the  insect  visitors, 
which,  on  sucking  the  honey,  cause  the  adhesive  discs,  with  the  pollinia 
attached  to  them,  to  adhere  to  their  bodies  (generally  to  the  probosces)  and  so 
carry  them  away  to  other  flowers.  In  some  species  parts  of  the  flower  are 
sensitive  or  irritable,  which  has  some  connection  with  the  pollination. 
Without  doubt  there  are  a  great  many  biological  differences  which  are  closely 
connected  with  the  infinite  multiplicity  of  forms ;  Darwin  (1862)  has  already 


FiQ.  321. — Chysia  bractescens. 

shown  an  enormous  variety,  never  even  dreamt  of  before,  in  the  European 
species.  The  genus  Catasetum  has  $  -  ?  -  and  $  -plants  with  flowers  of  such 
different  appearances  that  they  have  been  classed  in  various  genera  (Myanthus, 
Monacanthus).  Platanthera  is  pollinated  by  hawk-moths ;  Ophrys,  by  flies  ; 
Epipactis  latifolia,  by  wasps  ;  Orchis,  by  bees,  especially  humble-bees,  etc. 

The  DISTRIBUTION  OF  SEEDS  is  effected  by  the  wind,  the  seeds  being  so  ex- 
ceedingly small  and  light.  Many  species  moreover  have  peculiar,  elater-like, 
fine,  hygroscopic  hairs  in  the  ovary,  which  eject  the  seeds  in  a  manner  similar 
to  the  elaters  of  the  Liverworts.  , 

The  USES  are  few,  mostly  as  ornamental  plants  in  conservatories.  The 
tubers  of  several  Orcfa's-species  are  OFFICINAL  ;  they  contain  starch  and  mu- 
cilage and  are  used  as  "  salep."  The  fruits  of  Vanilla  planifolia  are  used  as 
condiments  and  differ  from  other  Orchid-fruits  in  being  rather  fleshy  and 
in  dehiscing  irregularly ;  the  seeds  are  very  small,  shining  and  black. 


334  DICOTYLEDONES. 

Class  II.    Dicotyledones. 

In  this  class  THE  EMBRYO  lias  2  seed-leaves,  a  rule  from  which 
there  are  few  exceptions  (e.g.  Ficaria,  Cyclamen,  Pinguicula, 
certain  species  of  Corydalis,  with  only  1 ;  and  a  few,  mostly  para- 
sitic forms,  e.g.  Monotropa,  Orobanche,  Pyrola,  entirely  without 
cotyledons).  On  germination  the  cotyledons  nearly  always  raise 
themselves  above  the  ground  as  green,  assimilating  leaves  and  are 
then  termed  aerial  or  epigean,  in  contradistinction  to  the  under- 
ground or  hypogean  which  are  always  buried.  The  structure  of 
the  seed  varies  (endospermous  or  exendospermous)  ;  the  embryo 
may  be  straight  or  curved.  In  many  instances  the  primary  root 
grows  as  a  vigorous  tap-root,  with  weaker  branches  arising  acro- 
petally  (in  annuals,  biennials,  many  perennials,  especially  woody 
plants)  ;  but  in  a  large  number  of  herbaceous  perennials,  which 
have  rhizomes,  the  root  behaves  very  much  as  in  the  Monocoty- 
ledons. The  roots  generally  increase  in  thickness  by  means  of  a 
cambium. 

THE  STEM,  when  seen  in  transverse  section,  has  its  vascular 
bundles  arranged  in  a  ring ;  in  reality,  however,  they  form  a 
kind  of  cylindrical  network  in  the  stem  ;  the  bundles  are  open, 
and  thickening  takes  place  by  means  of  a  cambium  ;  annual  rings 
are  formed  in  the  perennial  stems.  There  is  a  rich  and  very  varied 
form  of  branching.  The  two  first  leaves  of  a  shoot  (fore-leaves) 
are  placed  nearly  always  to  the  right  and  to  the  left ;  the  same 
arrangement  is  found  in  the  two  first  leaves  developed  on  the 
flower-stalk,  and  these  are,  as  a  rule,  the  only  two  ;  they  are  found 
below  the  calyx  and  are  usually  termed  the  "  bracteoles"  It  has 
become  customary  to  indicate  the  bracteoles  by  the  letters  a  and  /?, 
according  to  their  sequence  of  growth,  and  in  that  sense  these 
letters  will  be  employed  in  the  following  diagrams. 

THE  ARRANGEMENT  OF  THE  LEAVES  varies  very  much  ;  there  is 
also  a  great  variety  of  shapes  in  the  leaves  and  their  venation, 
but  the  linear  leaves,  with  parallel  venation,  so  frequent  in  the 
Monocotyledons,  are  seldom  met  with,  as  also  the  large  sheaths 
(though  the  sheath  is  well  developed  in  the  Umbelliferous  plants) ; 
stipules  occur  much  more  frequently. 

THE  FLOWER  is  most  commonly  cyclic,  but  acyclic  or  hemicyclic 
forms  also  occur.  The  type  which  may  be  taken  as  a  basis 
consists  in  the  majority  of  instances,  as  in  the  Monocotyledons,  of 
5  whorls,  of  which  the  4  outer  ones  (calyx,  corolla,  and  the  2 


DICOTYLEDONES.  335 

whorls  of  stamens)  are  most  frequently  4  or  5  in  number  and 
placed  in  regular  alternation,  whilst  the  innermost  one  (the  car- 
pels) has  generally  fewer  members,  probably  on  account  of 
space  (Figs.  3(50,  361,  421,  429,  487,  etc.).  Trimerous  (Figs.  384, 
387,  etc.)  flowers,  or  those  in  which  the  members  of  the  flower  are 
in  threes  or  a  multiple  of  three,  also  occur,  as  well  as  dimerous 
flowers  ;  other  numbers  are  rare.  It  is  of  the  greatest  importance 
in  connection  with  the  relative  position  of  the  members  of  the 
flower  to  the  axis  and  bract  (orientation),  whether  the  bracteoles 
are  typically  present  (even  though  they  may  not  be  developed),  or 
are  typically  absent.  If  there  are  2  bracteoles  present,  then  their 
position  in  a  pentamerous  flower  is  often  as  follows :  the  first  sepal 
turns  obliquely  forward,  the  second  is  posterior  and  median,  the 
third  obliquely  forward,  the  fourth  and  fifth  obliquely  backward  ; 
quincuncial  aestivation  is  often  found  in  these  buds  (Figs.  360,  429,. 
471,  475,  584).  The  first  and  third  leaves,  in  the  following  chapters,, 
are  most  frequently  alluded  to  as  the  "  anterior,"  the  fourth  and 
fifth  as  the  "  lateral  "  leaves.  The  reversed  arrangement,  with  the- 
median  sepal  in  the  front,  occurs  for  instance  in  Papilionacece 
(Fig.  511),  Lobeliacece  (Fig.  594),  Bhodoracece.  If  any  bracteoles  are 
present  below  a  tetramerous  flower,  the  relation  is  generally  that 
2  sepals  (the  first  ones)  stand  in  the  median  plane,  the  two  next 
ones  transversely  (Fig.  393),  and  the  corolla  then  adopts  a  diagonal 
position  (Fig.  397) ;  but  a  diagonal  position  of  the  calyx  generally 
shows  that  the  flower  is  not,  strictly  speaking,  tetramerous,  as  in 
Plantago  (Fig.  567),  Veronica  (Fig.  559  C)  and  others. 

If  the  bracteoles  are  not  typically  present,  then  the  position  of 
the  sepals  is  changed  accordingly,  and  the  two  outer  sepals  en- 
deavour to  assume  the  position  which  the  bracteoles  would 
otherwise  have  occupied,  e.g.  in  Primula  (Fig.  547).  Other 
positions  are  also  found  when  the  number  of  bracteoles  is  more  or 
less  than  two. 

The  leaves  which  follow  the  sepals  occupy  definite  positions 
with  regard  to  them,  which  we  may  consider  later.  An  arrange- 
ment must,  however,  be  mentioned  here;  when  the  flower  is- 
"  diplostemonous,"  that  is,  has  two  whorls  of  stamens  (thus,  Sn,  Pn, 
An  +  n),  these  may  be  arranged  in  two  ways.  Either  the  first- 
formed  whorl  of  stamens,  which  are  termed  the  "calyx-stamens," 
stands  directly  in  front  of  the  sepals  (that  is  "  episepalous "), 
and  is  the  outermost  whorl,  and  in  this  case  a  regular  alternation, 
takes  place  between  sepals,  petals  and  the  two  whorls  of  stamens, 
\v.  B.  z 


336  DICOTYLEDONES 

which  is  also  continued  into  the  carpels  if  their  number  is  the 
same  as  that  of  the  other  whorls  :  the  carpels  are  then  placed 
opposite  the  sepals  (Fig.  278)  and  the  flower  is  isomerous  and  Gn 
should  be  added  to  the  formula  above.  Or,  the  calyx-stamens 
form  the  innermost  whorl,  and  the  corolla-stamens,  which  are  sub- 
sequently formed  ("  epipetalous  "  stamens),  stand  outside  these 
(Figs.  360,  429)  ;  if  the  number  of  carpels  is  the  same  as  that  of 
the  preceding  whorls,  they  are  often  placed  right  in  front  of  the 
petals  and  the  jcorolla-stamens.  The  first-mentioned  arrangement 
is  termed  Diplostemonous,  and  the  second  ObdipLostemonous.  Both 
arrangements  may  be  found  in  one  and  the  same  order,  e.(f.  Carjophyllaceae. 
The  size  and  relation  of  the  members  of  the  flowers,  and  also  the  contact  with 
other  members  in  the  early  stages  of  their  development,  play  an  important 
part  in  determining  the  arrangement. 

The  great  number  of  structural  arrangements  found  in  this 
enormously  large  class,  may,  as  is  the  case  in  the  Monocotyledons, 
be  further  varied  by  suppression  and  division  of  certain  leaves 
(especially  the  stamens).  Instances  of  this  will  occur  in  the 
following  (Figs.  559,  568.— 426,  441,  445,  etc.). 

The  Dicotyledons  were  formerly  divided  into  3  sub-classes  : 
Apetalae  (those  without  corolla),  Sympetala3  or  Gamopetalae  (those 
with  the  petals  united),  and  Choripetalae  or  Polypetalae  (the  petals 
not  united).  This  division  has  now  been  abandoned  because  it  has 
been  proved  that  the  Apetalae  were  merely  reduced  or  incomplete 
forms  cf  the  Choripetalae,  and  they  have  therefore  been  distributed 
among  the  various  families  of  the  latter  sub-class. 

With  regard  to  the  Sympetalse  (or  Gamopetaloe)  it  may  be 
stated  that  they  form  to  a  very  great  extent  a  closely  connected  and 
natural  group,  having  in  common  not  only  the  character  that  the 
corolla  is  gamopetalous  and  the  stamens  united  with  it  (this  being 
also  found  in  the  Choripetalaa),  but  also  a  great  many  others  (such 
as  persistent  calyx,  cyclic  flowers  with  the  formula  S5,  P5,  A5  and 
as  a  rule  G2,  the  two  carpels  being  united  to  form  the  ovary  ;  seeds 
with  a  thick  integument  and  a  very  small  nucellus).  They  are 
therefore  considered  as  an  independent  sub-class,  and  must  be 
placed  at  the  close  of  the  system  of  classification  as  the  forms 
which  presumably  have  arisen  the  latest.  In  the  future  systems 
of  classification  this  arrangement  will  very  probably  be  changed, 
and  the  first  families  of  the  Sympetalae,  the  Bicornes  and  others 
will  for- instance  be  to  a  certain  extent  united  with  the  families  or 
.orders  of  the  Choripetalae.  The  Sympetalae  may  certainly  be  con- 


SALIC1FLORJ1.  337 

sidered  as  the  youngest  types,  the  strongly  pronounced  metamor- 
phosis supporting  this  theory,  as  also  the  formation  of  the  integu- 
ment of  the  ovule,  the  one  thick  integument  being  undoubtedly 
derived  from  the  coalescence  of  two — a  holochlamydeous  ovule,  etc. 

The  Apetalae  and  Choripetalae  are  united  into  one  sub-class.  The 
leaves  of  the  perianth  in  this  case  are,  as  a  rule,  free  from  each 
other,  the  structure  of  the  flowers  presents  many  differences,  and 
the  ovules  have  as  a  rule  2  integuments  and  a  large  nucellus.  Con- 
siderable uncertainty  still  prevails  regarding  the  arrangement  and 
the  relationship  of  the  individual  families  of  the  Choripetalee,  and 
some  of  the  following  families  are  hardly  quite  natural ;  but  the 
best  arrangement  arrived  at  so  far  has  been  adopted  here. 

At  the  beginning  of  the  book  a  review  of  the  orders  of  the 
Dicotyledons  will  be  found. 

Sub-Class  1.    Choripetalae.     Petals  free. 

Family  1.     Saliciflorse. 

Trees  and  shrubs,  which,  in  the  structure  of  the  vegetative  shoot 
and  the  catkin-like  inflorescences,  resemble  the  Querciflorae,  but 
the  structure  of  the  flower  differs  so  much  from  them,  that  the 


FIG.  322.— Male  and  female  catkins  of  Salix  caprea. 

only  order  brought  under  this  heading — Salicacece — well  deserves 
to  be  separated  and  to  form  a  family  of  its  own,  the  nearest 
relatives  of  which  are  still  doubtful.  As  Juglandaceas  and  Myricaceaj 
also  deserve  to  be  placed  in  a  special  family,  the  name  Amentacece  (CatJdn- 
bearers),  bitherto  applied  to  all  of  tbese  plants,  cannot  be  retained  as  tbe  name 
of  a  family. 

There  is  only  one  order. 


338 


DICOTYLEDONES. 


Order.  Salicaceae  (Willows).  Trees  with  simple,  scattered, 
stipulate  leaves.  Dioecious.  The  flowers  are  arranged  in  simple  inflor- 
escences (spikes  or  racemes)  which  are  termed  catkins,  and  which 
fall  off  as  a  whole  after  flowering  (  £ )  or  after  the  ripening  of  the 
fruit  (?)  (Fig.  322).  The  perianth  is  very  imperfect1  or  wanting, 
particularly  in  Salix  (Fig.  323  o)  ;  the  ^-flower  with  2-several 
stamens  and  without  any  trace  of  a  carpel  (a,  b,  c)  :  the  ?  -flower  has 
a  free  bicarpellate  ovary,  unilocular,  and  formed  from  2  lateral  car- 
pels with  2  parietal  (median)  placenta?  and  generally  co  ovules ;  the 
style  divides  into  two  stigmas  (d,  e,  f).  The  fruit  is  a  two-valved 
capsule  and  the  very  small  seeds  bear  a  tuft  of  hairs  at  the  base.. 
Endosperm  absent. —  The  catkins  are  situated  on  dwarf -branches,  which' in 
some  species  often  develop  before  the  leaves  and  bear  at  their  base  only 
scale-leaves  ;  in  others  foliage-leaves  are  borne  beneath  the  catkins.  The  vege- 


Fis.  323. — Salia;:  male  flowers  of  S.  pentandra  (a),  S.  durita  (b),  S.  rubra  (c);  female 
flowers  of  S.  aurita  (d),  S.  nigricans  (e),  S.  mollissima  (/). 

tative  bud  commences  with  2  bud-scales  which  are  united  on  the  anterior  side 
into  a 'scale.  The  capsule  opens  by  the  dorsal  suture.  The  seed-hairs  spring 
from  the  funicle. 

Salix  (Willow)  has  short-stalked,  most  frequently  lanceolate 
leaves  and  erect  catkins  with  undivided  bracts  (Fig.  322).  The 
flowers  are  naked ;  1  (o  in  a-f)  or  2  yellowish  glands  situated  in 
the  median  line.  In  the  £  -flower  generally  two  stamens,  situated 
laterally  like  the  carpels  in  the  ?  -flower.  Various  forms  are  seen  in 
Fig.  323.— The  terminal  bud  of  the  branches  often  aborts  regularly,  the  upper- 
most lateral  bud  taking  its  place. 

Populus  (Aspen,  Poplar)  has  long-stalked,  more  or  less  round  or 
cordate  leaves  with  drawn-out  apex ;  catkin  pendulous ;  lobed 

1  This  is  Eichler's  view. — According  to  Drude  the  perianth  is  absent ;  at  the 
base  of  the  bracts,  a  nectary  or  cup-like  disc.  Prantl  holds  the  same  view. 
According  to  Pax  the  perianth  is  absent,  but  there  is  a  disc  cup-like,  or  reduced 
to  a  single  toothed  scale. 


CASUARINIFLOKJi.  339 

bracts ;  perianth  cup-like  with  oblique  edge ;  stamens  usually 
numerous  ;  stigmas  often  divided. — P.  tremula  (Aspen)  has  received  its 
name  from  the  tremor  of  the  leaves  :  cf.  "  to  shake  like  an  aspen  leaf." 

POLLINATION.  The  Poplars  are  wind-pollinated.  The  Willows  Lave  sticky 
pollen  and  are  pollinated  by  insects.  The  catkins  of  the  Willows,  especially  the 
<? ,  are  more  conspicuous,  from  the  numerous,  closely-packed,  yellow  flowers, 
rich  in  honey  and  pollen.  The  catkins  often  appear  before  the  foliage  and  so  are 
much  more  easily  seen,  whilst  at  this  time  of  the  year  the  number  of  competing 
honey-flowers  is  smaller,  and  the  insect  visits  consequently  more  numerous. 
On  many  catkins  of  the  Willow  the  flowers  open  earliest  on  the  side  which  is 
turned  towards  tlie  sun  and  in  descending  order,  i.e.  the  upper  flowers  develop 
before  the  lower  ones.  Hybrids  frequently  appear. 

There  are  about  180  species  existing  in  the  northern,  cold  and  temperate 
latitudes.  Some  in  the  Polar  regions  are  scarcely  more  than  an  inch  in  height, 
and  have  a  creeping  rhizome  (Salix  herb  acea,  polar  is,  reticulata).  Fossil  forms 
are  found  in  the  Tertiary  and  perhaps  also  in  the  Upper  Cretaceous. 

USES.  Principally  for  ornamental  trees,  as  they  grow  very  quickly  and  are 
easily  propagated  by  cuttings,  S.  babylonica,  Weeping  Willow;  S.  purpurea  ; 
Populus  alba,  Silver  Poplar ;  P.  pyramidalis,  Pyramid  Poplar — a  form  of 
P.  nigra ;  P.  monilifera,  Canadian  Poplar.  The  wood  is  very  poor  and  little 
used  ;  the  branches  of  many  Willows  are  cultivated  for  basket-making,  etc.  The 
wood  of  the  Aspen  is  used  for  matches.  The  bark  contains  tannin  and,  in  many 
Willows,  a  very  bitter  extract,  Salicin  (S.  pentandra,  fragilis).  Salicylic  acid 
(officinal)  is  obtained  from  Salix.  Balsam  is  extracted  from  the  buds  of  many 
Poplars,  especially  when  the  leaves  are  shooting. 

Family  2.     Casuariniflorae. 

Trees  with  verticillate,  scale-like  leaves  forming  sheaths  at 
the  nodes.  Monoecious.  Flowers  unisexual,  ^-flowers  in  cat- 
kins; ?  in  short  spikes.  Pollen-tube  entering  the  ovule  at  the 
chalaza,  and  not  through  the  micropyle.  Ovary  1-seeded, 
uniiocular.  Carpels  uniting  into  a  multiple  fruit.  Only  one 
order. 

Order.  Casuarinaceae.  Trees  (30  species),  from  Australia 
and  certain  parts  of  S.E.  Asia,  with  peculiar,  equisetum-like 
appearance.  The  leaves  are  verticillate,  scale-like  and  united  into 
sheaths.  The  internodes  are  furrowed.  Branching  verticillate. 
The  unisexual  flowers  are  situated  in  catkins  or  short  spikes.  The 
^-flower  has  a  central  stamen,  surrounded  by  2  median,  scale- 
like  perianth-leaves  and  2  lateral  bracteoles.  The  $  -flower  has  a 
1-chambered  ovary  (2  ascending,  orthotropous  ovules),  no  perianth, 
but  2  large,  lateral  bracteoles  which  finally  become  woody  and 
form  two  valves,  between  which  the  nut-like  fruit  is  situated. 
The  multiple-fruits  therefore  resemble  small  cones. —  Casuarina 
equisetifolia,  cultivated,  gives  "  iron- wood." 


340  DICOTYLEDONES. 

[The  Casuarinas  differ  from  the  ordinary  Dicotyledons  in  many  important 
respects  which  may  be  briefly  summarised  thus  : — The  bicarpellate  ?  -flower  has 
a  well-pronounced  stylar-cylinder  terminated  by  two  stigmas,  but  the  cavity  of  the 
ovary  closes  very  soon  after  its  formation,  and  in  it  are  developed  two  parietal 
ovules  ;  these  are  united  by  a  bridge  of  cellulose  to  the  styJar-cylinder  or  summit 
of  the  ovary,  and  hence  the  ovules  are  connected  with  the  walls  of  the  ovary  by 
the  bridge  (above),  as  well  as  by  the  funicle  (below).  The  archespore  is  developed 
from  the  hypodermal  cells  at  the  summit  of  the  nucellus,  two  primordial 
mother-cells  are  first  formed  and  from  these  by  tangential  divisions  a  central 
cylindrical  mass  of  cells  (sporogenous-tissue)  is  produced  which  is  surrounded 
by  tapetal  cells.  The  cells  of  the  sporogenous  tissue  correspond  to  the  mother- 
cells  of  the  embiyo-sac  of  other  Angiosperms;  they  divide  transversely  and 
from  16-20  macrospores  are  formed  together  with  inactive  cells  which  are  not 
crushed  together  as  in  the  case  of  other  Phanerogams.  The  sexual  apparatus 
is  developed  from  a  single  cell,  but  the  number  of  cells  composing  this  tippar- 
atu-<  is  subject  to  variation,  the  oosphere  being  accompanied  by  one  or  two- 
neigbbouriug  cells  wbich  resemble  canal-cells  rather  than  synergidae.  The 
sexual  apparatus  is  found  in  the  majority  of  the  macrospores,  but  in  most  of 
these  it  remains  as  a  number  of  naked  cells  ;  while  in  the  fertile  macrospores- 
the  cells  are  invested  by  walls  of  cellulose  (usually  only  one  fertile  macrospore  is. 
found  in  each  ovule).  Antipodal  cells  are  never  developed.  The  macrospores- 
elongate  considerably  towards  the  chalaza,  into  which  some  penetrate.  The 
pollen-tube  traverses  the  stylar  cylinder  and  enters  the  ovules  at  the  chalaza,. 
3ts  passage  through  the  the  tissue  of  the  nucellus  being  assisted  by  the  pro- 
tongation  of  the  macrospores.  About  the  centre  of  the  nucellus  the  pollen- 
tube  is  ruptured ;  the  apical  portion  which  alone  takes  part  in  the  fertilisation 
being  firmly  attached  to  the  macrospore.  Although  the  actual  impregnation 
has  not  been  observed,  Treub  considers  that  the  endosperm  begins  to  be- 
formed  before  fertilisation.] 

Family  3.     Querciflorse. 

Trees  and  shrubs  with  small,  unisexual,  monoecious  flowers, 
having  no  perianth  or  a  simple  inconspicuous  one.  The  $  and  ? 
flowers  are  very  different  and  generally  placed  in  separate  inflor- 
escences. The  ^-flowers  are  most  often  adnate  to  the  tracts.  The 
stamens  are  placed  opposite  the  perianth-leaves,  whera  tfoey  are  pre- 
sent in  equal  numbers.  The  ?  -flower  is  naked,  or  has  a  superior 
perianth.  The  ovary  at  the  base  is  2  or  3- (-6)  locular  with  1  or 
2  pendulous  ovules  in  each  loculus,  only  one  of  which  is  developed; 
the  fruit  is  a  one-seeded  nut ;  endosperm  absent ;  embryo  straight. 
The  inflorescences,  which  are  either  compound  and  mixed  (small 
dichasia  in  spikes)  or  simple,  are  here  also  termed  catkins;  but,, 
strictly  speaking,  this  term  is  applied  to  the  (^-inflorescences  only. 
In  all  Querciflorse  the  leaves  are  scattered  (usually  in  2  rows) 
simple,  and  penninerved,  and  with  deciduous  stipules. 


QUERCIFLORJ!. 


341 


It  is  worthy  of  remark  that  in  Betnlacece,  Corylacece  and  Quercus  the  ovules, 
and  to  some  extent  the  loculi  of  the  ovary  are  not  developed  till  after  pollin- 
ation, so  that  the  development  of  the  pollen-tube  proceeds  very  slowly.  The 
smallness  of  the  flowers,  the  absence  of  honey,  the  dryuess  and  lightness  of  the 
pollen,  the  size  of  the  stigma  and  the  abundance  of  hairs  found  on  many 
stigmas  are  all  adaptations  for  wind-pollination.  It  is  also  an  advantage  that 
the  flowers  are  generally  pollinated  before  the  foliage-leaves  are  developed,  thus 
preventing  the  pollen  being  entangled  by  the  leaves. 

The  two  orders  Betulacece  and  Corylacece  mentioned  here  are  by  other  authors 
united  into  one  order.  [It  is  doubtful  whether  these  two  should  be  retained  in 
the  family  Querciflorae,  as  recent  researches  (p.  273)  have  shown  that  they 
differ  from  the  Cupu'iferae  in  many  important  points,  and  agree  with  the 
Casuavinas  in  the  fact  that  the  pollen- tube  enters  the  ovule  through  the 
chalaza.J  ^ 

Order  1.  Betulaceae  (Birches).  Monoecious,  with  thick, 
cylindrical,  compound  $  and  $  inflorescences  (2-  or  3-flowered 
dichasia  in  a  spike  with 
spirally-placed  floral- 
leaves)  (Figs.  324,  326, 
328).  When  the  perianth 
in  the  (^-flower  is  com- 
pletely developed,  it  is 
composed  of  4  somewhat 
nutted  leaves,  which  are 
placed  opposite  the  4 
stamens  (Figs.  325,  326/1). 
The  female  flowers  are 
naked  ;  the  ovary  is  bi- 

locular,     with     two     Styles  FIB.  32*.— 4lnun  glutino«w.    Brau,-h  of  Alder  with 

<J  -(n)  and  ?  -(m)  catkins :    fc  bud  ;    6  fruit-bearing- 
catkin  ("cone.") 


and  one  pendulous  ovule  in 
each    loculus.       The    sub- 


tending floral-leaves  unite  with  the  bracteoles  and  form  a  3-5-lobed 
cover-scale;  which  is  not  attached  to  the  fruit  (Figs.  325  D,  326$). 
Fruit  a  nut  without  cupnle  (see  Corylacece  and  Cupuliferw).  In  the 
bud  the  leaves  are  flat.  The  stipules  are  deciduous.  On  germination  the 
cotyledons  are  raised  above  the  ground.  Terminal  buds  are  only  found  on  old 
Alder  trees  ;  the  Birch  has  s.ympodial  branches. 

Alnus  (Alder)  (Figs.  324-326).  In  the  majority  of  species  the 
^  -  and  ?  -catkins  are  both  developed  in  the  year  previous  to  their 
flowering,  and  pass  the  winter  naked  and  bloom  before  the  leaves 
expand.  £  -flower  :  4  stamens.  $  -flower  :  the  5-lobed  cover- 
scales  of  the  ?  -catkin  are  woody  and  remain  attached  to  the  axis, 
so  that  the  entire  catkin  when  ripe  resembles  a  small  cone  (Fig. 


342 


DICOTYLEDONES. 


FIG.  325.— Alnus  glutinosa:  A  dicha^ium  of  rf -flowers  seen 
from  the  front ;  B  the  same  from  inside  ;  C  the  same  from  the 
back ;  D  dichasium  of  ?  -flowers  with  subtending-leaf  and  four 
liracteoles.  The  letters  b,  a,  ft,  ft',  ft,  are  the  same  as  in  Fig. 


324  ?>).  Each  cover-scale  supports  two  winged  or  wingless  nuts. 
In  the  native  species  of  Alder  the  buds  are  stalked  (Fig.  324  k).  The  bud- 
scales  are  formed  by  the  stipules  of  the  lowest  leaves. 

-  Betula  (Birch).  The  <$  -catkins,  in  the  native  species,  appear  in 
autnmn,  the  ?  -catkins  in  the  flowering  year  on  leaf- bearing,  short- 
lived shoots.  <£ -flowers  :  2  stamens,  divided  (Fig.  328  A).  The  3- 

lobed  cover-scales 
(Fig.  327  a)  of 
the  $  -catkin  are 
detached  from, 
the  axis ;  each 
cover-scale  sup- 
ports 3  broadly- 
winged  nuts  (6). 
The  stem  has  cork 
with  annual  rings. 
The  young  twigs  and 
leaves  have  aromatic 
resin  glands. 

THE  INFLORESCEN- 
CES OF  THE  ALDER. 
— In  the  axil  of  each 
cover-scale  [b  in  the  Figs.]  is  situated,  in  the  # -catkins  (Figs.  326  A,  325  A-C) 
a  3 -flowered  dichasium,  the  flowers  of  which  have  a  4-partite  perianth,  the 
posterior  perianth-segments  being  sometimes  almost  suppressed,  and  4  stamens 
with  undivided  filaments.  In  the  ?  -catkin  (Figs.  325  D,  326  C)  a  2-flowered 

dichasium  is  found,  the  middle  flower 
being  suppressed  (indicated  by  a  star  in 
C).  In  both  instances  the  inflorescences 
have  two  bracteoles  (ct-/3)  and  the 
flowers  borne  in  their  axils  have  each 
one  bracteole  (£'),  the  other  one  (a')  be- 
ing suppressed  and  therefore  in  326  A 
and  C  only  represented  by  a  dotted 
line ;  these  four  bracteoles  unite  with 
the  cover-scale  (6)  which  supports  the 
entire  dichasium,  to  form  the  5-lobed 
"  cone-scale  "  (Fig.  326  B)  which  in  the 
?  -catkin  eventually  becomes  woody. 

THE  INFLORESCENCES  OF  THE  BIRCH. — 
A  3-flowered  dichasium  is  situated  in 
the  axil  of  the  cover-scale  in  both  <J- 
and  ?  -catkins  (Fig.  328  A,B)',  only  the 
central  flower  has  bracteoles  (a-/J)  (the 
lateral  flowers  having  no  bracteoles),  and  these  bracteoles  unite,  as  in  the  Alder, 
with  the  supporting  cover-scale  (b),  and  form  a  three-lobed  cone-scale  (Fig.  327  a). 


FIG.  326.  —  Alnna  gluiinosa:  diagram 
of  dicbasia  of  <J  (A)  and  ?  (C)  catkins; 
B  a  cone-scale.  All  the  bracteoles  in  A 
and  C  are  slightly  pressed  from  their 
normal  position. 


QUERCIFLOR&.  343 

While  the  ?  -flower  exactly  resembles  that  of  the  Alder,  the  reduction  of  the 
cf  -flower,  already  described  in  the  Alder,  is  carried  further,  so  that  often  only 
the  2  median  perianth-leaves  are  developed  (Fig.  328  A) ;  there  are  also  only 
2  stamens,  these  being  deeply  cleft,  while  the  other  2  are  suppressed. 

About  50  species ;  N.  Temp. — Fossil-forms  certainly  occur  in  the  Oligocene. 
During  the  Glacial  period  the  Dwarf -birch  (B.  nana)  extended  over  Europe  ;  at 
the  present  time  it  is  confined  to  the  moors  and  mountains  of  N.  Europe  and 
N.  America  and  Asia.  Wind-pollinated. 

USES. — Important  forest  trees.  The  bark  contains  tannic  acid.  The  tar  of  the 
Birch  is  used  in  the  preparation  of  Russia  leather  ;  whilst  its  spring  sap  is  very 
saccharine,  and  is  used  in  some  places  for  making  a  fermented  di'ink.  Its  ex- 
ternal bark  is  used  for  roofing,  for  baskets,  etc. 


FIG.  327.— Betula  vermcosa  :  a  cone-scale;  FIG.  328.— Diairrams  of  dichnsia  in 

b  fruit.  the  <J  -(A)  a  ml  ?-(#)  catkins  of  Birch. 

Order  2.  Corylaceae  (Hazel-nuts).  Monoecious.  The  <£- 
catkins  are  long  and  cylindrical ;  the  $ -flowers  are  placed  singly 
in  the  axil  of  the  subtending-leaf  (cover-scale)  ;  they  are  naked 
and  formed  of  a  number  of  divided  stamens,  which  are  partly 
united  with  the  cover-scale,  4  in  the  Hazel,  apparently  8  (Figs. 
330  A,  329  B,  0),  more  on  the  Hornbeam.  The  ?  -flowers  have 
a  very  small,  superior  perianth ;  in  the  axil  of  each  cover-scale  a 
2-flowered  dichasium  (Fig.  329  D)  is  present,  of  which  the 
central  flower  (*  in  Fig.  330  B)  is  suppressed.  The  gynceceum  is 
bicarpellary  as  in  the  Birches  ;  the  ovary  is  bilocular,  with  two 
long  styles  (Fig.  329  D-F)  •  the  loculi  have  1  (-2)  ovules  (Fig. 
3305).  Each  single  ?  -flower  and  fruit  is  surrounded  by  a  leaf- 
like  covering,  the  cupule  (husk),  which  is  formed  of  three  floral-leaves 
(namely,  the  bract  of  a  lateral  flower,  and  its  own  bracteoles  ; 
thus  in  Fig.  330  .35,  a,  a',  ft  form  the  cupule  for  the  left-hand 
flower,  and  /?,  a,,  (3n  the  cupule  for  the  right-hand). 

Corylus  (Hazel-nut,  Fig.  329).  The  long,  cylindrical  (£ -catkins 
pass  the  winter  naked,  2-3  together,  on  short  branches.  The  very 


344 


DICOTYLEDONES. 


small  ?  -catkins  are  enclosed  in  buds,  in  which  they  pass  the 
winter ;  these  buds  are  situated  in  the  axils  of  the  fallen  foliage- 
leaves,  and  it  is  only  by  their  larger  size  that  they  may  be  dis- 
tinguished from  the  ordinary  foliage-buds.  In  spring  the  $  -catkins 
are  easily  recognised  by  their  red,  projecting  stigmas  (Fig.  329  A). 
The  cupule — the  "husk" — is  tubular, fringed,  and  envelopes  the  nut. 
The  leaves  are  alternate  and  unsymtuetrical,  the  external  side  being  larger 
than  the  inttrnal ;  this  is  connected  with  the  vernation,  the  blade  being  con- 


FIG.  329.— Corylusavellana:  A  branch  at  the  time  of  flowering  with  cJ-aud  ? -catkins; 
B  (J -flower  with  subtending-leaf  (bract)  and  two  bracteoles ;  C  the  same  without  t'-e 
anthers;  D  view  of  interior  of  ?-dichasium  shortly  after  fertilisation;  E  young  fruit 
with  cupule;  f  similar  one  with  the  cupule  opened ;  G  mature  ? -fruits;  H  nut. 

duplicate  in  the  bud  ;  the  stipules  are  deciduous  The  bud- scales  are  formed 
of  stipules,  the  most  internal  having  a  leaf -blade  attached  to  them  which  is 
suppressed  in  the  external  ones.  The  cotyledons  remain  underground  on  germi- 
nation. 

Carpinus(0.  betulus,  Hornbeam).     The  <$  -  and  $ -catkins  do  not 
appear   till    the    leaves   are    shooting.      The     ?  -catkin    in    this 


QUERCIFLORJi).  345 

instance  is  also  long  and  cylindrical.  The  cupule  in  C.  betulus  is 
3-lobed,  and  to  a  slight  extent  only  embraces  the  base  of  the 
ribbed  nut  (Fig.  331)  ;  each  lobe  corresponds  to  a  floral-leaf. 
Whilst  the  carpels  are  placed  medianly  in  Corylus,  in  Carpinus,  on  the  other 
hand,  they  are  situated  transversely,  as  in  the  case  of  the  Betnlacece.  The 
lamina  of  the  leaf  is  not  conduplicate  in  the  bud,  but  flat,  and  folded  only  along 
the  lateral  veins,  which  are  also  indicated  in  the  form  of  the  fully-developed  leaf ; 
otherwise  the  vegetative  characters  are  essentially  the  same  as  in  the  Hazel. 
The  cotyledons  are  aerial. — Ostrya  resembles  the  Hornbeam,  but  the  cupule 
completely  envelopes  the  nut,  as  a  sac  open  at  the  apex  (Eur.,  N.  Am.,  Japan). 

N.  Am.,  Asia,  and  Europe ;  25  species.— Fossil  forms  in  the  Oligocene.  Wind- 
pollinated.  USES.  As  timber  (Carpinus  betulus)  and  firewood.  The  fruits  of 
C.  avellann  (ordinary  Hazel-nut),  C.  tubulosa  (Lambert's  nut)  and  C.  column 
(Turkish  Filbert)  are  edible. 


FIG.  330.— Dingrams  of  the  <J -flovrcr  (;!)  of  Con/lus  FIG.  331.— Nm.  of  the  Hornbeam 

and  the  dichasiumof  the  ?  -flowers  (£).  with  cupule. 

Order  3.  Cupuliferae.  Monoecious.  The  inflorescences  make 
their  appearance  with  the  leaves,  arising  in  the  axils  of  the  leaves 
of  the  same  year.  A  woody  cupule  furnished  externally  with  scales 
or  spines  is  common,  and  surrounds  1-several  flowers  (the  cupule 
in  the  Corylacese  never  encloses  more  than  a  single  flower  or  fruit). 
The  ^-flower  has  a  united  perianth,  which  is,  however,  4-6  partite, 
and  encloses  an  indefinite  number  of  undivided  stamens.  The  ?  - 
flower  has  a  superior,  6-merous  perianth  (3  +  3,  compare  Figs. 
332  D,  334)  ;  the  gynceceum  is  formed  of  3  (or  in  Castanea  4-6) 
carpels  with  a  corresponding  number  of  stigmas  (Figs.  332  D, 
H ;  334,  335)  ;  and  the  ovary  has  at  the  base  3  (-6)  loculi  (Fig. 
333),  each  of  which  has  2  pendulous  anatropous  ovules  ;  the  fruit 
is  a  one-seeded  nut  (Figs.  332  E,  336). 

The  cupule  of  the  Cupulifera3,  according  to  the  opinion  of 
Eichler,  is  formed  by  united  bracteoles  (compare  Fig.  333,  where 


346 


DICOTYLEDONES. 


the  four  valves  in  the  cupule  of  Castanea  are  considered  as 
bracteoles  of  the  lateral  flowers  of  the  dichasium)  ;  according  to 
another  view  (see  Prantl,  in  Engler's  Bot.  Jahrb.,  viii.,  1889),  it  is 
a  ring-like  axial  outgrowth  independent  of  the  bracteoles  of  the 
flower,  whose  scales  and  spines  are  floral-leaves.  The  cupule  in 
the  Oak  only  encloses  the  base  of  the  fruit,  but  in  the  Eating- 
chestnut  and  Beech  the  fruit  is  completely  enclosed,  and  con- 


FIG.  332.— Castanea  vnaca :  A  branch  with  inflorescences;  B  <J. flower;  C  young  cnpule 
with  three  ? -flowers;  D  ? -flower;  E  the  same  in  longitudinal  section;  F  cupule  with 
3  nuts  (diminished) ;  G,  H  nuts  (£  in  longitudinal  section  to  show  embryo). 

sequently  the  cupule  must  divide  into  a  number  of  valves  (gener- 
ally 4)  to  allow  the  fruit  to  escape.  In  the  3-flowered  dichasia 
of  Pasauia,  Sect.  Eupasania  (Trop.  Ind.),  each  individual  flower 
has  its  own  cupule  of  the  same  structure  and  development  as  in 
Quercus,  and,  moreover,  each  group  of  flowers  has  externally  the 
typical  six  bracteoles. 

Castanea  (Eating-chestnut,  Fig.  332).      The  catkins  are   erect 


QUERCIFLOKJE.  347 

(A),  cylindrical,  with  the  ?  at  the  base  and  the  $  at  the  top,"  or 
some  are  entirely  £  and  composed  of  small  dichasia.  The  cupule 
((7,  F)  is  4<-valved,  provided  with  spines,  and  entirely  envelops  the 
3  nuts  ;  it  is  already  developed  at  the  time  of  flowering. —  <J -flowers 
are  most  frequently  borne  in  7-flowered  dichasia,  and  have  a  well  developed 
perianth,  most  frequently  consisting  of  6  leaves  in  two  whorls  (Fig.  332  B),  and 
a  large  number  of  stamens.  $  -flowers  are  most  frequently  borne  in  3-flowered 
dichasia  (Figs.  332  C,  333) ;  the  letters  in  Fig.  333  indicate  the  older  theory, 
according  to  which  the  4  bracteoles  (a'-ft)  of  the  two  lateral  flowers  are  thick 
and  united  into  a  single  4-valved,  woody  cupule,  which  surrounds  the  3  nuts, 
and  is  furnished  externally  with  spines ;  the  spines  are  well  developed  hair- 
structures. — 6  carpels  in  two  whorls. — The  leaves  in  the  vertical  shoots  have 
a  divergence  of  f ,  f ,  T5^ ;  on  the  horizontal  shoots  they  are  alternate.  The 
cotyledons  remain  underground  on  germination. 

Fagus  (Beech).  The  <£  -catkins  are  pendulous,  capitate ;   the  <$  - 
flowers  have  an  obliquely  bell-shaped,  fringed  perianth,  with  6-20 


FIG.  333.— Diagram  of  the  cupule  of  Castanea.         FIG.  33*.— Female  flower  of  Fagus  (mag.) 

stamens.  ?  -catkins  erect,  2-flowered,  borne  singly  in  the  axil  of 
foliage-leaves  of  the  same  year;  the  $ -flower  has  a  gyriceceum 
formed  of  3  carpels,  bearing  an  epigynous,  6-leaved  perianth  (Fig. 
334).  In  this  genus  the  dichasium  has  only  2  flowers,  the  central 
one  being  suppressed.  The  cupule  contains,  therefore,  only  2  tri- 
angular nuts  ("  mast  ").  All  the  shoots  have  the  leaves  arranged  in  two 
rows;  the  rows  are  on  the  underside,  being  only  about  90°  distant  from  each 
other  ;  the  buds  on  the  other  hand  approach  each  other  towards  the  upper  side. 
The  bud-scales  are  stipules  without  laminae  ;  in  vernation  the  laminae  are  folded 
-along  the  lateral  ribs,  the  upper  lateral  portion  being  the  largest  (as  in  Horn- 
beam and  Chestnut).  The  cotyledons  are  folded,  and  at  germination  are 
aerial,  large,  and  reniform.  4  species  (Europe,  Japan,  N.  Am.) — Notliofagus 
(S.  Am.,  New  Zealand,  S.  Austr.) 

Quercus  (Oak,  Fig.  335).  Catkins  simple.  $  -catkins  long, 
thin,  pendulous,  few-flowered.  $  -catkins  erect ;  the  cupule  is  cup- 
like,  entire,  and  encloses  only  the  base  of  the  solitary  nut  ("  acorn  "). 
—The  <?-  flower  has  a  similar  construction  to  that  of  the  Chestnut.  The- 


348 


DICOTYLEDONES, 


?  -catkin  has  not  more  than  5  flowers  (single-flowered  dichasia,  in  which  only 
the  central  floicer  is  developed}.  The  scales  on  the  cupules  are  no  doubt  leaf  - 
structures  in  this  case  also.  According  to  another  theory,  the  scales  are  hair- 
structures  ;  they  arise  on  the  internal  face  of  the  young  cupule  apparently  in 
descending,  but  really  in  ascending  order.  The  rim  of  the  cupule  gradually 
expands.  In  the  ?  -flower  (Fig.  335)  the  loculi  of  the  gynceceuai,  together  with 
the  ovules,  are  not  developed  until  after  pollination. — The  leaves  in  all  cases 
have  a  divergence  of  £ ;  the  lowermost  leaves  on  the  shoots  are  reduced  to 
stipules  which  serve  as  the  bud-scales  (5  rows).  The  laminae  are  conduplicate, 
as  in  Corylus,  and  the  external  side  is  the  broadest.  The  cotyledons  are  fleshy 
and  remain  underground.  .  200  species. — Pasania  (100  species). 

368  species,  in  temperate  climates,  especially  in  Europe  and  N.  America. 
Authenticated  forests  have  been  found  in  the  Oligocene.  The  Beech  has  one 
species,  Fagus  sylvatica,  in  Europe  ;  it  is  a  most  important  forest  tree  (in 
Denmark  the  most  important)  and  reaches  its  most  northern  limit  near  Alve- 
sund  in  Norway  (60°  N.L.),  its  northern  boundary  line  passing  from  Alvesund 
in  a  zig-zag  line  through  Ludwigsort,  south  of  Kouigsberg,  in  Prussia,  to- 
wards the  Crimea.  According  to  Steeustrup  and  Vaupell,  the  Beech  did  not 


B 


FIG.  335.— Quercus :  A  g  -flower  in  its  cupule 
(mag.) ;  B  longitudinal  section  through  A, 
showing  cupule,  perianth,  and  inferior  ovary. 


FJG.  330.— Fruit  of  Quercus. 


make  its  appearance  in  Denmark  until  a  comparatively  recent  time,  the  Oak 
then  being  partially  supplanted.  Other  species  of  Beech  are  found  in  N. 
America  and  Japan.  Several  species  of  Nothofagus  occur  in  the  South  West 
of  S.  America,  and  in  the  colder  regions  of  the  southern  hemisphere.  The 
Oaks  grow  especially  in  temperate  regions,  e.g.  in  Western  Asia,  N.  America, 
and  the  mountains  of  Mexico.  Evergreen  species  are  found  in  Tropical  Asia, 
Himalaya,  Japan  and  the  Mediterranean  region.  In  this  country  there  is  one 
species  of  Oak  (Q.  robur),  of  which  there  are  three  varieties  (Q.  peduncnlaia, 
intermedia,  sessili  flora).  The  Eating-chestnut  is  found  in  the  South  of  Europe, 
but  is  cultivated  in  the  midland  and  southern  counties  of  England. — USES. 
The  wood  of  these  trees  is  very  useful  as  timber.  The  wood  of  Q.  tinctoria. 
has  a  yellow  colouring  matter  (Quercitron-wood).  The  bark  of  the  Oak  contains 
a  large  quantity  of  tannic  acid,  and  is  used  for  tanning ;  for  this  purpose 
also  the  cupules  of  Q.  vallonea,  cegilops,  grceca,  and  others  from  the  Eastern 
Mediterranean,  are  used  under  the  name  of  "  Valloons."  The  Cork-oak 
•(Q.  suber ;  S.W.  Europe)  is  the  most  important  tree  from  which  cork  is  obtained, 


JUGIANDIFLOR^. 


349 


its  bark  being  very  largely  developed  and  stripped  for  cork.  Gall-nuts  are  found 
on  many  species  ;  those  of  Q.  lusitanica,  var,  infectoria  (Eastern  Mediterranean) 
are  officinal,  and  likewise  the  fruits  (acorns)  and  the  bark  of  Quercus  pedunca- 
lata  and  sessiliflora.  Oil  is  obtained  from  the  Beech  "  mast."  The  nuts  of  the 
Chestnut  tree  are  edible. 

Family  4.    Juglandiflorse. 

This  family  resembles  the  Querciflorae  in  the  catkin-like  inflores- 
cences, the  imperfect,  unisexual  flowers,  the  epigynous  perianth 
and  the  woody  shoots  with  scattered  leaves,  etc.,  though  it  is  in 
other  respects  very  dissimilar ;  one  point  of  difference  is  the  presence 
of  aromatic  compounds,  but  a  more  imporant  divergence  is  found 
in  the  structure  of  the  gynoeceum,  which  is  formed  of  two  carpels 
with  one  loculus  and  has  one  basal,  orthotropous  and  erect  ovule, 


Fio.  337. — Juglans  regia:  A  £ -flower  seen  from  below  with  bract  (cover-scale)  (b), 
bracteoles  (a  and  /3),  perianth-leaves  (p)  ;  B  the  same  from  the  front ;  C  lateral  view  of  the 
same  ;  D  diagram  of  A  ;  E  $ -flower  with  bract,  the  bracteoles  are  united  with  the  ovary, 
their  edge  being  visible  as  an  indented  line  below  the  perianth  ;  F  2  ? -flowers  at  the  end  of 
a  foliage-shoot;  G  fruit  (without  the  fleshy  covering,)  in  longitudinal  section;  H  trans- 
verse section  of  the  same.  % 

which,  as  in  the  Quercifloree,  does  not  become  developed  until 
after  pollination ;  the  fruit  too  is  very  different,  being  generally  a 
drupe.  Endosperm  absent. 

Order  1.  Juglandaceae  (Walnuts).  Leaves  scattered,  impari- 
pinnate,  rich  in  aromatic  compounds.  Stipules  absent.  Flowers  uni- 
sexual. Monoecious.  The  ^-catkins  are  lateral,  generally  on  naked 
branches  of  the  previous  year,  cylindrical,  pendulous,  many-flowered; 
the  two  bracteoles  and  the  2-4-leaved  perianth  of  the  $  -flower  unite 
with  the  subtending  bract ;  the  £  -flower  has  indefinite  stamens 
.  (6-20  in  Juglans,  Fig.  337  A-D).  The  $  -catkins  are  terminal,  gene- 
rally on  branches  of  the  same  year,  few-flowered  (Fig.  337  F)  ;  the 


350  DICOTYLEDONES. 

$  -flowers  have  a  superior,  4-leaved  perianth,  a  bicarpellate  gynoe- 
ceum,  two  styles  with  stigmas  on  the  internal  surface.  The  ovary, 
bracteoles  and  bract  all  unite  together  (Fig.  337  E).  The  fruit  is 
generally  a  green  or  black  drupe,1  whose  flesh  (outer  soft  portion)  in 
Carya  and  Juglans  ruptures  more  or  less  irregularly,  and  frees  the 
stone  ("Walnut"). — The  stone  in  Juglans  is  divided  internally  by  one  true 
(Fig.  337  H)  and  by  several  false,  low  partition  walls  into  several  incomplete  com- 
partments, so  that  the  two  large  cotyledons  become  lobed  and  incised  to  fit  like 
a  cast  into  the  irregularities  of  the  inner  surface  of  the  stone ;  the  embryo  is 
exendospermous  and  covered  with  a  thin  testa. — THE  LEAP  SCARS  are  large  and 
cordate  with  3  groups  of  vascular  bundles.  The  PITH  in  Juglans  and  Pterocarya 
is  divided  into  chambers.  The  stone  ruptures,  on  germination,  along  the 
dorsal  suture  into  2  valves';  the  cotyledons  remain  underground.  In  Juglana 
regia  a  long  row  of  accessory  buds  is  found  on  the  lowest  internode  (epicotyl) 
above  the  axils  of  the  cotyledons.  Pollination  by  the  wind.  Both  protogynous 
and  protandrous  examples  of  Juglans  regia  occur. — 33  species,  mostly  in  tem- 
perate North  America. — USES.  Walnuts  are  obtained  from  J.  nigra  and  regia  ; 
Hickory  from  North  American  species  of  Carya.  The  oil- containing  seeds  of 
several  species  are  edible.  Pterocarya  and  others  are  cultivated  as  ornamental 
plants. 


FIG.  338.— Myrica  gale  :  a  young  fruit ;   x  the  bracteoles  with  numerous  glands  ;  b  longi- 
tudinal  section  of  fruit. 

Order  2.  Myricacese.  To  this  order  belong  shrubs  or  trees 
which  have  penninerved,  simple,  at  most  lobed  or  pirmatifid  leaves, 
with  or  without  stipules,  and  with  yellow,  aromatic,  resin  glands 
(Fig.  338  a).  The  flowers,  situated  in  catkin-like  spikes,  are- 
unisexual  and  naked,  and  supported  by  scale-like  floral-leaves. 
£  -flower  :  4-6  (-16)  stamens  with  short  filaments;  $  :  generally 
situated  singly.  The  gynoeceum  has  a  short  style  with  2  long 
stigmas,  and  unites  with  the  bracteoles,  which  form  wing-like 
outgrowths  on  the  ripe  drupe  as  in  Pterocarya,  in  the  Juglandaceae 
(Fig.  338).  Cotyledons  fleshy  (Fig.  338  fy.—Myrica;  Comptonia. 

1  The  fruit  of  the  Walnut  is  thus  a  false  fruit ;  and  the  term  drupe  must 
therefore  not  be  used  in  the  same  sense  as  in  the  Bosacese. 


URTICIFLOR^E.  351 

40  species  ;  Temperate. — Myrica  gale  (Sweet-gale,  Bog-myrtle)  has  been  used 
in  the  preparation  of  beer  (Sweet-willow  beer)  on  account  of  its  resinous  essential 
oil.  M.  cerifera  (N.  America)  and  species  from  the  Cape,  M.  quercifolia  and 
others,  form  wax  on  the  fruit  which  is  used  in  the  preparation  of  candles. 

Family  5.     Urticiflorae. 

The  flowers  are  regular,  hypogynous,  nearly  always  unisexual, 
small  and  insignificant,  with  single,  green  perianth  of  4-5  leaves. 
Stamens  4-5,  placed  opposite  the  leaves  of  the  perianth.  Ovary 
formed  of  1  or  2  carpels,  most  frequently  unilocular,  with  one 
ovule  (Fig.  340).  The  fruit  isanut,  more  rarely  a  drupe,  with  one 
seed,  generally  endospermous.  The  Nettles  are  the  sole  order  in  the 
family  which  has  only  one  carpel  (1  stigma) ;  this  turns  the  posterior  side  to 
the  front  (Fig.  340).  The  others  have  two  carpels  (2  stigmas)  but  the  anterior 
only  is  fertile  (Fig.  346)  except  in  a  few  Ulmaceaa  and  Moraceae. 

The  majority  arj  trees  or  shrubs  with  petiolated  leaves,  stipu- 
late ;  rough  hairs  are  very  frequently  developed  upon  the  leaves 
The  flowers  are  very  often  crowded  together  in  the  inflorescence, 
which  is  rarely  catkin-like.  Peculiar  aggregations  of  fruits  are 
found  in  some  orders.  Latex  and  tough  bast,  which  is  used  techni- 
cally, are  also  frequently  found.  Cystoliths  are  found  in  the 
epidermis  of  many  species  of  Ficus,  Urtica,  and  others.  Wind-  or 
self-pollination  is  most  common,  as  in  the  Querciflorae  and  JuglandifloraB.  In 
the  Urticaceae,  Murus  and  some  others,  the  stamens  lie  incurved  in  the  bud, 
and  when  ripe  straighten  themselves  suddenly  and  elastically,  and  thus  small 
clouds  of  pollen-grains  are  ejected  with  considerable  violence  on  to  the  stigmas, 
which  are  often  provided  with  brush-like  hairs  (Fig.  341).  The  formation  of 
honey  does  not  take  place. 

Order  1.  Ulmaceae  (Elms). — Trees  or  shrubs  without  latex. 
Leaves  simple,  arranged  in  two  rows  (divergence  |-),  oblique  (the 
inner  side,  nearer  the  axis,  being  the  larger),  strongly  penni- 
nerved,  dentate,  hispid;  stipules  deciduous.  In  opposition  to  the 
other  Nettle-like  plants  the  flowers  are  often  £  with  a  united 
cup-  or  saucer-like,  generally  4— (5)-6-divided  perianth,  and  a 
corresponding  or  larger  number  of  opposite  erect  stamens.  The 
gynceceum  has  two  carpels  (2  stigmas),  generally  one  loculus  with 
one  pendulous,  anatropous  or  amphitropous  ovule,1  seldom  two 
loculi  and  2  ovules.  Fruit  one-seeded  (nut  or  drupe).  Embryo- 
without  endosperm. 

A.  TJiMVJE.  The  fruit  is  a  winged  nut  (Fig.  339),  the  embryo 
straight,  without  endosperm.  Anthers  extrorse. — Ulmus  (Elm). 

1  The  pollen-tube  in  Ulmus  does  not  enter  the  ovule  through  the  micropyle. 
W.  B.  A  A 


352 


DICOTYLEDONES. 


The  flowers  are  situated  in  inflorescences  which  develop  from 
the  lower  buds  of  the  shoot  of  the  preceding  year.  The  lowermost 
bud-scales  are  empty,  the  uppermost  support  either  solitary  flowers,  or  small, 
dichasial  or  unipared  scorpioid  inflorescences.  The  terminal  bud  on  the  vege- 
tative shoot  quickly  falls  off,  and  the 
upper  lateral  bud  continues  the  growth 
sympodially.  Flowering  takes  place 
before  the  leaf-buds  open.  The  flowers 
are  wind-pollinated  and  Lave  no  honey. 
Fossil  species  have  been  found  in  the 
Oligocene. 

20  species  ;  North  Temp.  (2  species 
in  this  country).  Important  as  timber. 
The  Cork-elm  (U.  suberosa)  has  a  rather 
thick  cork,  which,  however,  is  of  no 
technical  use.  The  bast  is  used  as 
Lime-bast. 

B.  CELTIDEJE.  The  fruit  is  a  drupe, 
the  embryo  curved,  with  folded  or  rolled 
up  cotyledons,  with  or  without  endo- 
sperm. The  anthers  are  introrse.  The 
flowers  are  borne  on  a  shoot  of  the 
same  year.  Planera  (N.  America) ; 
Zelkova. — About  114  species ;  especially 
N.  Temp.,  Trop. 


FIG.  339.— -A  Ulmus  campcstris,  flower 
with  exceptionally  aborted  gynoeceum; 
B,  U.  effuxa,  flower  with  8  stamens  ;  C,  IT. 
campestris,  fruit  opened  in  front  to  show 
the  seed  pendulous  from  the  apex  of  the 
loculus;  one  loculns  is  aborted. 


Order  2.  Urticacese 
(Nettles).— The  majority  of  spe- 
cies are  herbs  with  simple,  stipu- 
late leaves  ;  they  have  no  latex ; 
stinging  hairs  abundant.  The  flowers  (Fig.  340)  are  unisexual,  gene- 
rally 2-merous  and  arranged  in  clusters,  which  are  united  into 
catkin-like  inflorescences.  The  perianth  is  composed  very  often  of 
4  (2  +  2)  free,  or  in  the  ?  -flowers  generally  united,  green  leaves ; 
the  4  (2  +  2)  stamens  are  opposite  the  perianth-leaves,  the  filaments 
are  bent  inwards  in  the  bud  and  throw  themselves  elastically 
towards  the  outside.  The  gynceceum  has  one  style  and  i,ne  stigma 
(capitate  or  brush-like,  Fig.  341)  ;  the  ovary  is  unilocular,  with  one 
orthotropous,  erect  ovule  (all  other  orders  of  this  family  have  inverted 
or  curved  ovules).  Fruit,  a  nut  or  drupe.  Endosperm  present 
(in  JJrtica  very  little),  oily.  Embryo  straight.  The  STINGING  HAIRS 
are  club-shaped,  very  turgid,  and  provided  with  a  siliceous,  brittle  apex, 
which  breaks  off  in  an  oblique  direction  and  allows  the  poisonous  cell- sap  to 
be  forced  out.  In  many  tropical  Nettles  this  is  so  strong  that  it  may  produce 
partial  paralysis.  There  is  no  rudiment  of  an  ovary  in  the  <£  -flowers  (Fig.  340 
A).  The  PERIANTH  in  the  ?  -flower  differs  from  that  of  the  <?  in  having  the  two 


URTLCIFLOR51.  353 

internal  leaves  generally  much  larger  and  enveloping  the  fruit  (340  B)  ;  it 
often  happens  that  all  the  perianth-leaves  are  united  to  form  a  gamophyllous 
envelope.  £  -flowers  may  occur  among  the  others. — THE  INFLORESCENCES  among 
our  native  species  are  dicliasia,  which  become  transformed  into  unilateral 
scorpioid  cymes  by  the'  development  ot  the  bud  of  the  2nd  bracteole.  In 
Parietaria  they  are  more  pressed  together,  and  the  floral-leaves  at  the  same 
time  are  also  raised  on  their  axillary  shoots  to  just  beneath  the  flower.  As  a 
rule,  not  only  in  this  order  but  also  in  those  related  to  it,  a  small  vegetative 
branch  is  situated  in  the  axil  of  the  foliage-leaf,  and  this  bears  an  inflorescence 
on  each  side  at  its  base. 

Urtica  (Nettle)  has  opposite  leaves  with  distinct  stipules  and 
stinging  hairs.  The  perianth-leaves  of  the  ?  -flower  are  free 
(Fig.  340). — Parietaria  (Pellitory)  has  scattered  leaves  without 
large  stipules,  and  stinging  hairs  are  absent.  The  $  -perianth  is 
4-toothed,  flask-  or  bell-shaped. — Film  is  a  tropical  genus  with  trimer- 
ous,  zygomorphic  £  -flowers,  the  posterior  perianth-leaf  being  much  larger  than 


FIG.  340.— Diagram  •of    <f-  and  ?•  FIG.  341. — Parietaria dijitsa;  hermaphrodite 

•flowers  of  Urtica  dioica.  flower  :  a  in  the  female,  b  at  the  commence- 

ment of  the  male  stage;  the'stigma  has  fallen 
off,  but  the  anthers  have  not  yet  dehisced. 

the  two  others,  and  more  or  less  hood  shaped. — The  flower  of  Forskohlea  is  the 
most  reduced;  the  <?  -flower  has  only  one  stamen,  and  the  ?  -  as  well  as  the  <?  - 
flowers  have  a  one-sided,  tongue-like  perianth  (?).  Pouzolzia. 

WIND -POLLINATED.  The  pollen  is  shot  out  of  the  anthers,  when  they  spring 
forward,  and  is  caught  by  long  stigmatic  hairs.  Parietaria  diffusa  is  protogy- 
nous  (Fig.  341). 

500  species  ;  chiefly  in  the  Tropics,  although  the  few  species  which  occur 
in  Europe  are  represented  by  a  much  larger  number  of  individuals. — USES. 
The  bast  of  the  native  species  Urtica  dioica  and  urens,  of  U.  cannabina 
(Siberia),  etc.;  of  Boehmeria  nivea  "Ramie"  and  "China-grass"  (from  Sunda 
Is.,  China),  and  others,  is  used  in  the  manufacture  of  muslin. 

Order  3.  Moracese,  (Mulberries).  Nearly  all  trees  or 
shrubs,  seldom  herbs,  generally  with  latex.  The  leaves  are  scat- 
tered, and  not  infrequently  lobed.  The  flowers  are  unisexual 
(monoecious  or  dioecious)  and  arranged  in  catkin-  or  capitulum-like, 
compound  inflorescences.  Perianth-leaves  2-6,  generally  4,  with  an 
-equal  number  of  stamens  opposite  to  them,  as  in  the  .Nettles.  The 


354  DICOTYLEDONES. 

ovary  is  1-  seldom  2-locular,  and  has  2  stigmas  (it  is  thus  formed 
from  2  carpels)  seldom  only  one  style  with  one  stigma.  One 
ovule  in  each  loculus,  more  or  less  curved,  and  pendulous ;  micro- 
pyle  directed  upwards.  Fruit  usually  a  drupe.  The  embryo  is 
generally  curved  inside  i\\Q  fleshy  endosperm,  or  it  is  exendospermous. 
A.  MORE.E.  The  filaments  are  incurved  in  the  bud.  Leaves 
folded  in  the  bud. — Morus  (Mulberry)  (Figs.  342-344).  Monoecious. 
The  inflorescences  are  catkin-like  in  appearance,  but  in  reality 
composed  of  many  small  dichasia.  The  flowers  are  similar  to  those 
of  the  Nettle,  but  with  2  carpels :  in  the  $  with  perianth  2  +  2, 
and  stamens  2  +  2  (Fig.  342),  in  the  ?  ,  perianth  2  +  2,  and  2 
carpels  in  regular  alternation.  The  small  drupes  are  enveloped 
by  the  perianth,  which  eventually  becomes  fleshy,  and  as  all  the 
flowers  on  the  axis  very  accurately  fit  together,  the  collection  of 


FIG.  342.— Mortis  alba  FIG.  343.— Morus  alba  FIG.  344.—  Morus  nigrn 

$  flower  (6/i;.  ?  inflorescence.  fruits. 

fruits  is  formed,  which  we  call  a  Mulberry  (Fig.  344) .  The  leaves 
are  folded  in  the  buds,  and  have  small  stipules.  The  following  are 
allied  to  Morus  : — Madura,  Broussonetia  (the  Paper-mulberry  tree)  which  has 
spheroid  $  inflorescences  (made  up  of  dichasia),  etc. 

Dorstenia  presents  an  interesting  transitional  form  to  the  Fig  in  its  flat, 
open,  and,  in  some  instances,  lobed  inflorescence  on  which  the  <J  and  ? 
flowers  are  sunk  in  grooves.  Indications  of  a  somewhat  similar  structure  are 
found  in  certain  Nettles,  the  sympodial  axes  of  the  dichasia  becoming  flatly 
expanded.  The  fruits  are  1-seeded,  but,  nevertheless,  spring  open  and  eject 
their  seeds. 

B.  ARTOCARPE^;.  Filaments  straight  in  the  bud;  foliage- 
leaves  with  convolute  vernation.  An  interpetiolar  leaf-sheath 
(ocrea)  formed  in  the  axil  of  each  leaf  by  the  connate  stipules, 
covers  the  younger  leaves  as  a  hood.  It  falls  off  as  the  leaf 
expands,  and  leaves  a  ring-like  scar  on  the  stem. — Ficus  (the 
Fig).  The  inflorescence  (the  so-called  syconus)  has  a  pear- 


URTICIFLOR.E. 


355 


shaped,  fleshy,  but  hollow  axis,  on  the  interior  surface  of  which  the 
flowers  are  situated  (Fig.  345).  It  is  a  kind  of  capital  urn,  with 
a  hollow  receptacle,  whose  "  involucral "  leaves  close  over  the 
entrance  to  the  interior ;  it  is  not,  however,  a  simple  capitulum, 
but  a  coalescence  of  cymose  inflorescences.  The  edible  parts  are 
the  fleshy  stem-portion  and  perianth-leaves.  The  <$  -flower 
has  a  2-6  divided  perianth,  1-2  (-6)  stamens  ;  the  $  -flower 
has  an  oblique  ovary.  The  fruits  are  drupes,  with  thin  flesh. 
— Many  species  have  aerial  roots,  and  some  live  as  epiphytes  oa  trees. 
POLLINATION,  in  the  edible  Fig,  is  effected  by  a  small  Gall- wasp  (Cynips 
psenes  L.),  which  lays  its  eggs  in  the  Fig,  and  hence  carries  the  pollen  away. 
Even  in  very  ancient  times  it  was 
customary  to  hang  infected  wild  Figs 
on  the  branches  of  cultivated  ones,  so 
that  the  young  Gall-wasps,  as  they 
emerged,  could  immediately  effect 
the  pollination  (caprification).  Ficus 
carica,  and  other  species,  have  two  kinds 
of  ?  -flowers,  besides  the  <?  -flowers. 
One  kind  has  a  short  style  and  no  stig- 
matic  hairs,  and  it  is  only  in  the  ovaries 
of  these  that  the  wasps  lay  their  eggs 
(gall-flowers)  ;  the  other  kind  has  a  long 
style  and  well-developed  stigmatic-hairs, 
but  the  wasps  cannot  reach  their  ovaries 
— these  are  "seed-flowers."  There  are, 
moreover,  two  kinds  of  plants  of  Ficus 
carica  ;  £  -plants,  which  have  only  seed- 
flowers,  and  bear  the  edible  Figs,  and 
<£  -plants  (called  "Caprificus"),  which 
bear  inedible  fruits,  and  have  $  -flowers 
at  the  upper  part  of  the  Fig,  but  gall- 

flowers  at  the  base.  [The  Caprificus,  at  r'6'  3*5-A  Fi* in  l<"#tadin.l  section. 
Naples,  bears  three  crops  of  inedible  Figs  each  year,  viz.  Mamme  (April),  Pro- 
ftchi  (June),  Mammoni  (August).  The  $  -flowers  are  produced  especially  in  June, 
the  first  Figs  being  almost  entirely  $  ,  and  the  last  having  but  few  $  -flowers. 
Each  crop  produces  a  new  generation  of  Fig- wasps.  The  female  wasp  enters 
the  Figs  on  the  Caprificus,  and  lays  one  egg  in  each  flower,  with  the  result  that 
the  flower  developes  into  a  kind  of  gall.  The  mother- wasp  dies  within  the  Fig. 
The  male  wasp  is  wingless  ;  it  bites  a  small  passage  into  the  ovaries  containing 
the  female  wasps,  and  impregnates  them ;  the  female  wasps  then  escape  from 
the  Fig,  those  in  the  Profichi  carrying  pollen  away  with  them  as  they  pass  out. 
They  then  enter  another  Fig,  lay  their  eggs,  and  die.  The  edible  Fig-tree 
similarly  has  three  crops  in  the  year,  Fiori  di  Jico,  Pedagnuoli,  Cimaruoli. 
The  wasps,  entering  these  Figs,  are  unable  to  lay  their  eggs  in  the  ovary,  but, 
nevertheless,  they  effect  cross-pollination  on  entering  the  Pedagnuoli,  which 
bear  fertile  seeds.] 


356  D1COTYLEDONES. 

The  flowers  of  lirosimum  are  the  most  reduced.  The  perianth  is  -wanting, 
and  the  $  -flower  has  only  1  stamen.  Cecropia  (Trumpet-tree),  in  S.  Am.,  has 
its  pith  divided  into  chambers ;  these  are  inhabited  by  ants,  which  feed  upon 
small  food-bodies  formed  on  the  swollen  base  of  the  petioles.  The  leaves  are 
petiolated,  often  shield-like,  fringed  or  lobed,  and  sometimes  with  white  felted 
hairs.  They  serve  as  food  for  Bradypus  (the  Sloth).  Sorocea;  Castilloa. 

About  300  species  exclusively  in  the  warmer  climates.  The  white  Mulberry 
(M.  alba,  from  China,  India,  Mongolia)  is  cultivated  for  the  sake  of  its  leaves, 
which  are  the  indispensable  food  for  silkworms.  The  black  Mulberry  (M.  nigra, 
W.  Asia)  is  cultivated  for  its  fruits,  which  are  used  for  the  officinal  Mulberry 
juice.  The  ordinary  Fig-tree  (Hens  carica)  is  from  the  Mediterranean.  The 
fruit  of  the  well-known  Oriental  Sycamore  (F.  sycomorus)  is  edible.  The  Bread- 
fruit tree  (Artocarpus  incisa)  and  the  Jack  (A.  integrifolia)  have  their  home 
in  the  South  Sea  Islands,  and  are  cultivated  in  tropical  countries.  The 
Bread-fruit  is  morphologically  the  same  as  the  Mulberry.  It  has  a  very  large, 
spheroid  inflorescence,  whose  floral-leaves  and  perianth  become  fleshy  and 
united  into  one  nutritious  mass,  together  with  the  axis,  which  is  also  fleshy. 
The  milky  juice  of  the  India-rubber  tree  (Ficns  elastica,  East  Indies,  a  com- 
mon house-plant),  and  of  Castilloa  elastica  (Am.)  is  the  raw  material  of  India- 
rubber.  The  milky  juice  of  Galactodendron  utile,  (Cow-tree,  S.  Am.)  is  sac- 
charine and  nutritious,  but  in  Antiaris  toxicaria  (the  Upas-tree,  of  Java)  it 
is  a  strong  poison.  The  bast  of  the  Paper-Mulberry  tree  (Br.  papyrifera, 
Eastern  Asia)  is  used  in  Japan  for  paper.  Shellac  is  obtained  from  a  small, 
hemipterous  insect  (Coccus  lacca),  which  lives  upon  Ficus  laccifera  and  F. 
religiota  (the  Bo-tree,  sacred  to  Buddha),  E.  India.  The  wood  of  Madura 
aurantica  (Am.)  has  a  yellow  colour,  and  is  known  as  yellow  Brazilian  wood. 

Order  4.  Cannabaceae.  The  plants  which  belong  to  this 
order  are  aromatic  herbs,  either  annuals  or  perennials,  without 
latex.  Leaves  palminerved,  and  more  or  less  divided,  hispid,  and 
with  free,  persistent  stipules.  Flowers  always  dioecious-,  $  -flowers 
in  panicles,  formed  of  dichasia,  passing  over  into  uniparous  scor- 

pioid  cymes.  They  differ  from  the 
Nettles,  particularly  in  the  5-leaved 
perianth  of  the  ^-flower,  the  5  sta- 
mens  (Fig.  346-351)  with  filaments 
erect  in  tne  bud,  and  in  the  ?  -flower 
by  the  small,  entire,  cup-like  perianth, 
FIG.  3i6.-Diagram  of  male  and  which  surrounds  the  base  of  the  ovary 

female  flowers  of  the  Hop  and  (Fig.  346,  p.  352).  The  Ovary  has 
Hemp :  b  the  bract,  p  the  perianth.  .  .  ,  ,  . 

The  position  of  the  embryo  is  two  styles,  or  one  divided  into  two, 
indicated.  with  two  stigmas  and  a  pendulous, 

curved  ovule  (Fig.  346  JB,  352  B] ;  the  fruit  is  a  nut ;  the  embyro 
is  curved  (Hemp,  Fig.  353),  or  rolled  (Hop,  Fig.  349),  without 
endosperm. 


URTICIFLOM;. 


357 


Only  2  genera  with  3  species  (Asiatic),  of  which  two  are  culti- 
vated. —  Humulus  lupulus  (Hop,  Figs.  347-349)  is  a  twining, 
perennial  plant,  twisting  to  the  right,  with  opposite,  palmilobed, 
rough  leaves,  and  large,  interpetiolar  stipules.  The  ?  -flowers 
are  situated  in  closely-flowered,  cone-like,  compound  inflores- 
cences, with  ultimately  large,  thin,  imbricate  floral-leaves  (Fig. 
348)  which  bear  the  yellow,  glandular  hairs,  containing  lupuliiu 


317.  348. 

FIG.  347-348.—  Humulus  lupulus :  347,  twining  stem  ;  348,  branch  with  strobiles. 

This  inflorescence  is  made  up  as  follows  : — The  most  external  floral-leaves  are 
situated  in  pairs,  and  are  the  persistent  stipules  of  a  leaf,  the  blade  of  which 
has  become  suppressed,  or  in  any  case  is  rudimentary.  Such  a  pair  of  stipules 
supports  4  (2-6)  flowers  in  a  double  uniparous  cyme,  whose  central  axis  does 
not  develope  into  a  flower.  The  bracts  of  these  flowers  (bracteoles  of  the 
partial  inflorescence)  become,  at  maturity,  very  large,  spathe-like,  and,  together 
with  the  stipules,  produce  a  cone-like  appearance. 

Cannabis  sativa  (Hemp,  Figs.  350-353)  is  an  East  Indian  herb, 
with  palmilobed  leaves,  and  differs  from  the  Hop  in  being  annual, 


358 


DICOTYLEDONES. 


erect,  arid  in  having  its  leaves  opposite  at  the  base  and  scattered 
above.  The  ?  -inflorescence  is  not  cone-like  as  in  the  Hop,  but  the 
flowers  are  similar  in  construction.  The  main 
difference  is  to  be  found  in  the  axillary  shoot,  which  was 
suppressed  in  the  Hop,  and  is  in  the  Hemp  developed 
into  a  leaf-bearing  shoot  which  on  each  side  bears  only 
one  ?  -flower,  and  in  the  fact  that  the  bracts  are  not  so 
strongly  developed. 

The  "  Hops  "  (the  female  inflorescences)  are  used  in 
brewing,  and  medicinally  on  account  of  the  yellow 
glands  which  contain  lupulin.  The  Indian  variety  of 
Cannabis  sativa  contains  an  abundance  of  glandular 
hairs  and  resin.  The  withered  inflorescences  are  used 
in  medicine  and  are  officinal.  The  bast  of  the  stems 
of  the  Hemp  is  also  used  and  the  fat  oil  of  the  seeds.  In  Oriental  countries 
the  entire  plant  is  use  1  in  the  preparation  of  an  intoxicating  drink  (haschisch), 
the  narcotic  material  being  found  in  the  glandular  hairs. 


FIG.  349.  —  Humulus 
lupulus :  fruit  in  longi- 
tudinal section. 


350. 


352. 


353. 


FIGS.  350-354.— Cantialis  sativa:  350,  $  -plant;  351,  $  -flower;  352,  ? -flower,  entire  and 
in  longitudinal  section;  353,  fruit  in  longitudinal  section. 


Family  6.     Polygoniflorse. 

This  family  is  on  one  side  closely  allied  to  the  TTrticacece  by  its 
solitary,  basal,  vertical,  and  straight  ovule,  and  by  the  conical  ocrea 
which  envelopes  the  younger  leaves  in  the  bud,  similar  charac- 


POLYGONIFLORJE. 


359 


fcers  being  present  in  the  Urticacese.  On  the  other  side  it  is 
related  to  the  Curvembryge.  The  flowers  are  small,  often 
trimerous,  regular  and  slightly  perigynous  (in  Chloranthacece,  if  they 
properly  belong  to  this  family,  and  Houttuynia,  more  or  less  epigynous). 
Syncarps  are  present  in  some  Piperaceae,  but  the  fruit  is  generally 
a  single  fruit,  one-seeded  berry,  nut  or  drupe.  The  leaves  are 
generally  scattered. 

Order  1.  Polygonaceae.  The  majority  are  herbaceous  plants 
with  round,  often  jointed  stems,  scattered  leaves  and  ocrea,  that  is 
a  membranous,  tubular,  ligular  or  stipular  structure  inside  the 
base  of  the  leaf,  which  clasps  the  stem  and  axillary  bud  ;  the 
edges  of  the  lamina  are  rolled  backwards  in  the  bud.  The  flowers 
are  regular,  small,  generally  £ ,  slightly  perigynous,  with  incon- 
spicuous, simple,  green  or  white  perianth  of  5-6*  free  segments ; 
stamens  5-9  (Fig.  354)  sometimes 
arranged  in  two  series  ;  gynoaceum 
2-3  carpels,  ovary  unilocular  with 
one  basal,  straight  (orthotropous) 
ovule,  2-3  free  styles.  The  fruit  is  a 
2-3-angular  nut ;  the  embryo,  with 
mealy  endosperm,  is  straight  or 
curved  (Fig.  355  fl"),  often  unsym- 
metrical. — The  inflorescences  are  com- 
pound, and  generally  branch  from  the 
axils  of  the  bracteoles,  so  that  the  last 
partial-inflorescences  become  coiled,  uni- 
parous  scorpioid  cymes ;  in  Polygonum 
the  two  bracteoles  unite  into  a  mem- 
branous tube  ;  in  FJieum  and  Rumex  there 
is  only  one  bracteole. 


FIG.  354.— A  Diagram  of  Rheum  ;  E  of 
Rumex;  Cot  Polygonnm  J'agopyrum;  D 
of  P.  lapathifolium.  The  ovules  are  in- 
dicated inside  the  ovaries ;  bracts  and 
bracteoles  are  not  shown. 


Eheum  (Rhubarb,  Fig.  354  A)  has  a  6-leaved,  petaloid  perianth 
(Pn  3  +  3)  and  9  stamens  (A32 +  3).  The  3-winged  nut  is  not 
enclosed  by  the  perianth. 

Rumex  (Dock,  Fig.  354  B)  has  6  stamens  (A32  +  0);  the  perianth 
is  6-leaved  (Pr3  +  3),  green  or  red,  and  the  triangular  nut  is  en- 
veloped by  the  3  interior  perianth-leaves,  which  point  upwards 
and  continue  to  grow  after  flowering.  These  perianth-leaves  often 
have  warts  on  their  outer  surface.  The  following  are  monoecious :  R. 
acetosa  and  E.  acetosella. 

Polygonum  (Knot-grass,  Figs.  354  C,  D ;  355).  The  petaloid 
perianth  is  most  frequently  5-merous  (-f  spiral)  ;  5-8  stamens. 
The  nut  is  triangular  (Fig.  354  C,  355),  or  lenticular  (Fig.  354  D). 


360 


DICOTYLEDONES. 


There  are  two  whorls  of  stamens,  the  external  with  introrse,  and  the  internal 
with  extrorse  anthers.     The  gynoeceum  is  often  bi-carpellate  (Fig.  354  D). 

The  flowers  may  be  considered  as  constmcted  upon  the  monocotyledonous 
type.  Pterostegia  has  a  perfectly  monocotyledonous  flower  with  5  trimerous 
whorls.  Kheunt  likewise,  but  here  the  external  staminal  whorl  is  doubled  (Fig. 
254  A).  O&yria  has  a  dimerous  I?/«eim-flower  (4-leaved  perianth,  6  stamens. 
2  stigmas).  Eumex  has  a  J?/ie?tm-flower  with  the  suppression  of  the  internal 
whorl  of  stamens  (Fig.  354  B) ;  Emex  is  a  dimerous  Rumex.  Polygonum,  to 
which  Coccolob  -,  Miililfnbeckia  and  others  are  related,  differs  from  Rheum 


uiutra 


FIG.  355.— Polygonum  fagopymm :  A  branch  with  flower  and  Iruits  (nat.  size);  B  flower; 
C  the  same  in  longitudinal  section  ;  D  anterior  and  posterior  view  of  stamen  ;  E  gynoeceum ; 
F  fruit  (mag.) ;  G  fruit  in  longitudinal  section;  H  transverse  section,  showing  the  curved 
cotyledons  embedded  in  the  endosperm ;  I  the  embryo. 

chiefly  in  having  one  of  the  leaves,  which  in  the  latter  takes  part  in  the  forma- 
tion of  the  perianth,  developed  in  this  case  into  a  bracteole  (so  that  the  perianth 
is  reduced  to  rive  members),  and  several  or  all  the  stamens  in  the  inner  whorl 
become  suppressed. — The  perianth  in  Coccoloba  and  Miihlenbeckia  is  more  or 
less  perigyuous  and  becomes  fleshy,  enclosing  the  fruit.  Miihlenbeckia 
vlatyclada  has  flat  branches  with  rudimentary  leaves;  sometimes  branches 
with  normal,  arrow-shaped  leaves  are  found.  Atraphaxis. 


POLYGON1FLORJ3.  361 

POLLINATION.  Rumex  is  wind-pollinated,  the  stigmas  are  therefore  large  and 
bru-h-Hke  (indicated  in  Fig.  354  B).  Rheum  and  Polygonum  are  insect-pollinated 
and  have  therefore  capitate  stigmas,  etc.;  honey-glands  are  situated  at  the  base 
of  the  stamens  (d,  in  Fig.  354  C,  and  n  in  Fig.  356) ;  a  few  small-flowered 
Polygomi m -species  are  self-pollinated  ;  Buckwheat  (P.  fagopyrum)  is  dimorphic 
and  has  long-styled  and  short-styled  flowers  (Fig.  556).  Pol.  bistorta  is  protan- 
drous  and  homostyled. 

About  750  species,  most  of  which  are  found  in  the  temperate  regions  of  the 
Northern  Hemisphere,  some  reaching  as  far  as  the  snow  line  or  into  the  Arctic 
regions  (Oxyria,  Konigia).  Trees  and  shrubs  are  found  in  the  Tropics : 
Coccoloba,  Triplaris.  Rheum  is  Central  Asiatic. — The  thick  rhizomes  of  R. 
officinale  (Rhubarb)  are  officinal.  The  rhizomes  of  the  ordinarily  cultivated 
species,  R.  undulatum  and  rhaponticum,  are  used  in  veterinary  medicine.  The 
following  are  cultivated  as  culinary  plants  for  the  sake  of  their  leaves  : — Rumex 
acetosa  (Sorrel),  R.  patientia,  R.  scutatus,  and  Rheum  undulatum  (petioles). 
Several  species  of  Polygonum  (P.  hydropipef  and  others)  have  a  sharp,  pungent 
taste.  "Buckwheat  "  is  the  mealy  fruit  of  Polygonum  fagopyrum  (Central  Asia) 
and  is  of  value  as  a  farinaceous  food.  P.  cuspidatum  (P.  sieboldi,  Japan)  is  an 
ornamental  plant. — Calligojium  in  sandy  and  stony  deserts. 


FIG.  356. — Flower  of  Polygonum  fagopyrum  in  longitudinal  section:  1,   long-styled  ;  2, 
short-styled ;  a  the  anthers  ;  st  the  stigmas  ;  n  nectary. 

Order  2.  Piperaceae  (Peppers).  Shrubs  or  herbs,  often 
with  nodose,  jointed  stem  ;  leaves  simple,  entire,  often  with  curved 
veins ;  stipules  wanting  (Peperomia)  or  intrapetiolar  and  cap- 
like,  often  enclosing  the  terminal  buds  (Piper).  The  flowers  in 
the  group  Piperece  (Piper,  Fig.  357,  and  Peperomia)  are  borne  in 
spikes  with  fleshy  axes  (club-Like),  seldom  in  racemes,  the  outer 
ones  are  crowded  and  are  5  or  unisexual,  always  small,  naked  and 
without  bracteoles;  generally  stamens  3+3,  and  gynoaceum  3,  but  the 
number  of  the  stamens  may  be  reduced  by  suppression  to  2,  and  the  carpels  to  1. 
The  flowers  are  situated  in  the  axils  of  the  small,  generally 
shield-like  floral-leaves.  The  ovary  is  always  unilocular  and  has 
one  upright,  orthotropous  ovule.  Fruit  a  berry  or  drupe.  Both 
endosperm  and  perisperm  are  present,  the  latter  being  especially 
well  developed  (Fig.  359). 

Piper ;    generally  shrubs   with    scattered   leaves,   and   terminal 


ODi:  ODICOTYLEDONES. 

inflorescences  which  are  crowded  to  one  side  by  the  development 
of  the  highest  lateral  bud,  so  that  they  are  situated  opposite  the 
leaves  (Fig.  357).  Many  species  have  stems  with  an  abnormal 
anatomical  structure. — Peperomia;  chiefly  succulent  herbs,  often 
epiphytes,  with  opposite  or  verticillate  leaves  having  aqueous 
tissue  on  the  upper  side. 

The  group  Saururece  (considered  by  some  as  an  order,  and  perhaps  representing 
a  more  original  type)  has  3-4  carpels  with  many  ovules.  Lactoris  stands  the 
highest  with  regular  3-merous  perianth,  3  +  3  stamens  and  3  carpels,  which  are 


FIG.  357. — Piper  nigrum  -.  branch  with,  fruit  (^). 

united  at  the  base.  Fruit  a  capsule  with  several  seeds.  (It  has  one  specieb 
from  the  island  of  Juan  Fernandez,  and  is  also  placed  in  an  order  of  its  own, 
LactoridacetB,  allied  to  the  Magnoliacese,  through  Drimys). — Saururus  has  naked 
flowers  ;  most  frequently  6  stamens,  and  4  carpels,  free  or  united  at  the  base, 
each  with  2-4  orthotropous  ovules.  Fruit,  small  berries. — Houttuynia ;  stamens 
situated  a  little  upward  on  the  ovaries ;  placentation  parietal ;  capsule  many- 
seeded. 

About  1,000  species  ;  entirely  tropical,  especially  from  South  America  and  East 
India.  They  are  found  chiefly  among  the  underwood  in  damp,  shady  places  ; 
some,  which  are  fleshy  (Peperomia),  live  as  epiphytes  on  trees;  a  few  climb  by 


CUJRVEMBRY-E.  363 

roots. — USES.  Several  Piperaceae  are  used  medicinally  and  for  spices  on  ac- 
count of  their  pungent  properties  and  the  essential  oils  found  in  nearly  all 
parts  of  the  plant.  The  following  are  officinal :  "  Black-pepper  "  (the  unripe, 
dried  fruits)  and  "  White-pepper"  (the  seeds  of  the  ripe  fruits)  of  Piper  nigrum 
(climbing  shrub,  East  Indian) ;  "  Cubeb  "  berries  of  P.  cubeba  (climbing  shrub, 
Java.  "  Long-pepper  "  is  the  unripe  inflorescence  of  P.  longum,  East  India. 


358.  359. 

FIG.  358.— Piper  nigrum  (Diagram),    in  FIG.  359. — Piper  nigrum:  Fruit  in  longi- 

addition  to  the  bract  there  are  two  struc-  tudinal  section,  showing  the  endosperm, 

tures  resembling  bracteoles.  perisperm,  and  pericarp. 

The  leaves  of  P.  anyustifolia  (Matico)  are  officinal.  The  leaves  of  the  Betel- 
pepper  (East  India)  are  used  together  with  the  nuts  of  the  Areca-palm  to  form 
the  well-known  East  Indian  intoxicating  compound  "Betel."  A  good  many 
others  are  also  used. 

Order  3.  Chloranthaceae.  (Chloranthus,  Hedyosmum)  have  opposite  leaves, 
with  stipules  more  or  less  united  at  the  base,  and  inferior  "  drupes."  Ovules 
pendulous.  Only  endosperm.  About  33  species,  Tropical. 


Family  7.     Curvembryse. 

The  plants  in  this  family  have  a  curved  ovule,  and  most  fre- 
quently a  kidney -shaped  seed  (generally  provided  with  fine,  cuticular, 
projecting  warts,  Fig.  362  _B),  with  a  curved,  peripheral  embryo  en- 
closing the  endosperm  which  is  most  frequently  floury  (Figs.  362  G> 
365  H;  for  exceptions,  see  Fig.  366)  ;  the  seeds  in  all  cases  are 
borne  on  a  centrally-placed,  and  in  most  cases  free,  placenta  (they 
are  "  basal  "  when  there  is  only  1  ovule  in  the  ovary,  Fig.  364). 
The  flower  is  regular,  hypogynous  or  perigynous  (Fig.  364)  (.only 
rarely  epigynous)  and  usually  5-merous.  The  flower  which  is  most 
complete  has  5  whorls  (So,  Po,  A5  + 5,  G2  — 3  — 5),  as  in  some 
genera  of  the  Caryophyllaceae  (Figs.  360,  361)  ;  but  from  this 
type  it  becomes  reduced,  the  petals  and  stamens  being  suppressed, 
so  that  finally  5  perianth-leaves,  5  stamens  (opposite  the  perianth- 
leaves),  and  2  carpels  (Fig.  361  F)  only  are  present ;  for  example, 


364  DICOTYLEDONES. 

in  certain  genera  of  the  Carijophyllacex,  in  the  Chenopodiacese, 
Amarantacese,  and  others.  When  the  number  of  stamens  is  in- 
creased to  more  than  5  in  the  whorl,  it  is  always  possible  to  show 
that  some  of  the  stamens  have  been  divided.  The  number  of  the 
carpels  and  ovules  also  becomes  reduced  ;  in  the  highest  there  is  a 
central  placenta,  not  free  in  its  early  stages,  with  a  large  number 
of  ovules ;  in  those  which  are  most  reduced  there  is  only  a  single 
ovule,  which  is  placed  centrally  at  the  base  of  the  ovary  (Fig.  364). 
Somewhat  corresponding  changes  are  found  in  the  fruit,  which  is  a 
many-seeded  capsule  in  those  which  have  many  ovules,  but  a  one- 
seeded  nut  where  there  is  one  ovule.  In  the  most  reduced  forms 
the  flowers  are  generally  unisexual. — Similar  features  are  also  pre- 
sent in  the  vegetative  parts.  Almost  all  the  species  are  herba- 
ceous, the  leaves  are  simple  and  most  frequently  without  stipules. 
The  structure  of  the  stem,  especially  in  Chenopodiaceas,  Amarantacese,  Nyctagi- 
niaceae  and  others,  often  differs  from  that  of  the  ordinary  Dicotyledon.  In  the 
woody  portion  of  the  stem  and  root  several  rings  are  sometimes  formed  which 
resemble  annual  rings  hut  which  are  formed  by  new  cambium-rings  arising  out- 
side the  old  ones  which  then  cease  to  divide. 

Order  1.  Caryophyllacese.  Herbaceous  plants,  with  round, 
nodose  stem ;  leaves  opposite,  slightly  amplexicaul,  simple,  with 
sessile,  undivided,  entire  lamina;  stipules  nearly  always  absent ;  the 
inflorescences  are  dichasia  passing  over  into  unipared  scorpioid 
cymes.  The  flowers  are  regular,  $  or  unisexual,  hypogynous  or 
perigynous,  5-  (or  4-)  merous  with  2-3-4-5  carpels ;  calyx  persistent ; 
corolla  polypetalous.  The  ovary  is  unilocular  (or  originally,  and 
sometimes  also  in  the  later  stages,  plurilocular  below,  e.g.  Viscaria}, 
with  free  styles  and  1-several  curved  ovules  on  a  central,  free 
placenta.  The  fruit  is  a  nut  or  a  capsule  opening  apically  with 
long  or  short  valves  (teeth,  Fig.  362),  equal  to  or  double  the  carpels. 
For  the  seeds  refer  to  the  family.  In  Dianthus  the  embryo  is  straight. 

The  flowers  which  are  most  complete  have  Sn,  Pn,  An  +  n  (obdiplostemo- 
nous),  Gn  where  n  —  5  (Figs.  360,  361  A)  or  4  (Fig.  361  B)  ;  the  carpels  may  be 
placed  opposite  to  the  sepals  (Fig.  360)  or  opposite  to  the  petals  (Fig.  361-4,  B). 
Without  any  change  taking  place  in  the  position  of  the  other  whorls,  the 
carpels  are  next  found  reduced  to  2-3-4  (see  the  genera)  ;  their  number  may 
easily  be  recognised  by  that  of  the  styles.  This  is  the  construction  in  the 
majority  of  the  genera  in  the  two  first  groups.  Stellaria  media  differs  consider- 
ably. It  may  have  (a)  the  flower  as  described  above,  with  G3  ;  (b),  the  corolla 
only  absent,  or  (c)  only  the  petal-stamens  (A5  +  0,  Fig.  361  C),  or  (d)  all  these  as 
well  as  some  of  the  sepal-stamens.  The  same  applies  to  Sagina,  Alsine,  Ccras- 
tiurn,  and  others,  and,  finally,  a  series  of  genera  are  formed,  with  certain  condi- 


CURVEMBRY2E. 


365 


tions  of  reduction  which  have  become  constant,  and  by  a  gradual  series  of  steps, 
lead  to  the  most  reduced  form,  which  has  only  5  sepals  and  5  (or  even  as  far  as 
only  1)  sepal-stamens  (Fig.  361  D,  E,  F).— The 
PETALS  in  the  Alsinea  are  often  deeply  bifid.  The 
sepal-stamens  are  most  frequently  the  longest,  and 
bear  nectaries  at  the  base  (Fig.  363  st).  In  the 
most  complete  forms  the  ovary  has  partition-walls 
in  the  lower  portion  (Fig.  360)  ;  these  do  not, 
however,  reach  to  the  top,  and  generally  soon  dis- 
appear. The  ovules,  when  numerous,  are  situated 
on  the  placenta  in  as  many  double  rows  as  there 
are  carpels.  In  the  number  of  ovules  a  reduction 
from  many  to  1  takes  place  (Fig.  361).  A  com- 
parison proves  that  the  "  free,  centrally  placed  " 
placenta  is  formed  by  the  ventral  portion  of  the  carpels.  The  single  basal  ovule 
in  Herniarin  (Fig.  364),  Scleranthux,  and  others,  is  also  borne  on  the  carpels. 

The  vegetative  branching  is  characteristic.  One  of  the  leaves  in  a  pair  is 
formed  before  the  other,  and  has  a  more  vigorous  axillary  bud  ;  these  stronger 
leaves  stand  in  a  J-spiral,  the  fifth  above  the  first  one,  and  the  branches  are 
consequently  arranged  in  the  same  manner.  In  the  inflorescence,  however,  it 
is  the  upper  or  second  bracteole  (/3)  whose  axillary  bud  (w  in  Fig.  361)  is  most 
advanced.  The  bud  of  the  first  bracteole  (a)  becomes  sometimes  entirely  sup- 
pressed, or  in  some  this  bracteole  itself  is  suppressed. 


FIG.  360.— Diagram  of  Lych- 
nis :  a,  (3  bracteoles. 


FIG.  361. — A-F  Diagrams  of  flowers  of  the  Caryophyllaceaa :  A  Agrostemma ;  B  Sagina; 
C  Stellaria;  D  Corci'g/ola  j  E  Paronychia;  F  Herniaria. 

The  most  original  type  appears  to  be  represented  by  the  Alsineas.  From  this 
form  on  one  side  the  Sileneaa,  adapted  in  a  higher  degree  for  insect-pollina- 
tiou,  are  developed,  and  on  the  otber  side  the  Paronychiese,  with  various  re- 
ductions 

i.  ALSINE^:,  STITCHWORT  GROUP.  Sepals  free,  and  connected 
with  them  stellately  expanded,  slightly  unguiculate  (white  or  in- 


366 


DICOTYLEDONES. 


odorous)  petals;  these,  however,   often  become  suppressed  (Fig. 
363).     The  fruit  is  a  capsule. 

a.  As  many  carpels  as  sepals  (4  or  5).  Cerastium  (Chick- 
weed).  The  petals  are  bind.  Capsule  cylindrical,  frequently 
curved  at  the  top,  and  opening  by  10  teeth  (Fig.  362).—  Malachium 


A.  R  C 

FIG.  362.— Cerastium  aruense:  A  fruit;  I?  seed;  C  section  of  seed. 

differs  only  in  the  5-toothed  capsule  with  bifid  teeth.— Spergula  (Spurry). 
The  petals  are  not  bifid,  capsule  5-valved ;  seeds  winged.  The 
leaves  are  linear,  and  appear  as  if  placed  in  large  numbers  in  a 
whorl,  a  branch  being  situated  in  the  axil  of  each  with  leaves 
placed  very  close  together  at  its  base ;  stipules  membranous. — Sagina 
has  Sn,  Pn,  An+n,  or  An,  Gn,  where  n=4  or  5.  The  corolla  is  often  wanting. 

b.     3  (rarely  2)  carpels   (Fig.  361   G).      Stellaria  (Stitchwort) 
has  deeply  cleft  petals.    The  number  of  stamens  varies  (see  above! 


FIG.  363.— Arenaria  (H'llianthus)  peploides  :   ?  (A)  and  cj -flower  (B,  C). 

— Arenaria  has  entire  petals.  (To  this  group  belong  AUine,  Moehringia^ 
Halianthus,  or  Honckenya  (Fig.  363),  which  differ  from  each  other,  especially  in 
the  form  of  the  seed  and  number  of  the  capsular  valves.)  Spergularia  has 
membranous  stipules,  as  in  Spergula. — Holosteum. 


2.     PARONYCHIE-E   (Figs.  361  D,  E,  F;  364).     Small, 'greenish 


CURVEMBRYJ;. 


367 


plants.  The  leaves,  in  the  majority,  are  opposite,  with  membran- 
ous stipules.  The  flowers  are  most  frequently  arranged  in  small 
dichasia  ;  they  are  small  and  insignificant,  perigynous  (Fig.  364) 
or  hypogynous.  The  corolla  is  in  most  cases  wanting,  and  when 
present  is  very  small ;  in  general  the  calyx-stamens  are  developed, 
but  the  corolla-stamens  may  be  represented  by  small  scales  (Fig. 
364).  Ovary  most  frequently  with  1  ovule.  Fruit,  a  nut,  rarely  a 
capsule ;  it  is  enclosed  by  the  strongly  perigynous  floral  axis 
(torus). 

Scleranthus  (Knapwell)  is  perigynous  with  bell-shaped  torus  ; 
no  corolla;  corolla-stamens  are  wanting  or  rudimentary;  some 
calyx-stamens  may  also  be  absent. — Corrigiola  (Fig.  361  _D)  ; 
Illecebrum;  Paronychia  (Fig.. 361  _E7);  Herniaria  (Figs.  361  F,  364). 

3.  SILENE^E,  PINK  OR  CARNATION  GROUP. 
This  has  a  gamosepalous  calyx  and  un- 
guiculate,  white  or  red,  petals,  with  out- 
growths (ligule,  corona,  paracorolla)  at  the 
throat  of  the  corolla.  These  structures 
are  not  found  in  the  other  groups,  and 
are  merely  outgrowths  at  the  junction  of 
the  limb  and  claw.  The  corolla,  stamens 
and  ovary  are  frequently  raised  above  the 
calyx,  upon  a  lengthened  internode  (gyno- 
phore).  The  flower  has  S5,  P5,  A5  +  5; 
fruit  a  capsule  with  many  seeds. 

a.  5-  (rarely  3-4)  carpellate  ovary. — 
Lychnis  (Campion,  Fig.  360).    The  corolla 
is  longer  than  the  calyx  ;  corona  present. 
The  capsule  is  10-  or  5-toothed,  completely 
1- chambered     or    5 -chambered     at    the 
base, — the  genus  has  been  divided  accord- 
ingly into    several  genera :    Melandrium 

Lychnis,  Viscaria.  Some  species  are  unisexual  by  the  abortion  of  stamens 
or  carpels  (L.  vespertina,  diurna).  Agrostemma  (A.  githago,  Corn-cockle, 
Fig.  361  A)  has  a  long- toothed  calyx,  the  teeth  exceeding  the 
corolla;  corona  absent ;  o-toothed  capsule. 

b.  Tricarpellate. — Silene    (Catch-fly).       Six-toothed    capsule  ;. 
corona  present  in  the  majority. — Cucubalus  has  berry -like  fruits  which. 
finally  become  dry  but  do  not  dehisce. 

C.  Bicarpellate  (2  styles,  4-toothed  capsule). — Dianthus  (Pink)  ;. 
at  the  base  of  the  calyx  1-several  pairs  of  floral-leaves  are  situated  j 


FIG.  364.— Herniaria  glalra : 
(^flower;  b1  longitudinal  gec- 
tion through  the  flower;  c1  stig- 
ma with  two  pollen-grains. 


W.  B. 


BB 


368  DICOTYLEDOXES. 

corona  absent.  The  straight  embryo  is  a  peculiar  exception. — Gypsopliila 
has  a  campanulate,  open  calyx,  5-nerved,  membranous  between  the 
nerves;  corona  absent;  the  flowers  are  generally  small  and 
numerous,  in  a  large,  paniculate  dichasia. — Saponaria  (Soapwort) 
has  corona. 

POLLINATION.  Alsinece  has  ordinary  nectaries  at  the  base  of  the  calyx- 
stamens  (Fig.  336)  :  they  are  frequently  protanclrous  but  may  often,  in  the 
absence  of  cross-pollination  (in  the  less  conspicuous  species)  pollinate  them- 
selves. Their  open  flowers  are  accessible  to  many  kinds  of  insects  (particu- 
larly flies  and  bees).  Gynodicecious  flowers  are  found  in  several  species,  and 
the  £  -flowers  are  then  generally  more  conspicuous  than  the  $  -flowers.  That 
the  ?  -flowers  have  descended  from  £  -flowers  is  seen  by  the  large  staminodes 
found  in  them  (Fig.  363).  Arenaria peploides  is  dioecious  (Fig.  363).  The  Silenfce 
are  as  a  rule  adapted  for  pollination  by  insects  with  long  probosces — especially 
butterflies, — and  they  are  frequently  protandrous,  so  that  at  first  the  calyx- 
stamens  open,  later  on  the  corolla-stamens,  then  the  stigmas  expand.  The 
honey  is  secreted  by  a  ring-like  nectary  round  the  base  of  the  ovary  or  by 
nectaries  at  the  base  of  the  stamens.  Some  only  blossom  and  emit  scent  at 
night  or  in  the  evening  (Lychnis  vespertina,  Silene  nutans,  Saponaria  officinalis) 
and,  like  other  night-flowers,  are  of  a  white  or  pale  colour. 

DISTRIBUTION.  1,100  species,  especially  in  temperate  climates,  fewer  in  the 
colder  zone,  less  still  in  the  Tropics.  The  Paronychieae  are  especially  found  in 
dry,  sandy  fields. 

USES.  "  Soap-root "  (with  Saponin,  forming  a  lather  in  water)  from  Saponaria 
officinalis  was  formerly  officinal,  and  Gypsophila  struthium.  The  seeds  of 
Agrostemma  githago  are  said  to  be  poisonous. — The  following  are  ornamental 
plants :  species  of  Pinks  (D.  caryophyllus,  garden  Pink,  often  with  double 
flowers;  D.  barbatus,  plumarius,  etc].  Lychnis,  Gypyophila,  Silene,  Cerastinm 
{C.  tomentosum  as  edging  for  borders),  Saponaria  officinalis  (often  coronate). — 
Spergula  arvensis  is  sometimes  cultivated. 

Order  2.  Amarantaceae.  The  flowers  are  essentially  the  same  as  in  the 
Chenopodiacece  and  the  extremely  reduced  Caryophyllaceae  (Fig.  361  F) ;  they 
are  regular,  hypogynous,  generally  £ ,  have  5  free  (rarely  slightly  united) 
perianth-leaves  ;  in  front  of  these  5  stamens,  which  are  often  united  at  their 
base  into  a  shorter  or  longer  tube  and  have  stipule-like  teeth  between  them  (the 
division  Gomphretiea;  has  2-locular  anthers,  each  of  which  opens  longitudinally) ; 
and  a  2-3  carpellate  gynoeceum  with  one  loculus  and  most  frequently  one,  moro 
rarely  several,  ovules;  the  fruit  is  a  nut,  more  rarely  (in  Celosia,  Amarantii*, 
Gomphrena)  a  capsule,  dehiscing  irregularly,  or  like  a  pyxidium.  The  characters 
which  especially  separate  them  from  the  allied  orders  are  found  in  the  perianth. 
The  perianth-leaves  are  not  green  and  herbaceous,  but  membranous,  dry,  and 
often  coloured  ;  they  are  frequently  produced  into  a  bristle  or  awn ;  they  have 
also  both  subtending  floral-leaves  and  2  large  bracteoles  similar  to  the  perianth; 
all  these  dry  leaves  persist  without  alteration  after  the  withering  of  the  flower. 
— The  flowers  are  without  scent.  They  are  arranged  in  spike-  or  capitulum- 
like  inflorescences;  sometimes  placed  singly,  sorm times  aggregated  in  the 
panicle-like  inflorescences ;  in  others,  on  the  contrary,  in  dichasia.  The 


•CURVEMBRY2E.  369 

majority  are  herbs,  some  are  shrubs.     The  leaves  are  scattered,  or  opposite, 
but  always  simple  and  without  stipules  ;  some  are  smooth,  others  hairy. 

450  species  ;  especially  in  the  Tropics,  principally  S.  Am.  and  E.  Ind. : 
few  are  found  outside  these  countries. — Only  a  few  are  used;  some,  chiefly 
E.  Indian  species,  are  cultivated  as  ornamental  plants:  Amaranthus  (Fox- 
tail); Gomphrena  globosa  ;  Celosia  cristata  (Cock's-comb)  remarkable  for 
its  fasciated  inflorescence ;  Alternanthera.  Some  are  employed  as  culinary 
plants  in  the  Tropics,  and  in  a  few  of  the  E.  Indian  species  the  seeds  are 
farinaceous,  arid  used  for  food. 

Order  3.  Chenopodiaceae.  Generally  herbaceous  plants  like 
the  Caryophyllaceee,  but  the  leaves  are  arranged  spirally  (except 
Salicornia),  and  are  simple,  exstipulate  ;  they  are  generally  fleshy 
and  like  the  stem  "  mealy,"  that  is,  covered  with  small  hairs,  whose 
large  spherical  terminal  cell  readily  falls  away;  otherwise  they 
are  seldom  hairy.  The  inflorescences  are  generally  flower-clusters 
borne  in  panicles.  Bracteoles  generally  absent.  Flowers  gene- 
rally unisexual :  with  the  single  exception  of  Beta  the  flowers  are 
hypogynous ;  they  are  regular,  small  and  inconspicuous,  with  single, 
green,  5-leaved,  but  more  or  less  united  perianth ;  5  stamens  opposite 
the  perianth,  and  a  2-5-carpellate,  unilocular  ovary  with  1  basal, 
curved  ovule;  but  in  some  genera  the  number  of  the  perianth-leaves 
and  stamens  is  reduced  to  3-2-1-0.  The  fruit  is  generally  a 
nut, — thus  this  flower  and  fruit  are  the  same  as  in  the  reduced 
Caryophyllaceas  (Fig.  361  F).  The  seed  is  similar  to  that  generally 
found  in  the  family  (for  exceptions  see  the  genera). 

The  floral  diagram  most  frequently  present  is  the  same  as  in  Fig.  361 F.  There 
is  no  indication  of  corolla  or  of  corolla-stamens,  which  may  be  supposed  to  have 
belonged  to  the  plant,  but  which  are  now  entirely  and  completely  suppressed. 
This  order  appears  to  have  been  an  offshoot  from  the  Caryophyllacete. — The 
perianth  persists  after  the  withering  of  the  flower,  and  envelopes  the  nut ;  it  is 
very  variable,  and.  together  with  the  position  of  the  seed,  the  form  of  the 
embryo,  the  sex  of  the  flowers,  etc.,  gives  the  characters  of  the  genera. 

I.  CHEXOPODIE^E,  GOOSEFOOT  GROUP  (Fig.  365),  has  £"  (or  poly- 
gamous) flowers,  with  regular  5-parted  perianth  (C)  ;  the  em- 
bryo is  ring-like  (J?).  The  leaves  have  the  ordinary  flat  forms. 
— Chenopodium  (Goosefoot).  The  flower  is  hypogynous,  and  the 
fruit  (which  is  compressed)  perfectly  free  ;  Mulberry-like  collec- 
tions of  fruits  are  formed  in  some  species  (sub-genus  Blitum}  by 
the  perianth  becoming  finally  fleshy  and  coloured. — Beta  (Beet, 
Mangold,  Fig.  365)  differs  from  all  genera  in  the  perianth,  which 
finally  becomes  cartilaginous,  being  epigynous  (I)).  Small,  most 
frequently  2-3-flowered  clusters  without  bracteoles,  situated  in  a 


370 


DICOTYLEDONES. 


long,   interrupted  axis  (-4,  B)  ;  the   flowers   and   fruits   in   each 
cluster  are  more  or  less  united  individually,  and  fall  off  together — 


FIG.  365.— Beta  vulgaris. 

they  are  commonly  known  as  seeds  (.EJ,  F).     The  seed  lies  horizon- 
tally.— Hablitzia  (H.  tamnoides). 

2.     SALSOLE^],  SALTWORT  GROUP,  has  cylindrical  or  semi-cylindri- 
cal leaves.     Perianth  as  in  the  preceding  group  ;  the  fruit  is  most 


FIG.  366.  —Salsola  soda :  embryo. 

frequently  compressed.      The  two  first  mentioned   genera   differ 
from  most  of  the  others  in  the  order  in  having  a  spirally-coiled,. 


CURVEMBRY.E.  371 

and  not  a  ring-like  embryo,  so  that  the  endosperm  is  slight  or 
wanting  (Fig.  366).  These  plants  are  sometimes  placed  as  a 
group  by  themselves,  SPIROLOBEJ; — in  contradistinction  to  which 
the  others  are  termed  CYCLOLOBE^;. — Salsola  (Saltwort)  ;  -leaves 
subulate,  with  spiny  tips  ;  the  flowers  have  2  spinous  bracteoles : 
during  the  ripening  of  the  fruit  a  tough  leathery  wing  is  de- 
veloped transversely  to  the  back  of  the  perianth. — Chenopodina 
deviates  from  Chenopodium  chiefly  in  the  embryo  and  want  of  endosperm. — 
Kochia  has  a  somewhat  similar  perianth  to  Salsola,  but  a  ring-like  embryo ; 
it  differs  from  the  others  in  being  hairy. 

3.  SALICORNIE^;,  GLASSWORT  GROUP.     Salicornia  (Glasswort)  has 
a  very  different  appearance.     The  stems  are  succulent,  jointed,  and 
almost  leafless  ;  the  leaves  opposite,  very   small,  sheath-like  and 
connate  ;  there  is  a  depression  in  the  axil  of  each  leaf,  in  which  a 
small    3-flowered    dichasiam    without    bracteoles    is   sunk ;    the 
flowers  have  a  trimerous  perianth,  1-2  stamens  and  1  carpel.     No 
endosperm.     S.  herbacea  on  clayey  beaches. 

4.  ATRIPLICE^E.     This    group    has   most    frequently    unisexual 
flowers  ;  the  ^ -flower  has  a  4-5  partite  perianth,  but  the  $  -flower 
differs    from    it.      Atriplex   is  monoecious  or  polygamous,  the  $  - 
flower  is  naked,  but    has    2  large,   herbaceous  bracteoles    which 
expand  during  the  ripening  of  the  fruit,  and  often  become  warted 
and  fringed,  enveloping  the  compressed  nut.     The  section  Dichospermum 
has  two  kinds  of  9  -flowers,  one  like  those  just  described,  the  other  similar  to  the 
Chenopodium- lowers,  which  have  been  deprived  of  their  stamens,  and  the  fruits 
of  which  are  depressed,  not  pressed  together  from  the  sides  ;    some  (e.g.  A. 
hortensis)  have  even  three  kinds  of  nuts.     All  the  flowers  of  Atriplex,  which 
present  vertical  fruits,  are  accessory  shoots,  which  stand  beneath  the  ordinary 
flower-clusters,  a  rather  singular  relation.— Spinacia  (Spinach)  is  dioecious  ; 
(£ -flower:  perianth,  4  (-5)  ;  stamens,  4  (-5);    $ -flower:  tubular, 
2-4-partite  perianth,  hardening  during  the  ripening  of  the  fruit, 
and  uniting  with  the  compressed  nut;  in  8.  oleracea,  it  also  forms 
thorns ;    4     long    stigmas. — Halimus   has    the    2    long  bracteoles 
almost   entirely    united    and    ultimately   adhering  firmly   to   the 
fruit, 

5.  BASELLE.E.     A  somewhat  exceptional  group  with  more  or  less  perigynous 
flowers  and  2  bracteoles.     Hasella,  Boussincjaultia,  Ullucus.     The  perianth  is 
sepaloid ;  ovary  1-ovuled.     In  Basella  the  perianth  is  fleshy,  enveloping  the 
nut,  and  the  cotyledons  are  so  rolled  together  that  a  tranverse  cut  divides  them 
in  two  places  (as  in  Spirolobeae).     Herbaceous  climbing  plants. 

POLLINATION.  Wind-  and  self-pollination,  as  far  as  is  known ;  the  insignifi- 
cant flowers,  devoid  of  honey,  appear  to  exclude  insect-pollination.  —  520 
species.  Most  of  them  are  annual  (out  of  26  native  species  only  5  are  peren- 


372  DICOTYLEDON  ES. 

nial) ;  inhabiting  salt-marshes  and  salt-steppes,  and  growing  as  weeds  (most 
frequently  on  garden  or  field  soil  containing  manure)  in  this  country,  especially 
species  of  Chenopodium  and  Atriplex.  The  majority  are  found  outside  the 
Tropics,  and  play  a  very  important  part,  for  example,  in  the  Asiatic  salt- 
steppes.  They  grow  gregariously  in  large  masses. 

USES.  Comparatively  few.  The  only  important  one  is  Beta  vulgar  is  (from 
the  Mediterranean  basin),  with  its  different  varieties,  viz.  Beet-root,  Cattle  beet 
or  Eed-beet,  Sugar-beet,  and  others.  These  are  biennial,  making  in  the  first 
year  a  root  which  acts  as  a  reservoir  of  reserve  material,  with  a  rosette  of  leaves, 
and  in  the  second  year  using  this  material  in  the  production  of  a  long  stem, 
leaves  and  flowers.  The  primary  root  has  been  developed  by  cultivation  into  a 
very  thick  and  fleshy  tap-root ;  its  mode  of  increase  in  thickness  deviates  from 
that  of  other  roots,  concentric  rings  of  vascular  bundles  being  formed  from  a 
cambial  ring  developed  outside  the  previous  ring.  In  this  way  several  rings, 
of  vascular  bundles  separated  by  medullary  rays,  alternating  with  rings  of 
parenchyma,  may  be  found  in  the  root  of  a  Beet.  Besides  Beta  vulgaris,  var. 
hortensis  (Beet-root),  the  following  are  also  cultivated  :  var.  cicla  (Leaf-beet, 
"Mangold,"  or  "  Koman  Spinach"),  Spinacia  oleracea  and  Atiiplex  hortensis 
as  Spinach  ;  a  form  of  the  latter  and  of  Spinach  are  grown  as  ornamental 
plants.  The  tubers  of  Ullucus  tuberosus  are  used  as  potatoes ;  Chenopodium 
quinoa,  in  Chili  and  Peru,  is  an  important  farinaceous  plant.  Soda  is  made 
from  some  (Salsola  kali,  Chenopodina  maritima  and  others).  Aromatic  proper- 
ties are  rare :  Chenopodium  ambrosioides  and  botrys. 

Order  4.  Batidaceae.  Batis  maritima,  a  bushy  West  Indian  maritime 
plant. 

Order  5.  Phytolaccaceae.  The  £  (eometimes  unisexual),  regular,  some- 
times slightly  perigynous  flowers  are  inconspicuous  and  have  a  single  sepaloid 
or  coloured  4-5-leaved  perianth  (generally  united  at  the  base) ;  stamens  either 
in  1  whorl  in  the  spaces  between  the  perianth-leaves  or  in  1  whorl  opposite  the 
perianth-leaves,  or  in  2,  one  of  which  alternates  with  these ;  but  the  number 
may  be  increased  by  the  splitting  of  one  or  of  both  the  whorls  to  as  many  as 
10-15-20-25.  Carpels  sometimes  only  one,  sometimes  many  (4-10)  placed  in  a 
whorl,  either  free  or  united  into  a  gynoeceum  with  a  corresponding  number  of 
loculi  in  the  ovary  ;  but  in  all  cases  each  carpel  bears  only  its  own  style  and 
1  ovule.  The  fruit  is  a  berry  (or  nut,  capsule,  or  schizocarp). — Mostly  herbs 
or  herbaceous  shrubs,  with  scattered,  simple  leaves  without  stipules  (Petiveriece 
have  stipules).  Inflorescences,  most  frequently  racemes  or  spikes,  which  in 
some  instances  are  apparently  placed  opposite  to  a  leaf,  being  displaced  by  a 
more  vigorous  growth  of  the  axillary  bud.  Embryo  always  bent. — Petiverict 
has  a  straight  embryo  with  rolled  cotyledons. — Phytolacca,  Pircunia,  Microtea, 
Seguieria,  Rivina  (Pr4,  A4,  Gl;  berry),  and  others. 

The  following  plant  is,  with  some  doubt,  placed  near  this  order :  Tliely- 
gonum  cynocrambe  ;  monoecious.  <J  -flowers:  perianth,  2-leaved ;  stamens  indefi- 
nite. ?  -flowers :  perianth-leaves  united,  3-toothed ;  Gl,  style  gynobasic. 
Fruit  a  drupe.  An  annual  plant ;  Mediterranean.  Branching  anomalous. 

About  90  species ;  in  tropical  and  temperate  countries,  principally  America 
and  Africa. — The  red  juice  in  the  fruits,  especially  of  Phytol.  dccandra,  is  used 
for  colour  ug  wine. 


CURVEMBRYJE.  373 

Order  6.  Portulacaceae  (Portulacas).  The  flowers  are 
regular  (except  Montia),  hypogynous  (except  Portulaca)  and  §  . 
The  diagram  which  applies  to  the  majority  of  genera  is  that  in 
Fig.  367,  but  with  all  the  5  stamens  completely  developed:  it  may 
be  considered  as  the  Chenopodiaceous  diagram  with  the  addition 
of  2  bracteoles  in  the  median  line  (m-n,  these  by  some  are  con- 
sidered as  sepals),  and  with  a  petaloid  perianth  (usually  desig- 
nated "corolla").  The  "petals"  fall  off  very  quickly,  and  are 
sometimes  wanting.  Most  frequently  5  stamens,  situated  oppo- 
site the  "  petals,"  but  in  other  genera  the  number  varies ;  Montia 
has  only  3  stamens  (by  suppression  of  the  two  anterior  and 
lateral,  Fig.  367),  others  again  have  more  than  5,  some  a  large 
and  indefinite  number.  This  may  be  explained  partly  by  the 
appearance  of  a  second  whorl  of  stamens  alternating  with  the 
first,  and  partly  by  the  splitting  (dedoublement)  of  the  stamens. 
Gynceceum  most  frequently  tricarpellate,  ovary  unilocular  with 
1-several  basal  ovules  (sometimes  on  a 
branched  placenta,  as  in  certain  Caryo- 
phyllacece).  The  fruit  is  a  capsule,  more 
rarely  a  nut. — The  majority  are  annual 
herbaceous  plants  with  scattered,  entire 
leaves,  often  fleshy  and  smooth,  with  or 
without  rudimentary  stipules  (dry,  mern- 

J  *  FlG-  367.— Montt'a. 

branous,  modified  into  hairs).     Inflorescence  Diagram  of  flower 

cymose. 

Portulaca  (Portulaca) :  flower,  epigynous  or  semi-epigynous  ; 
fruit,  a  pyxidium.  The  stamens  vary  in  number,  and  are  most 
frequently  placed  in  groups  (in  consequence  of  splitting)  opposite 
the  petals. — Montia  :  the  corolla  is  slightly  gamopetalous,  but 
cleft  on  the  posterior  side  (Fig.  367),  and  as  a  consequence  of  the 
larger  size  of  the  lateral  petals,  slightly  zygomorphic ;  3  stamens. 
— Calandrinia  ;  Talinum  ;  Anacampseros  ;  Claytonia. 

125  species  ;  mostly  in  warm  and  temperate  countries,  especially  the  arid  parts 
of  S.  Am.  and  the  Cape.  Montia  fontana  (Blinks)  is  a  native  plant.  Portulaca 
oleracea  is  cultivated  as  a  pot-herb  in  the  south  of  Europe.  A  few  species  of 
Portulaca  and  Calandrinia  are  ornamental  plants. 

Order  7.  Nyctaginiaceae.  The  characteristic  feature  of  this 
order  is  the  single,  regular,  united,  and  often  petaloid  perianth,  the 
lower  part  of  which  generally  persists  after  flowering  and  em- 
braces the  fruit  as  a  false  pericarp.  The  upper  portion  is  most 


374  DICOTYLEDONES. 

•   • 

frequently  valvate  and  folded,  or  simply  valvate  in  aestivation. 
The  number  of  stamens  varies.  The  free  gynoeceum  is  unicar- 
pellate  and  has  1  ovule.  The  fruit  is  a  nut,  but  becomes  a  false 
drupe,  since  the  lower  persistent  portion  of  the  perianth  becomes 
fleshy  (as  in  Neea,  where  this  fleshy  part  is  almost  always  crowned 
by  the  upper  persistent  part  of  the  perianth.  In  the  majority  of 
the  Mirabileae  the  lower  part  liecomes  the  dry  anthocarp,  while 
the  upper  petaloid  part  falls  away  after  flowering).  Finally,  a 
peculiar  involucre  is  formed  around  the  flowers  by  free  or 
united  floral-leaves.  —  The  majority  are  herbs,  some  are  trees 
(Pisonia,  etc.)  ;  Bougainvillea  is  a  liane.  The  stems  are  often 
nodose  and  swollen  at  the  nodes ;  the  leaves  are  simple,  penni- 
nerved,  scattered,  or  opposite,  without  stipules.  In  some,  the  vascular 
bundles  are  scattered  ;  stem  anomalous. 

Mirabilis ;  the  structure  of  the  stem  is  abnormal.  Dichasial 
branching  with  continuation  from  the  second  bracteole,  thus  form- 
ing unipared  scorpioid  cymes.  The  perianth  is  petaloid,  funnel- 
shaped,  and  has  a  folded  and  twisted  aestivation  resembling  that 
of  the  corolla  of  the  Convolvulacece ;  the  upper  coloured  portion 
falls  off  after  the  flowering.  Outside,  and  alternating  with  it,  is 
a  5-partite,  sepaloid  involucre  of  5  spirally- placed  floral-leaves. — 
Oxybaphus ;  the  involucre  envelops  1-3  dichasial  flowers.  — 
JBougainvillea-,  the  involucre  is  rose-coloured,  3-leaved,  and  en- 
velops 3  flowers  (placed  laterally ;  the  terminal  flower  wanting). 
The  leaves  of  the  involucre  in  Boerhaavia,  Pisonia,  Neea,  and  others 
are  reduced  to  teeth  or  scales. 

157  species  ;  mostly  in  tropical  countries,  and  especially  S.  Am.  Species 
of  Mirabilis  (Am.)  are  ornamental  plants.  Them  is  found  in  Neea  the'ifera 
Oersted  (discovered  by  Lund  in  Lagoa  Santa,  Brazil),  which  may  be  used  as  a 
tea-plant. 

Order  8.  Aizoacese.  Only  3  ivhorls  are  found  in  the  flower,  which  alternate 
with  one  another  when  their  leaves  are  equal  in  number.  The  first  is  sepaloid, 
the  third  one  the  carpels,  and  the  intervening  one  is  either  uncleft,  in  which 
case  it  is  developed  as  stamens,  or  it  is  divided  into  a  large  number  of  members 
which  then  all  become  stamens  (arranged  in  groups),  or  the  outermost  ones 
become  developed  as  petals.  The  fruit  is  most  frequently  a  capsule  with 
several  loculi.  Most  of  the  species  are  herbs  with  thick,  fleshy  stems,  and 
exstipulate  leaves.  The  structure  of  the  stem  is  usually  anomalous. 

i.  AIZOIDE^E  have  hypogynous  or  perigynous  flowers  with  (4-)  5  perianth- 
leaves;  stamens  single,  or  (by  splitting)  in  groups  of  2-3,  alternating  with  the 
perianth-leaves.  The  gynoeceum  (with  3-5  carpels)  has  3-5  loculi  in  the  ovary, 
and  most  frequently  numerous  ovules  in  each  loculus,  borne  on  the  central 
placenta  formed  by  the  edges  of  the  carpels.  The  fruit  is  a  capsule.  The 


CACTIFLORjE. 


375 


inflorescences  are  dichasia  and  unipared  scorpioid  cymes. — Aizoon,  Mollugo, 
Sesuvium,  and  others  are  herbs  or  bushes,  most  frequently  hairy. 

2.  MESEMBBIANTHEME;E  have  semi-  or  wholly-epigynous  flowers. — Tetra- 
gonia.  The  perianth  is  4  (more  rarely  3-5-6)  -merous.  Stamens  single,  or  (by 
splitting)  in  groups  alternating  with  the  perianth- leaves.  There  is  an  indefinite 
number  of  carpels,  and  each  loculus  of  the  ovary  contains  only  1  pendulous 
ovule.  Fruit  a  nut  or  drupe.  The  flowers  arise  singly  in  the  leaf-axils,  with  an 
accessory  foliage-bud  below  them ;  in  some  instances  there  is  also  an  accessory 
flower  between  this  bud  and  the  flower.  Southern  hemisphere,  especially  at 
the  Cape ;  T.  expansa,  New  Zealand  Spinach,  is  a  fleshy  plant  which  is 
•cultivated  as  a  pot-herb  (Japan,  Austr.,  S.  Am.). — Mesembrianthemum  :  the 
flowers  are  5-merous;  the  numerous  linear  petals  and  the  still  more  nu- 
merous stamens  all  arise  by  the  splitting  of  5  or  4  protuberances  (primordia) 
alternating  with  the  sepals.  The  ovary  presents  another  characteristic  peculi- 
arity :  the  carpels  alternating  with  the  5-4  stamens  form  an  ovary  (with  several 
loculi)  with  the  ovules  at  first  borne,  as  in  other  cases,  on  the  inner  corner 
•of  the  inwardly-turned  carpels;  but  during  the  subsequent  development  the 
whole  ovary  is  so  turned  round  that  the  placentae  become  parietal  and  the 
ovules  assume,  apparently,  a  position  very  rarely  met  with  in  the  vegetable 
kingdom :  on  the  dorsal  suture  of  the  carpels.  Shrubs  or  under-shrubs,  more 
rarely  herbs  with  fleshy  stems  and  simple,  entire,  more  frequently  thick  or 
triangular  leaves,  containing  a  quantity  of  water.  The  flowers  open  about  noon, 
and  are  brightly  coloui'ed,  generally  red  or  red-violet,  but  odourless.  The 
capsules  dehisce  in  rainy  weather.  300  species,  mostly  found  at  the  Cape. 
Some  are  ornamental  plants.  M.  crystallinum  (the  Ice-plant)  and  others  are 
covered  with  peculiar,  bladder-like,  sparkling  hairs,  the  cell-sap  of  which  con- 
tains salt — these  serve  as  reservoirs  of  water. 

Family  8.     Cactiflorae. 

The  position  of  this  family  is  very  doubtful ;  but  it  seems  in 
many  respects  to  approach  Mesembrianthemum.  Some  botanists 
place  it  near  to  the  Ribesiaceae ;  others,  again,  to  the  Passifloraceas. 
Only  1  order. 

Order  Cactacese  (The  Cacti).  The  flower  is  epigynous,  £, 
regular,  and  remarkable  for  its  acyclic  structure;  there  are,  for 
instance,  a  large  number  of  spirally- placed  sepals  and  petals,  which 
gradually  pass  over  into  one  another,  and  which  in  some  species,  to 
a  certain  extent,  arise  from  the  walls  of  the  ovary  as  in  Nymphsea 
(Fig.  383  A,  B}.  The  petals  are  free;  rotate,  opening  widely  in 
Opuntia,  Pereskia,  and  Ithipsalis  ;  erect  and  united  at  their  base 
into  a  Shorter  or  longer  tube  in  Cereus,  Epiphyllum,  Mammillaria, 
Echinocactus,  Melocactus,  and  others  (Fig.  369).  Stamens  numerous, 
attached  to  the  base  of  the  corolla ;  gynceceum  formed  of  many 
carpels,  with  one  style,  dividing  into  a  number  of  branches  corres- 
ponding to  the  number  of  carpels  ;  the  ovary  has  one  loculus  with 


376 


D1COTYLEOONBS. 


many  parietal  placenta?  ;  the  ovules  are  anatropous,  on  long  and 
curved  f  unicles.  Fruit  a  berry  with  exendospermous  seeds.  The 
fruit-pulp  is  mainly  derived  from  the  funicles. — The  external 


FIG.  3G3.—  A  Etfnnocactus:  a  position  of  a  leaf-lamina ;  b  a  lateral  shoot  on  ihe  displaced 
axillary  bud.  B  Pereslda :  b  a  foliage-leaf  on  a  small  thorny  branch  -which  is  subtended 
by  a  foliage-leaf  which  has  fallen  off  and  left  a  scar  (a). 

appearance  of  the  Cactaceaa  is  very  peculiar;  Pereskia,  which  has 
thick  and  fleshy  leaves  (Fig.  368),  deviates  the  least;  foliage- 
leaves  of  the  usual  form  are  wanting  in  the  other  genera,  or  are 
usually  very  small,  and  quickly  fall  off  and  disappear  (Opuntia), 
or  are  modified  into  thorns  ;  the  stem,  without  normal  foliage- 
leaves, — so  characteristic  a  feature  in  this  order, — makes  its  appear- 
ance after  the  two  normally 
developed  cotyledons.  The 
stems  are  fleshy,  perennial, 
and  may  finally  become 
woody.  In  some  they  are 
elongated,  globose,  pointed, 
and  more  or  less  dichoto- 
mously  branched,  e.g.  in 
several  of  the  llliipsalis 
species,  which  live  mostly 
as  epiphytes  on  trees  ;  m 
others,  elongated,  branched, 
globose,  or,  most  frequently, 
more  or  less  angular  (pris- 
matic) or  grooved  and  pro- 
vided with  wings,  and  either 
columnar  and  erect  (as 
much  as  about  20  metres  in 
FIG.  3G9.— Ediinopsis.  height  and  1  metre  in  cir- 


POLYCARPICJE.  377 

cumferetice,  as  in  G.  giganteus  in  New  Mexico)  or  climbing  by  roots 
(Cereus  and  Ithipsalis-species) ;  in  others  again,  compressed,  more  or 
less  leaf-like,  often  with  a  ridge  in  the  centre  (winged),  branched 
and  jointed  :  Epiphyllum,  Pliyllocactus,  Opuntia,  some  species  of 
Ehipsalis ;  others  are  thick,  short,  spherical  or  ovoid,  unbranched  or 
only  slightly  branched,  and  either  studded  with  prominent  warts 
(mammill(E)  each  of  which  supports  a  tuft  of  thorns  (Fig.  368  A ; 
Mammillaria  and  others)  or  with  vertical  ridges,  separated  by 
furrows  (rows  of  mammillae  which  have  coalesced)  in  Melocactus^ 
Echinocactus,  Echinopsis  (Fig.  369) ;  at  the  same  time  the  ovary  in 
some  is  embedded  in  the  stem  so  that  leaves  or  leaf-scars,  with 
tufts  of  thorns  in  their  axils,  may  be  observed  on  the  ovary  just  as 
on  the  stem. — The  flattened  shoots  of  the  Cactacea3  are  formed 
in  various  ways,  either  by  the  compression  of  cylindrical  axes 
(Opuntia)  or,  as  in  Melocactus,  etc.,  from  winged  stems  in  which 
all  the  wings  are  suppressed  except  two. 

The  thorns  are  produced  directly  from  the  growing  points  of  the  axillary 
buds,  and  are  modified  leaves.  The  axillary  bud  is  united  at  its  base  with 
its  subtending  leaf,  which  as  a  rule  is  extremely  rudimentary ;  and  these 
together  form  a  kind  of  leaf-cushion,  larger  in  some  genera  than  in  others. 
This  leaf-cushion  attains  its  highest  development  in  Mammillaria,  in  which  it 
is  a  large,  conical  wart  (see  Fig.  368  A),  bearing  on  its  apex  the  tuft  of  thorns 
and  rudimentary  lamina. — The  seedlings  have  normal  cotyledons  and  a  fleshy 
hypocotyl. 

All  the  species  (1,000?)  are  American  (one  epiphytic  species  of  ftldpsulis  is 
indigenous  in  S.  Africa,  Mauritius  and  Ceylon),  especially  from  the  tropical 
table-lands  (Mexico,  etc.).  Some  species,  especially  those  without  thorns,  as 
Jihipealw,  are  epiphytes.  Opuntia  vulgaris,  the  fruits  of  which  are  edible,  is 
naturalized  in  the  Mediterranean.  The  cochineal  insect  (Coccus  cacti)  lives  on 
this  and  some  closely  allied  species  (O.  cocchiellifera,  etc.),  particularly  in 
Mexico  and  the  Canary  Islands.  Several  are  ornamental  plants. 

Family  9.     Polycarpicae. 

The  flowers  as  a  rule  are  $  ,  regular  and  liypogynous ;  however  in 
some  orders  they  are  unisexual,  e.g.  in  the  MyristicaceaB,  or 
zygomorphic  (in  Monkshood  and  Larkspur  in  the  Ranunculaceoe)  ; 
in  the  Lauracere,  (Fig.  386)  for  example,  perigynous,  and  in 
Nymph&a  (Fig.  383)  even  partially  epigynous  flowers  are  typical. — 
The  flowers  are  acyclic  in  very  many  of  the  genera  of  the  two  first 
orders,  if  not  completely  so,  at  any  rate  in  the  numerous  stamens 
and  carpels,  thus  denoting  an  old  type.  It  is  a  remarkable 
characteristic  that  in  the  majority  of  the  orders  the  number  3  pre- 
vails in  the  calyx  and  corolla ;  the  number  5  also  occurs,  but  the 


378 


D1COTYLEDONES. 


number  2  is  seldom  met  with.  Most  orders  have  a  double 
perianth ;  chorisis  does  not  occur,  suppression  is  rare,  and  the  parts 
of  the  flower  are  developed  in  acropetal  succession.  The  most 
characteristic  feature  in  the  order  is  the  free,  one-leaved,  as  a  rule 
numerous  carpels  (apocarpous  gynoeceum).  The  number  of  carpels 
in  some  of  the  last  mentioned  orders  dwindles  down  to  1  (e.g.  the 
Berlieridese  and  Myristicacese) .  The  carpels  in  Nymphseacete  become 
united  into  one  pistil  (syncarpous),  a  condition  which  we  also  find 
distributed  among  the  other  orders. 

Endosperm  occurs  in  almost  all  the  orders  (except  e.g.  Lauracese). 
The  nutritive  tissue  in  Cdbombese  and  Nymphseese  is  chiefly  peri- 
sperm. 

Order  1.  Ranunculacese.  Nearly  all  are  herbs  (except 
Clematis').  The  leaves  are  scattered  (except  Clematideas),  they 
have  a  large  sheath  with  broad  base  (no  stipules),  and  are  most 


FIG.  370.— Diagram 
of  Aquilegia  vulgaris: 
sp  spur.  A  cyclic 
flower. 


FIG.  371. — Diagram  of  a  dichasium 
of  Rununculus  acer :  alt  a1,  and  j8x, 
/31,  bracteoles  (the  buds  in  the  axils 
of  the  bracteoles,  a  and  a1,  are  con- 
tin  tied  antidromously).  The  flower 
has  cyclic  calyx  and  corolla,  but 
acyclic  (&)  stamens. 


Fio.372. — Diagram  of  an 
acyclic  Ranunculaceous 
flower  (only  3  stamens  are 
indicated).  The  spiral  of 
the  sepals  has  a  diver- 
gence of  | ;  that  of  the 
corolla  and  subsequent 


frequently  palminerved  with  palmate  lobes.  The  flowers  are 
hypogynous,  with  most  frequently  a  well  pronounced  convex  re- 
ceptacle (Figs.  374  JB,  380),  J ,  regular  (except  Delphinium  and 
Aconitum')  ;  their  structure  varies  very  much  ;  in  some  the  leaves 
are  verticillate,  in  others  arranged  spirally ;  in  others,  again,  both 
modes  of  arrangement  are  found.  It  is  a  characteristic  feature 
that  the  various  series  of  leaves  (especially  calyx  and  corolla)  are 
not  so  distinct  or  so  sharply  divided  as  is  usual.  The  leaves  of  the 
perianth  are  free,  imbricate  (except  Clematideas) ;  stamens  numerous, 
with  most  frequently  extrorse  anthers  ;  gynoeceum  free,  apocarpous 
(except  Nigella  and  partly  Helleborus) ,  with  1  or  several  ovules 
(Figs.  373,  378,  379)  borne  on  the  ventral  suture.  The  fruit  is 


POLTCARPIC.E.  379 

either  a  nut  or  a  follicle  (Acttea  has  berries).     The  seed  has  a  large^ 
oil-containing  endosperm  and  a  small  embryo  (Fig.  374). 

The  main  axis  generally  terminates  in  a  flower,  and  the  lateral  axes  branch 
in  a  cymose  manner  (Fig.  371).  The  flowers  show  the  following  differences  in 
constiuction:  VEBTICILLATE  (EUCYCLIC),  i.e.  constructed  all  through  of  alternating 
whorls:  Aquilegia  (Fig.  370),  Xanthorhiza,  and  sometimes  Eronthis.  SEMI- 
VERTICILLATE  (HEHicYCLic)  i.e.  with  sepals  and  petals  in  alternate  whorls,  and 
the  others  arranged  spirally:  Ranunculus  (Fig.  371),  Myosurus,  P&onia  and 
several  other  genera  entirely,  or  in  certain  species  only.  SPIRAL-FLOWERED 
(ACYCLIC)  i.e.  all  the  leaves  are  arranged  spirally,  so  that  sepals  and  petals  do 
not  alternate  the  one  with  the  other,  even  though  they  are  the  same  in  number : 
Adonis  (Fig.  372),  Aconitum,  Delphinium-species,  Nigella-species,  Helleborus. 
The  leaves  of  the  calyx  are  in  this  instance  arranged  on  a  spiral  of  § ;  those  of 
the  corolla  on  f,  f ,  ^  or  ^-,  and  stamens  and  carpels  likewise  on  higher 
fractions  of  the  same  series. 

The  genera  Caltha,  Anemone,  Thalictrum  and  Clematis  have  a  single  perianth, 
which  is  most  frequently  petaloid ;  it  is  thus  apparent  that  the  sepals  are 
petaloid,  and  the  leaves,  which  in  other  genera  have  developed  as  petals,  are  in 
these  instances  stamens.  The  calyx  is  similarly  petaloid  in  the  genera  Helleborus, 
Eranthis,  Nigella,  Delphinium  and  Aconitum  ;  but  the  petals  are  present  in  these 
instances  in  unusual  (hprn-like)  forms,  and  almost  entirely  given  up  to  the 
function  of  nectaries,  a  function  they  already  possess  in  Ranunculus.  According 
to  a  more  recent  theory  the  "  honey  -leaves  "  are  transformed  stamens,  which 
have  lost  the  function  of  reproduction  ;  the  perianth  is  then  single,  and  most 
frequently  petaloid.  [Those  leaves  in  the  flowers  of  many  Eanunculaceae 
which  bear  nectaries  are  termed  by  Prantl  honey-leaves,  and  comprise  those 
leaf-structures  of  the  flower  whose  essential  function  lies  in  the  production 
of  nectar,  and  which,  independent  of  the  differentiation  of  the  perianth  into 
calyx  and  corolla,  are  derived  from  the  stamens  by  the  loss  of  their  reproductive 
functions.  Clear  transitional  forms  are  found  between  the  two  series  of  the 
perianth  (e.g.  between  the  sepaloid  and  petaloid  perianth-leaves  of  Anemone 
japonica,  A.  decapetala,  Trollitis -species)  while  transitional  forms  are  never 
found  between  perianth-  and  honey-leaves  (with  the  exception  of  Aquilegia 
vulgaris,  var.  stellata).  In  Anemone  and  Clematis  the  honey-leaves  pass 
gradually  into  the  stamens,  and  agree  with  the  stamens  in  the  other  Ranun- 
culacea3  in  their  arrangement,  development,  and  scant  system  of  veins  (except 
Nigella}.  In  Delphinium,  sect.  Consolida,  the  two  honey-leaves  placed  in  front 
of  the  unpaired  perianth-leaf  are  united  into  one,  as  shown  by  the  veins  (twice 
three  veins  arranged  symmetrically).  The  honey-leaves  of  Aquilegia,  Callian- 
themum,  and  the  majority  of  the  Ranunculus-species  serve,  by  reason  of  their 
large  circumference,  as  organs  of  attraction,  and  on  this  account  are  considered 
as  petals  by  other  authors. — The  same  position  in  the  flower  which  the 
honey-leaves  assume  is  found  occupied  by  staminodes,  without  nectar,  in  some 
Coptis-species,  inAnemonopsis,  Actcca  sect.  Euactcca,  (e.g.  A.  racemosa),  Clematis 
sect.  Atragene;  in  the  last-named  they  closely  surround  the  stamens,  in  Actcea 
they  are  petaloid. — A  perianth,  sharply  differentiated  into  calyx  and  corolla, 
and  destitute  of  honey-leaves,  is  found  in  Anemone,  sect.  Knowltonia  (Cape),. 


880 


DICOTYLEDONES. 


Adonis,  Pceonla. — The  perianth  of  the  Rannnculaceae  is  considered  by  Prantl 
to  .be  usually  petaloid. — The  nectaries  arise  in  the  Ranunculacese  (1)  on 
normal  stamens  (Clematis  sect.  Viorna),  (2)  on  the  honey-leaves  (this  is 
generally  the  case),  and  (3)  on  the  carpels  (Caltha  and  the  majority  of  Trollius- 
species). — As  the  result  of  his  researches  upon  the  Ranunculaceae,  Prantl 
does  not  agree  with  the  view  advanced  by  Drtule  (Schenk,  Hand.  d.  Bot.  iii.) 
that  the  petals  in  general  have  proceeded  from  the  metamorphosis  of  the 
stamens  (K)~\ . 


FIG.  373. — Ovaries  in  longitudinal  section:  «the  ventral  suture;  d  the  dorsal  suture : 
A,  B  Clematis;  C  Ranunculus ;  D  Mijosurus. 

Tlie  most  primitive  form  of  fruit  is  undoubtedly  the  pod  formed  by  one  carpel, 
on  the  edges  of  which  (along  the  ventral  suture)  two  rows  of  ovules  are  situated  : 
Paeoniere,  Helleboreae,  Delphinieas  (Fig.  379).  In  a  great  many  genera  the 
number  of  ovules  has  been  limited  to  one  perfect  one,  which  is  placed  in  the 
central  plane  under  the  united  leaf-edges,  and  sometimes  also  some  barren 
ovules  above  it  (Fig.  373).  The  fruitlets  in  this  case  become  achenes,  and 
are  present  in  much  larger  numbers  than  when  there  are  follicles. 


FIG.  374. — Helleborus  niger  :  A  flower;  B  receptacle;  pet  petals  FIG.  375.— Caltha 

.(honey-leaves);   pi  stamens  and  carpels;  C  seed ;  D  anther  (cross  palustris:  fruit, 

section);  alb  endosperm. 

The  following  have  FOLLICLES  :  Pasoniese,  Helleborese  (except 
Actaeo)  and  Delphiniese ;  ACHENES:  Ranunculese,  Anemoneae  and 
Clematideas. 


POLYCARPICJ). 


381 


FIG.  376.—  Aquttegia  vulgaris. 


A.  Follicles  (Figs.  375,379),  with  many  ovules,  situated  in 
two  rows  along  the  ventral  suture.  Actaa  has  berries,  Nigella  has 
capsules  of  several  loculi. 

I.  P^EONIEJ;,  PEONY  GROUP.  This  has  regular,  acyclic  flowers 
with  a  normal,  most  frequently  5-leaved,  imbricate  calyx ;  large, 
coloured  petals,  and 
introrse  anthers. 
Slightly  perigynous. 
Surrounding  the 
base  of  the  carpels 
a  ring-like  swelling 
of  the  receptacle 
("  disc  ")  is  present, 
which  is  largest  in 
P.  maul  an.  The 
follicles  are  more 
or  less  fleshy  or 
leathery.  Mostly 
herbs,  with  pinnati- 
sect  or  decompound 
leaves  and  large, 
solitary  flowers;  a  gradual  transition  may  be  traced  from  the 
foliage-leaves  to  the  petals.  P&onia;  Hydrastis. 
'  2.  HELLKBOREJ;,  HELLEBORE  GROUP.  This  has  regular  flowers  with 
most  frequently  a  coloured 
calyx.  The  petals  (honey- 
leaves)  are  modified  into 
nectaries;  they  may  be  horn- 
like, provided  with  a  spur,  or 
of  a  similarly  unusual  form, 
or  they  may  be  entirely  ab- 
sent. Anthers  often  extrorse. 
— Trollius  (Giobe-flower1). 
The  flower  is  acyclic:  many 

1  According  to  Prantl,  some 
species  of  Trollius  (T.  europccus, 
and  asiatiacus)  have  a  perianth, 
differentiated  into  calyx  and  cor- 
olia,  which  does  not  pass  over  into 
the  honey-leaves.  The  outer  leaves 
of  the  perianth  have  frequently  an 
incised  apex,  the  intermediate  ones  FIG.  377  —  CaltJia  palustris  (nal.  size). 


382 


D1COTYLEDONES. 


petaloid  sepals,  succeeding  these,  most  frequently,  several  linear, 
dark  yellow  petals,  which  bear  a  naked  nectary  at  the  base; 
finally,  many  stamens  and  carpels  arranged  in  a  spiral  (-§-,  -/T). — 
Caltha  (Marsh-marigold,  Figs.  375,  377);  5  (-7)  yellow  sepals, 
no  petals.  The  foliage-leaves  have  a  large  amplexicaul  sheath. — 
Helleborous  (Hellebore)  has  pedate  leaves.  The  flower  is  acyclic, 
with  5  large,  regular,  persistent,  often  petaloid  sepals  (-| )  ;  small, 
hornlike  petals  (honey-leaves ;  most  frequently  13,  divergence  T8g) 
and  generally  few  carpels  (Fig.  374). —  Coptis. —  Isopyrum. — 
Eranthis  (Winter  Aconite),  like  Anemone,  has  a  3-leaved  invo- 
lucre and  most  frequently  trimerous  flowers,  6  large  petaloid  sepals, 
6  petals  (tubular  honey-leaves),  6  oblique  rows  of  stamens,  3-6  carpels. 
Aquilegia  (Columbine,  Fig.  376)  ;  the  flower  is  entirely  cyclic 


FIG.  378.—  Nigella  :  A,  B  fruit  of  N.  damascene,  entire,  and  cut  transversely.    C  Petal 
(honej-leat)  of  N.  arvensis.     D  Petal  of  N.  damascene. 

and  has  large  spurs  on  all  the  5  petals  (funnel-shaped  honey- 
leaves)  ;  S5  coloured,  P5,  A5  x  (8-12),  G5  in  regular  alternation 
(Figs.  376,  370)  ;  the  innermost  stamens  are  often  staminodes 
(Fig.  370).—  Nigella  (Love-in-the-mist,  Fig.  378)  has  5  sepals 
and  8  small,  two-lipped  petals  cleft  at  the  apex  (the  nectary  is 
covered  by  the  under-lip;  Fig.  378  (7,  D).  The  5  carpels  are 
more  or  less  completely  united;  and  a  many-carpellate  ovary 
with  free  styles  is  formed  in  some.  Large  air-chambers  in  the 
external  wall  of  the  ovary  are  formed  in  N.  damascena  (Fig.  378). 
—  Acted  (Baneberry)  has  coloured  sepals,  either  no  petals  or  an 


sometimes  present  transitional  forms  to  the  inner,  and  sometimes  there  is  a 
distinct  boundary  between  them. 


POLYCARPICJfi. 


333 


indefinite  number,  and  only  1   carpel.     The  fruit  is  a  berry   (or 
follicle). — Cimicifuga,  Garidella,  Xanthorhiza  (85,  P5,  A5+5,  G5). 

3.  DELPHINIE.E,  LARKSPUR  GROUP.      Zygomorphic  flowers  with 
coloured  calyx  ;   the  2  posterior  petals   (honey-leaves)  are  trans- 
formed into  nectaries,  the  others  are  small  or  absent  altogether. — 
Aconitum  (Monkshood)  ;  5  sepals,  of  which  the  posterior  one  (Fig. 
379  A]  is  helmet-shaped  ;  most  frequently  8  petals  (as  in  Fig.  372), 
of  which  the  two  posterior  ones  (honey-leaves)  are  developed  into 
long-clawed  nectaries  (Fig.  379  A,  fe)  enveloped  by  the  helmet-like 
sepal ;  the   others  are  small,   or  are  to  some  extent  suppressed. 
Stamens  on  a  spiral  of  f-^ ;   generally  3   carpels.     Perennial  herbs. — 
Delphinium  (Larkspur)  ;  very  closely  allied  to  Aconitum,  but  the 
anterior  4  petals  are  most  frequently  wanting,  and  the  2  posterior 
ones  have  each  a  spur,  which 

is  enclosed  by  the  posterior 
sepal,  the  latter  being  also 
provided  with  a  membranous 
npur.  Stamens  and  carpels  ar- 
ranged on  a  spiral  of  §,  ^53,  ^-. 
In  V.  ajacis  and  consolida  there  is 
apparently  only  1  petal  (by  the 
fusion  of  4)  and  1  carpel. 

B.  Fruit  achenes.  Many 
carpels,  each  with  only  1 
ascending  (Fig.  373  G),  or 
pendulous  (Fig.  373  D),  per- 
fect ovule  ;  often  also  rudi- 
mentary ovules  above  it  (Fig. 
873  A,  B).  Fruit  achenes. 

4.  RANUNCULE^E,   BUTTER- 
CUP GROUP,  has  double  peri- 
anth.    Myosurus  and  Adonis  have 
pendulous  ovules  as  in  Anemoneae 

(Fig.  373  D) ;  Ranunculus,  with  Batrachium  and  Ficaria,  erect  ovules  (Fig.  273  C) 
and  downwardly-turned  radicle. — Ranunculus.  Most  frequently  S5,  P5, 
many  spirally-placed  stamens  and  carpels  (Figs.  371,  380).  The 
petals  (honey-leaves)  have  a  nectary  at  the  base,  covered  by  a  small 
scale.  Batrachium,  Water  Eanunculus,  deviates  by  the  achenes  being  trans- 
versely wrinkled  ;  dimorphic  leaves.  Ficaria  has  3  sepals  and  7-8  petals  arranged 
in  §-§.  F.  ranunculoides  (the  only  species)  has  tuberous  roots,  which  spring  from 
the  base  of  the  axillary  buds,  and  together  with  these,  serve  as  organs  of  repro- 
duction. The  embryo  has  only  1  cotyledon.— Myosurus  (Mouse-tail)  has- 
W.  B.  C  ° 


FIG.  379.— Aconitum  napellus.  A  Flower  in 
longitudinal'section,  below  are  the  2  bracteoles  ; 
a  half  of  helmet-like  sepal;  b  and  c  other 
sepals;  7c  nectary;  /  carpels.  B  Ovary  in 
longitudinal  section ;  C  the  samrj  transversely  ; 
d  dorsal  suture ;  v  veutral  suture. 


384 


DICOTYLEDONES. 


small  prolongations  from  the  5  sepals  ;  5  narrow  petals  which  bear 
the  nectaries  near  the  apex  ;  sometimes   only  5  stamens,  and  an 

ultimately  very  long  recep- 
tacle, with  numerous  spirally- 
arranged  achenes  (Fig.  381). — 
Adonis  is  acyclic  (Fig.  372)  ; 
most  frequently  5  sepals  with 
a  divergence  of  -|,  8  petals  of 
-|,  indefinite  stamens  and  car- 
pels of  f  or  ~.  The  corolla 
has  no  nectary. 

5.  AXEMOXE.E,  ANEMONK 
GROUP,  has  a  single  perianth. 
'{Pendulous  ovules  (Fig.  373  D),  radicle  turned  upward) — Anemone  has  a 
single,  petaloid,  most  frequently  5-6-leaved  perianth,  and  beneath 
the  flower  most  frequently  an  involucre  of  3  leaves,  placed  close 
together  in  the  form  of  a  whorl.  In  A.  nemorosa,  ranunculoides, 
•etc.,  the  involucral  leaves  resemble  foliage-leaves ;  in  A.  Tiepatica 


PIG.  380.— Flower  of  Rammculus  sceleratus 
in  longitudinal  section. 


PIG.   381.— Myosurus  minimus:  c  cotyledons;    m  the  foliage-leaves;   /' the  floral  axis 
-with  the  carpels,  and  g  the  same  without;  y  insertion  of  perianth. 

they  are  situated  close  under  the  perianth,  and  resemble  sepals, 
and  in  the  sub-genus  Pulsatilla  they  stand  between  the  foliage- 
leaves  and  floral-leaves.  The  style  of  Pulsatilla  finally  grows  out, 


POLYCARPIC^E.  385 

in  the  form  of  a  feather.  The  main  axis  of  A.  hepatica  has  unlimited 
growth  (it  is  biaxial),  and  the  flowers  are  borne  laterally  in  the  axils  of  the 
scale-leaves  ;  in  the  others  (uniaxtal)  the  flower  is  terminal,  and  the  rhizome 
becomes  a  sympodium  after  the  first  flowering — Thalicrtrum  (Meadow 
Rue)  has  no-  involucre  ;  4-5-leaved,  greenish  perianth.  The  re- 
ceptacle is  flat.  The  stamens  are  brightly-coloured  and  have  long  filaments ; 
1-5  accessory  flowers  may  occur  in  the  leaf-axils  of  the  panicle-like  inflorescence. 

6.  CLEMATIDEJ:,  CLEMATIS  GROI?P.  This  differs  from  all  the 
others  in  the  valvate  aestivation  of  the  calyx  an  1  its  opposite  leaves. 
There  are  4  (-several)  petaloid  sepals  ;  petals  are  absent,  or 
linear  (Atragene).  Ovule  1,  pendulous.  Achenes,  often  with 
prolonged,  feathery  style.  The  majority  of  the  genera  are  shrubs, 
and  climb  by  their  sensitive,  twining  leaf-stalks.  —  Clematis ; 
Atragene. 

POLLINATION.  The  flowers  are  conspicuous  either  by  coloured  petals  (honey- 
leaves)  (Ranunculus,  Pteonia)  or  coloured  sepals  (Helleborus,  Anemone,  Caltliu, 
etc.),  or  by  both  (Aquilegia,  Delphinium),  or  by  the  coloured  stamens  (Tlialic- 
trum).  Some  have  no  honey  (Clematis,  Anemone,  Thalictrum),  and  are  gener- 
ally visited  by  insects  for  the  sake  of  their  pollen.  Others  have  nectaries 
on  the  corolla  (Ranunculus,  Trollius,  Helleborus,  Nigella,  Aconitum,  etc.), 
more  rarely  on  the  stamens  (Pulsatilla,  Clematis-species),  or  the  carpels 
(CaltJia),  or  the  calyx  (certain  species  of  P&onia).  The  honey  is  readily 
accessible  in  the  flat,  open  flowers,  and  these  flowers  also  may  easily  pollinate 
themselves.  There  is  marked  protandry  where  the  honey  lies  deeply  hidden,  as 
in  Aquilegia,  Delphinium,  and  Aconitum.  Helleborus  and  some  Ranunculus- 
species  are  protogynous. 

About  680  species  ;  especially  in  northern  temperate  climates,  and  extending 
to  the  Polar  and  Alpine  regions.  Only  the  Clematidece  are  tropical. 

The  order  has  an  abundance  of  acrid,  vesicant  properties  (R.  acer,  sceleratus, 
etc.),  and  poisonous  alkaloids  (Helltborus  niger  is  poisonous).  OFFICINAL  : 
Aconitum  napellus  (aconitine ;  leaves  and  tuberous  roots) ;  the  rhizome  of 
Hydrattit  canadensis  from  N.  Am.  (the  alkaloid  hydrastine).  The  order,  how- 
ever, is  best  known  for  its  ornamental  plants ;  almost  all  the  genera  have 
species  which  are  cultivated  for  their  beauty.  Sweet-scented  flowers  are 
absent. 

Order  2.  Nymphaeaceae  (Water  Lilies).  WATER  PLANTS; 
generally  with  large,  floating  leaves,  and  large  solitary  flowers  ; 
sepals  3-5,  petals  3-  00,  stamens  6-  00,  carpels  3-  00.  The  flower 
is  hypogyiious,  but  in  the  Nympliseeee  different  degrees  of  epigyny 
are  found,  and  from  this  fact,  as  well  as  from  the  carpels  being 
united  into  one  pistil,  the  family  forms  a  lateral  offshoot  from 
the  Ranunculacese,  with  much  greater  modification.  The  seed 
often  has  an  aril,  and,  in  the  majority,  a  farinaceous  nutritive 


386 


DICOTYLEDONES. 


tissue,  partly  endosperm,  partly  perisperm  (Fig.  383  0).  The 
embryo  has  2  thick  cotyledons  and  a  small  hypocotyl  ;  the 
plumule  is  well  developed,  with  2-4  leaves. 

1.  CABOJIBE^E.  3-4  species  (Tropical  S.  Am.),  resembling  the  Water 
Bauunculus,  with  two  kinds  of  leaves,  the  submerged  being  dissected  and  the 
aerial  peltate.  The  flowers  are  eucyclic,  trimerous,  with  2-3  free,  epigynous 
carpels.  The  ovules  are  situated  on  the  central  line  of  the  carpel — an  almost 
unique  circumstance.  Endosperm  and  perisperm.  Cabomba ;  Brasenia. 

2.  NELUMBONEJ;.  The  leaves  are  peltate,  raised  on  long  stalks- 
high  above  the  water.  Large,  hypogynous  flowers  (Fig.  382); 
sepals  4-5  ;  petals  numerous ;  stamens  numerous;  carpels  several, 
distinct.  The  receptacle  is  very  remarkable,  being  raised  above 
the  stamens,  and  developed  into  an  inverted  conical  body  on 

the  apex  of  which  the  nut- 
like  fruits  are  embedded  in 
pits.  Endosperm  is  wanting,, 
but  the  embryo  is  large  and 
has  well  developed  cotyledons. 
—Nelumbo,  2  species.  N.  lutea  (N. 
Am.)  ;  N.  speciosa  (E.  Ind.)  was 
sacred  amongst  the  ancient  Hindoos- 
and  Egyptians  (the  Lotus  flower); 
its  seeds  are  used  as  food. 

3.  NYMPH J:EJE,  WATER  LILY 
GROUP.  The  carpels  are  united 
into  one,  many-locular  ovary, 
whose  numerous  ovules  are 
situated  on  the  surface  of  the 
partition  walls  (as  in  the  Pop- 
pies) ;  the  stigma  is  sessile 
and  radiating,  the  number  of  rays  corresponding  to  the  number 
of  carpels  (Fig.  383).  The  fruit  is  a  spongy  berry  with  many 
seeds,  which  have  a  large  perisperm  in.  addition  to  the  endosperm 
(Fig.  3S3  G). 

Sepals,  petals,  and  stamens  often  pass  gradually  over  the  one- 
into  the  other,  the  petals  becoming  narrower  by  degrees,  and 
bearing  anthers  on  each  side  of  the  apex,  which  gradually  become 
larger  anthers  in  proportion  to  the  filament,  until  the  perfect 
stamen  is  developed.  The  long-stalked  leaves  are  floating,  and 
most  frequently  cordate,  elliptical,  leathery,  with  a  shiny  surface,, 
sometimes  (as  in  Victoria  regia  and  Euryale  ferox)  with  strongly 


FIG.  382. — Nelumbo  nucifera:  vertical  section 
through  the  receptacle. 


POLYCARPICJE. 


387 


projecting  thorny  ribs  on  the  lower  surface.     In  the  intercellular 
passages  of  the  leaves  are  some  peculiar,  stellate  cells. 

Nuphar  has  5  sepals,  and  an  hypogynous  flower.  The  petals,  which 
are  small,  have  a  nectary  on  the  back ;  the  coloured  inner  side  of  the  sepals 
functions  as  petals  ;  the  ovate  gynoeceum  is  quite  free. — N.  luteum  is  a  native 
plant  (Yellow  Water-Lily),  with,  most  frequently,  13  petals  and  10-16  loculi  in 
the  ovary.  The  rhizome  is  horizontal,  as  much  as  5-6  cm.  in  thickness,  and 
bears  on  its  under  surface  a  number  of  roots,  which  on  dying-off  leave  deep 
scars ;  the  leaves  are  borne  in  spiral  lines,  and  the  flowers  are  solitary  in 
certain  leaf-axils.  The  construction  of  the  rhizome  is  very  peculiar ;  the 
vascular  bundles  are  scattered  and  closed  as  in  a  monocotyledonous  stem. 


A  EC 

Fio.  333. — Nymphcea;  A  flower  in  longitudinal  section,  the  most  external  leaves  being 
removed;  I?  fruit;  C  seed  of  Nnphar  (longitudinal  section);  the  perisperin  at  the  base, 
the  endosperm  at  the  top  surrounding  the  embryo. 

Nymplisea  has  4  sepals,  and  the  flower  is  more  or  less  <'pi<jy- 
nous.  Petals  and  stamens  are  inserted  at  different  heights  on  the 
ovary  to  just  beneath  the  stigma  (Fig.  383).  NympJisea  alba  (White 
Water-Lily).  Victoria  regia  from  the  Amazon,  and  Euryale  ferox 
from  Asia,  have  entirely  epigynous  flowers.  The  shield-like  leaves  of 
Victoria  are  as  much  as  2  metres  in  diameter,  and  the  edge  is  bent  up  to  a 
height  of  5-14  cm. ;  the  flowers  are  20-40  cm.  in  diameter,  and  change  in 
twenty-four  hours  from  white  to  rose-red.  A  development  of  heat,  as  much  as 
14°C.  above  the  temperature  of  the  air,  together  with  a  strong  formation  of 
car'»onic  acid,  has  been  observed  during  flowering. 

POLLINATION.  Ntjmphaa  alba  and  other  species  of  the  sub-genus  Symphy- 
topleura  are  self-pollinated ;  the  sub-genus  Leptopleura  is  insect-pollinated. 
Nuphar  and  Victoria  can  effect  self-fertilisation ;  Euryale  is  self-fertilised,  often 
in  entirely  closed  and  submerged  flowers. — The  dissemination  of  the  seeds  in 
Nuphar  luteum  is  effected  by  the  fruit,  which  rests  on  the  water,  becoming  de- 


388  DICOTYLEDONES. 

tached  from  its  stalk,  and  d«-hiscing  from  the  base  upwards  so  that  the  seeds  are 
set  free;  while  in  Nymphcea  albai\\e  spirally-twisted  stalk  draws  the  fruit  under 
water,  and  it  dehisces  by  its  upper  part  being  thrown  off  as  a  hood,  and  the 
seeds  which  are  enclosed  in  air-tight  sacs  rise  to  the  surface  of  the  water.  In 
this  condition  they  are  able  to  float  and  can  only  sink  to  the  bottom  when  the 
air  has  disappeared. 

53  species ;  in  fresh  water  in  all  parts  of  the  world,  but  especially  in  the 
Tropics. — The  rhizomes  and  seeds  of  some  may  be  used  as  food  ;  Euryaleferox 
is  even  cultivated.  Xymphcea  ccerulea  and  Lotus  were  sacred  among  the 
Egyptians. 

Order  3.  Ceratophyllacese.  About  3  species.  Aquatic 
plants,  submerged,  rootless ;  leaves  cartilaginous,  verticillate,  dis- 
sected into  repeatedly  dichotomous  branches  which  are  finely 
toothed  ;  only  one  of  the  leaves  in  a  whorl  supports  a  vegetative 
branch.  The  flowers  are  monoecious,  axillary.  Inside  ths  6-12 
perianth-leaves  is  situated  in  the  ^-flower  10-20  stamens  with 
thick  connective,  and  in  the  ?  -flower  a  gynceceum  formed  by 
one  carpel,  with  one  orthotropous  and  pendulous  ovule,  which  has 
only  one  integument.  Fruit  a  nut,  which,  in  some  species,  bears 
on  each  side  a  pointed  horn,  and  at  the  apex  a  similar  one,  formed 
by  the  persistent  style. — The  embryo  has  an  unusually  well  de- 
veloped plumule  with  several  whorls  of  leaves.  The  plant  is  root- 
less throughout  its  whole  life. — Ceratophyllum  (Horn- wort). 

Order  4.  Anonacese.  Sf-pals  3;  petals  3  +  3  (most  frequently  valvate) ; 
succeeding  these  (as  in  the  Ranunculacece)  are  numerous  acyclic  stamens  and  an 
apocarpous  gynceceum  ;  the  flowers  are'  hypogynous,  regular  and  $  ,  generally 
very  large  (2-3  cm.  in  diameter),  and  the  leaves  of  the  perianth  are  more  or  less 
fleshy  or  leathery.  The  majority  have  syncarps  with  berry-like  fruitlets,  but 
in  Anona  and  some  others  the  carpels  fuse  together  into  a  large,  head-like  fruit 
— a  kind  of  composite  berry.  The  seeds  have  ruminate  endosperm  as  in 
Myristica. — Trees  or  shrubs  with  alternate,  simple,  entire,  penninerved  leaves 
without  stipules.  450  (700  ?)  species  ;  especially  tropical.  The  best  known  are 
Anona  cherimolia,  squamosa  and  reticulata  (all  from  America)  cultivated  on 
account  of  their  large,  delicious  fruits.  Some  have  acrid  and  aromatic  pro- 
perties (Xylopia,  C  anew  go. — the  flowers  of  t"he  latter  yield  Ylaug-ylang) ; 
Artabotrys  odoratissima  ;  Asimina  (N.  Am.). 

Order  5.  Magnoliaceae.  Trees  or  shrubs  with  scattered,  often  leathery, 
entire  leaves,  generally  with  stipules,  which  (as  in  Ficus)  are  rolled  together  and 
form  a  hood  round  the  younger  internodes  above  them,  and  are  cast  off  by  the 
unfolding  of  the  next  leaf,  leaving  a  ring-like  scar.  The  endosperm  is  not 
ruminate.  Corolla  imbricate.  Fruit  a  syncarp. 

A.  MAGNOLIEJE.  The  flowers  are  borne  singly,  and  before  opening  are  en- 
veloped in  an  ochrea-like  spathe  which  corresponds  to  the  stipules  of  the  foliage- 
leaves.  The  perianth  generally  consists  of  3  trimerous  whorls,  the  external  one 
of  which  is  sometimes  sepaloid  (Liriodcndron,  and  the  majority  of  Magnolia- 


POLYCARPICJ:.  389 

species),  sometimes  coloured  like  the  others;  the  perianth  is  sometimes  many- 
seriate.  Numerous  spirally -placed  stamens  and  carpels.  The  latter  are  situated 
on  the  elongated,  cylindrical  receptacle,  and  are  individually  more  or  less  united, 
except  in  Liriodendron,  where  they  are  free.  This  last  genus  has  winged  achenes  ; 
the  fruitlets  in  Magnolia  open  along  the  dorsal  and  ventral  sutures,  and  the 
seeds  then  hang  out,  suspended  by  elastic  threads  formed  from  the  vascular 
bundles  of  the  funicle  and  raphe  ;  they  are  red  and  drupaceous,  the  external 
layer  of  the  shell  being  fleshy— a  very  rare  occurrence. 

B.  ILLICIE.E  has  no  stipules.  The  carpels  are  situated  in  a  whorl  on  a  short 
receptacle.  Follicles,  one-seeded.  The  leaves  are  dotted  by  glands  containing 
essential  oil.  Illicium ;  Dniiujs. 

70  species ;  in  tropical  or  temperate  climates ;  none  in  Europe  or  Africa.  They 
are  chiefly  used  as  ornamental  plants,  e.g.  the  Tulip-tree  (Liriodendron  tulipi- 
fera,  N.  Am.),  Magnolia  grandiflora  (N.  Am.),  M.  yulan  and  fuscata  (China), 
and  others.  The  remains  of  Liriodendron  occur  as  fossils  in  the  Cretaceous 
and  Tertiary  periods. — The  fruits  of  Illicium  anisatum  (Star-aniseed  from 
Eastern  Asia)  are  OFFICINAL.  The  bark  of  Drimys  winteri  (S.  Am.)  is  also 
strongly  aromatic. 

Order  6.  Calycanthaceae.  These  are  very  closely  related  to  the  Magnoliaceae, 
but  differ  in  having  perigynous  flowers  with  many  perianth-leaves,  stamens 
and  (about  20)  carpels  in  a  continuous  spiral,  seeds  almost  devoid  of  endos2)erm 
with  rolled  up,  leaf-like  cotyledons,  and  leaves  opposite  on  a  square  stem.— There 
are  some  species  in  N.  America  (Calycanthus  florida,  occidentalis,  etc.)  and 
1  in  Japan  (Chimonanthus  prcccox),  all  strongly  aromatic. 

Order  7.  Monimiaceae.  Aromatic  shrubs  with  opposite  leaves.  Perigynous 
flowers.  The  anthers  dehisce  by  valves  like  those  of  the  Lauracece,  and  the 
Mouimiaceas  may  thus  be  considered  as  an  apocarpous  form  of  this  order.  They 
are  also  closely  related  to  Calycanthacea.  150  species,  tropical. — Hedycarya, 
Mollinedia,  Monimia. 

Order  8.  Berberidaceae  (Barberries).  —  The  regular,  $, 
hypogyiious  flowers  are  dimerous  or  trimerous  and  have  regu- 
larly alternating  whorls  of  free  sepals,  petals,  and  stamens  and 


FIG.  334.— Diagram  of  Berleds.  FIG.  385. — Berbm's  :  carpel  with  2  stamens. 

1  unilocular  carpel :  the  corolla  and  stamens  have  each  2  whorls, 
the  calyx  at  least  2.  The  anthers  open,  as  in  Lauraceee,  by 
(2)  valves,  but  are  always  introrse  (Fig.  384).  The  pistil  has 


390  D1COTYLEDONES. 

a  large,  disc-like,  almost  sessile  stigma  (Fig.  385),  and  in  the  ovary 
several  erect  ovules  are  placed  close  to  the  base  of  the  ventral 
suture.  The  fruit  is  most  frequently  a  berry.  Seeds  endosper- 
mous. — Shrubs  or  herbs  with  scattered,  most  frequently  compound 
leaves  (without  stipules),  and  racemose  inflorescences. — They  show  a 
relationship  to  the  Lauraceae  in  the  number  of  the  parts  of  the  flower  and  the 
dehiscence  of  the  anthers. 

Berberis  is  a  shrub  ;  it  has  sepals  3  +  3,  petals  3  +  3,  stamens  3  +  3 
(Fig.  384).  The  petals  (honey-leaves)  bear  internally  at  the 
base  2  darkish-yellow  nectaries.  The  filaments  are  sensitive 
at  the  base,  and  suddenly  bend  inwards  if  touched  at  that  spot 
(Fig.  385).  The  racemes  often  have  a  terminal,  5-merous  flower;  they  are 
borne  on  dwarf-branches.  The  leaves  on  the  long-branches  develope  into 
thorns,  but  the  buds  in  their  axils,  in  the  same  year  as  themselves,  develope  as 
the  short-branches  with  simple  foliage-leaves,  articulated  at  the  base,  from 
which  fact  some  authorities  have  considered  that  the  leaf  is  compound  with  a 
single,  terminal  leaflet — Mahonia  has  imparipinnate  leaves.  The 
flower  has  3  whorls  of  sepals.  Otherwise  as  in  Berberis. — 
Epimedium ;  herbs  with  spurred  petals  ;  the  flowers  dimerous ;  4-5  whorls  of 
sepals,  2  of  petals  and  stamens.  Fruit  a  capsule.  Leontice,  fruit  dry.  The 
anthers  of  PodopJiyllum  dehisce  longitudinally. — Nandina.  Aceranthus. 

100  species;  North  temp.,  especially  Asia:  fossils  in  Tertiary.  Berberis 
vulgaris  is  a  native  of  Europe.  This  and  other  species,  together  with  Mahonia 
aquifolium  (N.  Am.),  Epimedium  alpinum,  etc.,  are  cultivated  as  ornamental 
plants.  Several  have  a  yellow  colouring  matter  in  the  root  and  stem. 
OFFICINAL  :  the  rhizome  of  Podophyllum  peltatum  (from  N.  Am.)  yields  po^o- 
phyllin. 

Order  9.  Menispermacese.  This  order  has  derived  its  name  from  the 
more  or  less  crescent-like  fruits  and  seeds.  Dioecious.  The  flowers  are  2-3- 
•merous,  most  frequently  as  in  Berberis  (S3  +  3,  P3  +  3,  A3  +  3),  with  the. 
difference  that  there  are  3  free  carpels,  each  with  1  ovule ;  in  some  genera, 
however,  the  number  is  different.  Stamens  often  united  into  a  bundle  (as  in 
Myristica) ;  anthers  dehiscing  longitudinally  ;  fruit  a  drupe. — The  plants  (with 
herbaceous  or  woody  stems)  belonging  to  this  order  are  nearly  all  twining  or 
climbing  plants,  and  have  scattered,  palmate  or  peltate,  sometimes  lobed  leaves 
without  stipules.  Structure  of  stem  anomalous.  Cocculus,  Menispermum, 
Cissampelos,  Anamirta. 

150  species  ;  Tropical ;  very  rich  in  bitter  and  poisonous  properties.  OFFI- 
CINAL :  Calumba-root  from  Jateorhiza  columba  (E.  Africa).  The  following  are 
cultivated  as  ornamental  plants : — Menispermum  canadense  (N.  Am.)  and  M. 
dahuricum  (Asia).  The  fruits  of  Anamirta  cocculus  (E.  Ind.)  are  very  poisonous 
<"  Graias-of-Paradise  "  ;  the  poisonous  matter  is  picrotoxine). 

Order  10.  Lardizabalaceae.  This  order,  by  the  free,  apocarpous  carpels, 
belongs  to  a  more  primitive  type,  and  by  the  united  stamens  to  a  more  developed 
one.  Akebia ;  Ilolliwllia  ;  principally  climbing  or  twining  shrubs.  About  7 
species  in  S.E.  Asia  and  S.  Am. 


POLYCARPIC^:. 


391 


Order  11.  Lauraceae  (True  Laurels).  Trees  or  shrubs; 
the  leaves,  always  without  stipules,  are  simple,  most  frequently 
scattered,  lanceolate  or  elliptical,  entire,  penninerved,  finely  reti- 
culate (except  Cinnamomum  with  3-5-veined  leaf),  leathery  and 
evergreen  (except,  e.g.  Cinnamomum)  ;  they  are  frequently  studded 
with  clear  glands  containing  volatile  oil.  The  flowers  are  borne  in 
panicles  and  are  small  and  of  a  greenish  or  whitish  colour.  They 
are  regular,  perigynous,  with  most  frequently  a  bowl  or  cup-shaped 
receptacle  (Fig.  386),  usually  §  ,  and  trimerous  (rarely  dimerous) 
through  all  {most  frequently  6-7)  whorls  ;  viz.  most  frequently, 
perianth  2  whorls,  stamens  3-4  and  carpels  1  (P3  +  3,  A3 +  3 +  3 
+  3,  G3)  in  regular  alternation  (Fig.  387).  Each  of  the  2  or  4 
loculi  of  the  anthers  open  by  an  upwardly  directed  valve  (Fig.  386) ; 
of  the  stamens,  the  2  outermost  whorls  are  generally  introrse, 


FIG.  386.— F:O«  er  of  the  Cinna- 
mon-tree (Cinnamomum  zeylani- 
cum)  (longitudinal  section). 


FIG.  387. — Typical  dia- 
pram  of  the  Lauraceae: 
g  staminodes. 


FIG  .  088.  —  Lauras 
iiobtlis  :  longitudinal 
section  of  fruit. 


the  others  extrorse,  or  1-3  whorls  are  developed  as  staminodes 
(Fig.  3870).  The  gynoeceum  has  1  loculus  with  1  style  and  1 
pendulous  ovule  (Fig.  386),  and  may  be  considered  as  formed  of  3 
carpels.  The  fruit  is  a  berry  (Fig.  388)  or  drupe,  which  often  is 
surrounded  at  its  base  by  the  persistent  receptacle  (as  an  acorn  by 
its  cupule),  which  becomes  fleshy  and  sometimes  coloured  during 
the  ripening  of  the  fruit.  The  embryo  has  2  thick  cotyledons,  but 
no  endosperm  (Fig.  388). 

The  Lauraceae  present  affinities  with  thePolygonacese,  in  which  there  is  found 
perigyny,  as  well  as  a  similar  number  of  parts  in  the  flower  and  a  similar 
gynoeceum,  but  with  erect  and  orthotropous  ovule.  From  their  general  charac- 
ters they  should  be  classed  among  the  Polycarpicae,  but  stand,  however,  isolated 


392 


DICOTYLEDONES. 


by  the  syncarpoas  gynceceum,  if  it  is  in  reality  formed  by  3  carpels  and  not  by  1 
only.  Hernandia,  which  has  epigynous  mono3cious  flowers,  deviates  most. — 
Cassytha  is  a  Cuscuta-like,  herbaceous,  slightly  green  parasite  with  twining, 
almost  leafless  stems.  The  flower  however  agrees  with  the  diagram  in  Fig.  387. 
Some  Lauraceee  have  curved  veins  or  palminerved  aud  lobed  leaves  (often 
together  with  entire  ones)  e.g.  Sassafras. 

'There  are  1000  species  ;  especially  in  the  forests  of  tropical  S.  America  and 
Asia,  of  which  they  form  the  principal  part.  Only  Lauras  nobilis  is  found  in 
Europe,  and  there  is  little  doubt  that  its  proper  home  is  in  Western  Asia. 


FIG.  389.— ZIyristica  :  fruit. 


Fio.  390.— Seed  with  aril  entire 
and  in  longitudinal  section. 


They  are  rare  in  Africa. — On  account  of  the  volatile  oil  found  in  all  parts  of  the 
plant,  they  are  used  as  tpices,  e.g.  the  false  Cinnamon  tree  (Dicypellium  caryo- 
pliyllatum,  in  the  Brazils).  The  OFFICINAL  ones  are — the  Cinnamon-tree  (Cinna- 
momum  zeylanicum  from  Ceylon,  E.  India,  Eastern  Asia),  which  is  also  culti- 
vated ;  the  Camphor-tree  (Cinnamomum  camphura.  Eastern  Asia).  The  Laurel- 
tree  (Lauras  nobilis,  Mediterranean),  the  berries  and  leaves  of  which  give 
laurel  oil,  is  medicinal. — Scented  wood  lor  furniture,  etc.,  is  obtained  from 
Sassafras  officinalis  (from  N.  Am.).  The  wood  ftcm  its  roots  is  officinal. 
Piclmrim  "  beans "  are  the  large  cotyledons  of  Neclandra  picJairy,  whilst  the 


RHCEADiNE.  393 

famous  "Greenheart "  wood  of  Demarara  is  the  wood  of  Nectandra  rodicei. 
The  pulp  and  seeds  contain  a  fatty  oil.  The  pear-like  fruit  of  Persea 
gratissima  (Mexico,  also  cultivated)  is  very  delicious.  Lindera  benzoin  is  a 
garden  shrub  ;  Laitrus  nobilis  likewise. 

Order  12.  Myristicaceae  (Nutmegs).  In  this  order  there  is  only  1 
genus,  Myristica.  Trees  or  shrubs.  The  leaves  agree  closely  with  those  of 
the  Lauraceee,  with  which  this  order  has  many  points  in  common.  The 
majority  of  the  species  are  aromatic,  having  in  their  vegetative  parts  pellucid 
glands  with  volatile  oils.  The  flowers  are  regular,  dioecious,  trimerous,  and 
have  a  single  gamophyllous  (cupular  or  campanulate)  3-toothed,  fleshy  perianth. 
In  the  $  -flowers  the  anthers  vary  in  number  (3-15),  and  they  are  extrorse 
and  borne  on  a  centrally-placed  column ;  in  the  ?  -flower  the  gynceceum 
is  unilocular,  unicarpellary,  with  1  ovule.  The  FKUIT  (Fig.  389)  has  tbe  form 
of  a  pear  ;  it  is  a  fleshy,  yellow  capsule,  which  opens  along  the  ventral  and 
dorsal  sutures,  exposing  the  large  seed.  This  seed  has  a  large,  red,  irregularly 
branched  aril — the  so-called  "mace";  the  "nutmeg,"  on  the  other  hand,  is 
the  seed  itself  with  the  inner  thin  portion  of  the  testa,  which  has  pushed  its  way 
irregularly  into  tbe  endosperm,  and  causes  the  marbled  appearance  of  the  cut 
seed  (Fig.  390)  ;  the  external,  dark  brown,  hard,  and  brittle  part  of  the  seed- 
shell  is  however  removed.  Mace  and  nutmeg  contain  volatile  and  fatty  oils  in 
abundance. — 80  species.  Tropical.  The  majority  are  used  on  account  of  their 
aromatic  seeds  and  aril,  the  most  important  being  M.  fragrans  (wosctiata),  from 
the  Moluccas.  This  is  cultivated  in  special  plantations,  not  only  in  its  native 
home,  but  in  other  tropical  countries  also.  Nutmegs  were  known  as  com- 
modities in  Europe  in  very  ancient  times  (e.g.  by  the  Romans),  but  it  was  not 
until  the  year  1500  that  the  tree  itself  was  known.  The  seed  is  OFFICINAL. 

Family  10.     Rhceadinse. 

The  plants  belonging  to  this  family  are  almost  exclusively  her- 
baceous, with  scattered,  exstipulate  leaves..  The  flowers  are  eucyclic 
di-  or  tetra-merous,  with  the  calyx  and  corolla  deciduous,  hypogynous, 
^  ^regular,  thegynoeceum  with  2-several  carpels  (generally  2,  trans- 
versely placed  (Figs.  391,  892,  393,  397).  The  ovary  is  unilocular 
with  parietal  placentas,  but  in  Cruciferse  and  a  few  others  it  becomes 
bilocular  by  the  development  of  a  false,  membranous  wall  between 
the  placentae.  The  stigmas  in  the  majority  of  cases  are  commis- 
sural,  i.e.  they  stand  above  the  placentae,  and  not  above  the  dorsal 
line  of  the  carpels.  The  fruit  is  Dearly  always  a  capsule,  which 
opens  by  the  middle  portions  of  the  carpels  detaching  themselves 
as  valves,  bearing  no  seed,  whilst  the  placentas  persist  as  the 
seed-bearing  frame.  Endosperm  is  found  in  Papaveracew  and 
Fumariaceie,  bat  is  absent  in  Cruciferse  and  Capparidaceds. — This 
family  through  the  Papaveraceae  is  related  to  the  Polycarpicse  (the  Nymphaaaceae), 
through  the  Capparidaceae  to  the  Eesedaceae  in  the  next  family. 

Exceptions  to  the  above  are :  EschschoUzia,  Subularia  (Fig.  403)  and  a  few 


394 


DICOTYLEDONES. 


Capparidacese,  in  which  perigynous  flowers  are  found.  A  few  Papaveracese  and 
Fumariaceae  have  trimerous  flowers.  'In  Fumaria  and  certain  Cruciferae,  the 
fruit  is  a  nut.  The  Fumariaceas  have  zygoinorphic  flowers.  Trees  and  shrubs 
are  almost  entirely  confined  to  the  Capparidaceas,  in  which  order  stipules  also 
are  found. 

Order  1.  Papaveracese  (Poppies).  Herbaceous  plants  with 
•stiff  hairs  and  latex ;  flowers  regular  (Fig.  391)  with  generally  2 
(-3)  sepals  (which  fall  off  as  the  flower  opens),  2  +  2  petals  (im- 
bricate and  crumpled  in  the  bad)  without  spur,  numerous  stamens  in 
several  alternating  whorls  (generally  a  multiple  of  2)  ;  carpels  2- 
several,  united  into  a  unilocular  gynoeceum.  Trimerous  flowers 
also  occur.  Capsule  with  very  numerous  seeds  on  the  parietal 
placentae ;  embryo  small,  with  large,  oleaginous  endosperm  (Fig. 
392).— The  leaves  have  no  stipules  and  are  generally  pinnately 
lobed. 


FIG.  391  — A  Diagram  of  the  flower  of 
Glaucium  and  tue  dichasium  (which 
•becomes  transformed  into  a  scorpioid 
•cyme).  £  Papaver  argemone,  transverse 
section  of  the  ovary  with  indication  of 
the  position  of  the  stigmas. 


FIG.  39.'.—  Papaver  somniferum :  A  cap- 
sule ;  st  the  stigma ;  v  valves ;  h  pores  ; 
B  seed  in  longitudinal  section  ;  alb  endo- 
sperm ;  emb  embryo. 


Papaver  (Poppy,)  has  large,  solitary,  terminal  flowers  ;  petals 
firmly  and  irregularly  folded  in  aestivation ;  gynceceuin  formed  by 
many  (4-15)  carpels  ;  stigmas  velvety,  sessile  and  stellate  (the  rays 
stand  above  the  placentae)  (Fig.  391  _&).  The  edges  of  the  carpels 
project  deeply  into  the  ovary,  but  do  not  meet  in  the  centre,  so 
that  it  remains  unilocular.  The  capsule  opens  by  pores  placed 
close  beneath  the  stigma,  and  formed  of  small  valves  alternating 
with  the  placentae  and  the  rays  of  the  stigma  (Fig.  391) .  P.  dubium, 
P.  nrgemone,  P.  rhueas. — Chelidonium  (Greater  Celandine)  has 
yellow  latex,  flowers  in  umbellate  cymes  (the  terminal,  central  flower 
opening  first)  and  only  2  carpels ;  the  fruit  resembles  the  siliqua 


RHCEADINJE.  395 

of  the  Cruciferse  in  having  two  barren  valves,  which  are  detached 
from  the  base  upwards,  and  a  seed-bearing  frame,  but  there  is  no 
partition  wall  formed  between  the  placentae.  Oh.  majus.—ThQ 
majority  of  the  other  genera  have,  like  Chelidonium,  2  carpels  (lateral  and  alter- 
nating with  the  sepals:  Fig.  391  A)  and  siliqua-like  fruit,  thus  :  Eschscholtzia 
(perigynous)  with  a  linear,  stigma-bearing  prolongation  extending  as  far  above 
the  placentae  as  above  the  dorsal  suture  of  the  carpels ;  Glaucium  (Horn-Poppy) ; 
G.  luteum,  whose  extremely  long,  thin  capsule  differs  from  that  of  Chelidonium 
by  the  formation,  during  ripening,  of  a  thick,  spongy  (false)  replum,  which 
persists  when  the  valves  are  detached ;  Sanguinaria  with  red  latex,  the  2  petals 
divided  into  8-12  small  petals  (perhaps  by  dedoublement) ;  Macleya  and 
Bocconia  (1- seeded  capsule)  with  2  sepals  and  no  petals. — Trimerous  flowers 
are  found  in  Argemone  and  Platystemon  (with  a  curious  fruit,  carpels  free,  and 
transversely  divided  and  constricted  into  joints  which  separate  as  nut-like  por- 
tions).— Meconopsis. — Hypecoum  (Fig.  393  C)  has  tri-lobed  and  three  cleft  petals, 
4  free  stamens  with  4-locular  anthers  and  a  jointed  siliqua;  it  presents  a  tran- 
sitional form  to  the  Fumariaceae,  with  which  order  it  is  sometimes  included. 

POLLINATION.  Paparer  and  Chelidonium  have  no  honey,  and  are  without, 
doubt  only  visited  by  insects  for  the  sake  of  the  pollen.  The  anthers  and 
stigmas  mature  about  the  same  time. — There  are  80  species ;  especially  from 
warm  climates.  OFFICINAL  :  Papaver  somniferum  (Opium-Poppy)  ;  the  latex 
of  its  unripe  capsules  is  obtained  by  incisions,  and  dried  (opium) ;  it  contains 
many  alkaloids:  morphine,  papa verine,  narcotine,  thebaine, etc.  The  oleaginous 
seeds  are  also  used  in  the  manufacture  of  oil.  Its  home  is  in  the  East,  where 
it  is  extensively  cultivated.  The  petals  of  the  Corn-poppy  (P.  rhceas)  are  also 
officinal.  Several  species  are  cultivated  as  ornamental  plants. 

Order  2.  Fumariacse  (Fumitories).  This  order  differs  from 
the  closely  allied  Papaveraceae  in  the  absence  of  latex,  a  poorer 
flower,  generally  transversely  zygomorphic  (Fig.  393  B),  in  which 
case  one  or  both  of  the  outer  lateral  petals  are  gibbous,  or  pro- 
longed into  a  spur ;  the  stamens  are  especially  anomalous.  Sepals 
2,  caducous ;  petals  2  +  2 ;  stamens  2,  tripartite  ;  each  lateral  anther 
is  bilocular  (Figs.  393  A,  B  ;  395)  ;  gynceceum  bicarpellate. 
The  fruit  is  a  nut  or  siliqua-like  capsule.  Endosperm. — Herbs  with 
scattered,  repeatedly  pinnately-divided  leaves  without  stipules, 
generally  quite  glabrous  and  glaucous  ;  the  flowers  are  arranged 
in  racemes  with  subtending  bracts,  but  the  bracteoles  are  some- 
times suppressed. 

Dicentra  (syn.  Dielytra)  and  Adlumia  have  a  doubly  symmet- 
rical flower,  with  a  spur  or  gibbous  swelling  at  the  base  of 
each  of  the  laterally-placed  petals  (Figs.  393  A,  394).  Corydalia 
has  a  zygomorphic  flower,  only  one  of  the  lateral  petals  having  a 
spur,  and  consequently  there  is  only  one  nectary  at  the  base  of  the 
bundle  of  stamens,  which  stands  right  in  front  of  the  spur  (Fig. 


396 


DICOTYLEDONES. 


393  B,  395,  396).  The  fruit  is  a  many-seeded  siliqua-like  capsule. 
A  peculiarity  of  the  flower  is  that  the  plane  of  symmetry  passes  transversely 
through  the  flowers,  whilst  in  nearly  all  other  zygomorphic  flowers  it  lies  in 
the  median  line.  Moreover,  the  flower  is  turned,  so  that  the  plane  of  symmetry 
ultimately  becomes  nearly  vertical,  and  the  spur  is  directed  backwards. — Many 
species  have  subterranean  tubers;  in  these  the  embryo  germinates  with  one. 


FiG.  393.— Diagram  of  Dtcenfro  (A),  Corydalis  (S),  and  Hypecoum  (C). 

coiyltdon,  which  is  lanceolate1  and  resembles  a  foliage-leaf.  The  tuber  is  in 
some  the  swollen  hypocotyl  (C.  card),  in  others  a  swollen  root  (C.  fabacea, 
etc.),  which  grows  down  through  the  precisely  similar  swollen  root  of  the 
mother-plant.  The  sub-genus  Cerat.ocapnos  has  dimorphic  fruits  (nuts  and 
capsules)  in  the  same  raceme.  Fumaria  differs  from  Corydalis  only  by 
its  almost  drupaceous,  one-seeded  nut  (Fig.  395). 

THE  STRUCTURE  OP  THE  FLOWER.  Hijpecoum  among  the  Papaveraceee  is  the 
•connecting  link  with  the  Fumariacese.  The  diagram  (Fig.  393  C)  corresponds 
both  in  number  and  in  the  relative  position  of  its  members  with  that  of  most 
of  the  other  Papaveraceae  (Fig.  391),  except  that  there  are  only  four  stamens 
(with  extrorse  anthers).  In  Dicentra  (Fig.  393  A),  the  two  central  (uppermost) 
stamens  are  absent,  but  each  of  the  two  lateral  ones  are  divided  into  three 
filaments,  of  which  the  central  one  bears  a  four-locular  anther,  and  each  of  the 


Fie.  394.— THcentra  spectdbilis:  A  flower  (£) ;  B  the  same,  after  removal  of  half  of  one 
outer  petal;  the  cap,  formed  by  the  inner  petals,  is  moved  away  from  the  anthers  and 
stigma;  the  insect  does  this  with  the  lower  side  of  its  abdomen,  and  thus  rubs  the  stigrna 
on  the  hairs  of  its  ventral  surface;  the  dotted  line  at  e  indicates  the  direction  of  the 
proboscis ;  C  androecium  and  gynoeceum  ;  D  stigma. 


RH(EADI]SL£. 


397 


others  a  two-locular  (half)  anther.  Corydalis  and  Fumaria  stand  alone  iu  the 
symmetry  of  the  flower,  differing  from  Dicentra  in  having  only  one  of  the  lateral 
petals  (Fig.  393  B,  sp)  prolonged  into  a  spur,  while  in  Dicentra  both  the 
petals  are  spurred.  This  structure  has  been  interpreted  in  various  ways. 
According  to  Asa  Gray  the  median  stamens  are  absent  in  the  last-named 
genera,  and  the  lateral  ones  are  split  in  a  similar  manner  to  the  petals  of 
Hiipecuurn.  Another,  and  no  doubt  the  most  reasonable  theory  (adduced  by 
De  c'andolle),  is  :  that  two  median  stamens  ate  split,  the  two  parts  move 
laterally,  each  to  their  respective  sides  and  become  united  with  the  two  lateral 
stamens ;  this  affords  a  natural  explanation  of  the  two  half-anthers,  and  estab- 
lishes a  close  relationship  to  the  Cruciferae.  A  third  interpretation,  held  by 
Eichler  and  others,  is  as  follows  :  the  median  stamens  are  always  wanting  ; 
when  they  appear  to  be  present,  as  in  Hypecoum,  it  is  due  to  the  fact  that  the 


FIG.  39-3.  —Fumaria  ojficin- 
alis :  A  the  flower  in  longi- 
tudinal section  ;  B  the  androe- 
cium  and  gynceceura  ;  nectary 
to  the  right. 


FIG.  396. — Coriidalis  cava:  a  a  flower  (lateral  viaw); 
?>  the  anthers  lying  round  the  stigma;  c  the  anthers 
shortly  before  the  opening  of  the  flower;  d  the  head 
of  the  stigma ;  e  relative  position  of  the  parts  of  the 
flower  during  the  visit  of  an  insect. 


side  portions  of  the  lateral  stamens  approach  each  other  (as  interpetiolar 
stipules)  and  coalesce  into  an  apparently  single  stamen. 

130  species  ;  mostly  from  the  northern  temperatures. 

POLLINATION.  Fumaria,  with  its  inconspicuous  flowers,  has  to  a  great 
extent  to  resort  to  self-pollination.  Corydalis,  on  the  other  hand,  is  dependent 
on  cross-pollination ;  C.  cava  is  even  absolutely  sterile  with  its  own  pollen. 
Corydalis  is  pollinated  by  insects  with  long  probosces  (humble-bees,  bees), 
which  are  able  to  reach  the  honey  secreted  in  the  spur ;  as  they  alight  on  the 
flowers  they  press  the  exterior  petals  on  one  side  (Fig.  396  e),  so  that  the 
stigma,  surrounded  by  the  anthers,  projects  forward ;  the  proboscis  is  intro- 
duced in  the  direction  of  the  arrow  in  the  figure,  and  during  this  act  the 
under-surface  of  the  insect  is  covered  with  pollen,  whioh  is  transferred  by  similar 


398  DICOTYLEDONES. 

movements  to   the  stigma  of    another   (older)   flower.  —  Ornamental  plants ; 
Dicentra  (spectabilis  and  eximia),  Adlumia,  Corydalis. 

Order  3.  Cruciferse  (Crucifers).  The  flowers  are  regular, 
5;  sepals  4,  free  (2  +  2),  deciduous;  petals  4,  free,  deciduous, 
unguiculate,  placed  diagonally  in  one  whorl,  and  alternating  with 
the  sepals  ;  stamens  6  ;  the  2  outer  are  short,  the  4  inner  (in  reality 
the  two  median  split  to  the  base)  longer,  placed  in  pairs  (tetrady- 
namia  of  Linnaeus)  ;  gynceceum  syncarpous  formed  by  2  (as  in  the 
previous  order,  lateral)  carpels,  with  2  parietal  placentae,  but 
divided  into  two  loculi  by  a  spurious  membranous  dissepiment 
(replum)  (Fig.  397).  Style  single,  with  a  capitate,  usually  two- 
lobed  stigma,  generally  commisural,  that  is,  placed  above  the 
parietal  placentas  (Fig.  397),  but  it  may  also  be  placed  above  the 
dorsal  suture,  or  remain  undivided.  Ovules  curved.  The  fruit  is 
generally  a  bivalvular  siliqua  (Fig.  398  B,  C),  the  valves  separat- 
ing from  below  upwards,  and  leaving  the 
placentae  attached  to  the  replum ;  other 
forms  of  fruits  are  described  below.  The 
oily  seeds  have  no  endosperm  (endosperm  is 
present  in  the  two  previous  orders)  ;  the 
embryo  is  curved  (Figs.  398  E,  F ;  399,  400). 
— In  general  they  are  herbaceous  plants, 
without  latex,  with  scattered,  penninerved 
leaves,  without  stipules ;  the  inflorescence 
FIG.  397.-Diag.-am  of  a  js  yerv  characteristic,  namely,  a  raceme 

Cruciferous  flower. 

with  the  flowers  aggregated  together  at  the 

time  of  flowering  into  a  corymb,  and  destitute  of  both  bracts  and 
bractt'oles. 

Many  are  biennial,  forming  in  the  first  year  a  close  leaf-rosette.  By  culti- 
vation the  tap-root  can  readily  be  induced  to  swell  out  into  the  form  of  a  tuber 
(Turnips,  Swedes,  etc.).  Stipules  are  found  indicated  by  small  glands  on  the 
very  young  leaves  ;  in  Cochlearla  armoracia  they  are  fairly  large  triangular 
scales.  Stellate  hairs  often  occur.  Floral-leaves  are  occasionally  developed. 
Terminal  flowers  are  never  found  in  the  inflorescences.  Iberis  and  Teesdalin. 
have  zypomorphic  flowers.  Subularia  (Fig.  403)  is  perigynous.  The  2  external 
sepals  (Fig.  397)  stand  in  the  median  plane ;  it  may  therefore  be  supposed  that 
there  are  two  bracteoles  outside  tbese  which,  however,  are  suppressed,  and  can 
only  in  a  few  instances  be  traced  in  the  young  flower ;  the  two  lateral  sepals 
•are  often  gibbous  at  the  base,  and  serve  as  reservoirs  for  the  nectar  secreted 
by  the  glands  placed  above  tbem  ;  they  correspond  in  position  to  the  extemal 
petals  of  the  Fumariaceas.  The  4  petals.which  follow  next  arise  simultaneously, 
:and  alternate  with  the  4  sepals;  if  it  could  be  shown  that  these  are  merely  2 
median  petals,  which  have  been  deeply  cleft  and  the  two  parts  separated  from 


EH(EADINJB. 


399 


each  other  and  displaced  to  the  diagonal   position,  there  would  be  a  perfect 
correspondence  with  the  Fumariaceous  flower ;  then  the  petaJs  would  be  followed 


FIG.  398. — Brassica  oleracea  : 


B.  C  siliqua ;  D  seed ;  E  embryo  ;  F  transverse 
section  of  seed. 


in  regular  alternation  by  the  2  lateral  small  stamens,  the  2  median  long  stamens,, 
which  it  has  been  proved  are  split  into  4  and  placed  in  couples,  and  the  2 
laterally-placed  carpels, — in  all  6  dimerous  whorls.  But  the  formation  of  the 


FIG.  399.— Transverse  section  of  seed  and 
embryo  -of  Cheiranthus  che\,ri. 


FIG.  400.— Transverse  section  of  seed  of 
Sisymbrium  alliaria. 


corolla  by  the  splitting  of  2  petals  does  not  agree  with  the  development  of  the 
flower  or  bear  comparison,  and  hence  the  only  fact  in  favour  of  this  theory  is 
W.  B.  D  D 


400  DICOTYLEDONES. 

the  otherwise  prevailing  correspondence  with  the  Fumariaceaa.  Yet  it  may  he 
observed  that  in  special  cases  each  pair  of  long  stamens  clearly  enough  arises 
from  one  protuberance  and  even  later  on  may  be  considerably  united  or  entirely 
undivided  (e.g.  Velio)  ;  in  other  instances  they  are  quite  distinct  from  the 
beginning,  and  it  is  possible  that  this  latter  condition  has  become  constant  in 
the  corolla.  Lepidium  ruderale  and  others  have  no  corolla.  Senebiera  didyma 
has  only  2  median  stamens.  Megacarpcea  has  several  stamens,  no  doubt  by 
dedoublement,  as  in  Capparidacea?. — The  number  of  carpels  may  also  be 
abnormally  increased  ;  Tetrapoma  barbareifolium  has  normally  4  carpels  with 
an  equal  number  of  placentae  and  repla.  It  is  supposed  to  be  a  variety  of 
Nasturtium  palustre. — The  2-4-8-10  greenish  glands,  which  are  found  at  the 
base  of  the  stamens,  are  nectaries,  morphologically  emergences,  and  not  rudi- 
mentary stamens.  The  forms  of  fruits  are  of  great  systematic  significance,  see 
the  genera.  In  some  species  dimorphic  fruits  are  present,  e.g.  Cardamine 
chenopodiifoU*  which  has  both  ordinary  Cardamine-siliq\iQ,a  and  1-seeded 
siliculas. 

The  curved  embryo  appears  in  five  forms,  which  have  systematic  importance  : 
1.  To  the  PLEUEOBHIZ^E  belong  those  genera  whose  radicle  (with  the  hypocotyl) 
lies  bent  upwards  along  the  edge  of  the  flat  cotyledons  (Fig.  399) ;  to  this  group 
belong  Cardamine,  Nasturtium,  Cheiranthus,  Matthiola,  Cochlearia,  Drabti, 
Iberis,  Thlaspi,  etc. ;  diagrammatic  transverse  section  :  Q— • — 2.  To  NOTO- 
RHIZ.E  belong  those  whose  radicle  lies  in  an  upward  direction  along  the  back  of 
one  of  the  flat  cotyledons  (Figs.  400,  413)  ;  e.g.  Hesperis,  Sisymbriitm,  Lepidium, 
Capsella,  Camelina:  Q  [|. — 3.  OBTHOPLOCE^E  differ  from  the  Notorhizeaa  in 
having  the  cotyledons  folded  (not  flat)  (Fig.  398  E,  f) ;  to  this  belong  Bras- 
sica,  Sinapis,  Raplianus,  Crambe,  etc. :  Q^X — 4.  SPIBOLOBE^E  :  the  radicle  lies 
as  in  the  Notorhizae,  but  the  cotyledons  are  so  rolled  together  that  a  transverse 
section  of  the  seed  cuts  them  twice  ;  Bunias  :  Q  II  II- — 5.  DIPLECOLOBEJE  :  the 
cotyledons  are  folded  forward  and  backward  so  that  a  transverse  section  cuts 
them  several  times;  Subularia,  Senebiera  :  O  II  II  II- 

On  germination  the  cotyledons  appear  above  the  ground  as 
green  leaves  ;  in  the  Orthoploceae  they  are  bilobed,  in  the'  Lepidium- 
species  divided. 

1.  Silicula,  broad  replum  (Siliculosse  latiseptse),  valves  flat 
or  slightly  vaulted,  and  the  replum  extends  through  the  greatest 
width  of  the  silicula  (Fig.  404).  The  seeds  are  situated  in  two  rows. 

O  —  :  Cochlearia  (Horse-radish)  :  the  siliqua  is  nearly  spheroid ; 
glabrous  herbs,  generally  with  fleshy,  stalked  leaves,  and  white  ' 
flowers. — Drdba  has  an  oblong,  lanceolate,  somewhat  compressed 
silicula  ;  herbs  with  small  rosettes  of  leaves,  most  frequently  with 
stellate  and  long-stalked  racemes. — Alyssum  and  Berteroa  are 
whitish,  on  account  of  the  stellate  hairs  ;  they  have  a  more  com- 
pressed and  round  or  elliptical  silicula.  Vesicaria ;  Aubrietia. 
Lunaria  (Honest}^,  Fig.  401):  very  broad  and  flat  silicula  with 
long  stalk  (the  receptacle  as  in  Capparidaceae). 


401 

O  ||  :  Camelina  (Gold -of -pleasure)  has  a  spheroid,  pear-shaped 
siliqua  with  a  small  rim  passing  right  round  (Fig.  402).  Subu- 
•laria  (Awl wort),  an  aquatic  plant  with  perigynous  flower  (Fig. 
403)  and  folded  cotyledons. 

2.  Silicula,  narrow  replum  (Siliculosse  angustisepta3),  i.e. 
the  replum.  is  much  shorter  than  the  arched,  more  or  less  boat- 
shaped  valves  (Figs.  405,  406,  407). 

O=:  Thlaspi  (Penny- Cress)  has  a  flat,  almost  circular  silicula, 
emarginate  or  cordate,  with  a  well-developed  wing  round  the  edge 
(Fig.  406).  Iberis  and  Teesdalia  •  the  racemes  during  flowering 


Fia.  401. — Lunaria  biennis. 
Fruit,  the  valves  of  which 
have  fallen  off. 


FIQ.  402. — Came- 
lina sativa.    Fruit. 


Fro.  403.— Subularia  aquatica. 
Longitudinal  section  through 
the  flower. 


are  especially  corymbose,  and  the  most  external  petals  of  the  outer 
flowers  project  radially  and  are  much  larger  than  the  other  two 
(the  flower  is  zygomorphic). — Biscutella,  Megacarpcea. 

O  ||  '  Capsella  (Shepherd's-Purse)  has  a  wingless,  obcordate  or 
triangular  silicula  (Fig.  407).  Lepidium  (Pepperwort)  has  a  few- 
(2-4)  seeded,  slightly  winged,  oval  silicula.  Senebiera  has  a  silicula 
splitting  longitudinally  into  two  nut-like  portions  ;  its  cotyledons 
are  folded. — Anastatica  hierochuntica  ("  Rose  of  Jericho  ")  is  an  annual,  sili- 
cula-fruited,  desert  plant  (Arabia,  Syria,  N.  Africa).  After  the  flowering  all  its 
then  leafless  branches  bend  together  upwards,  forming  a  kind  of  ball;  this 
spreads  out  again  on  coming  in  contact  with  water,  and  the  fruits  then  dis- 
seminate their  seeds,  which  germinate  very  quickly,  often  in  the  fruit. 


402  DICOTYLEDONES. 

3.  Siliqua  (Siliquosse).  The  fruit  is  a  true  siliqua,  several 
times  longer  than  broad.  The  seeds  in  -most  are  borne  apparently 
in  one  row. 

O^>x>  :  Brassica  (Cabbage).  The  seeds  are  placed  apparently 
in  one  row  in  each  loculus  (Fig.  398  (7) ;  the  style  is  long  and  round; 
the  valves  have  only  1  strong,  longitudinal  rib. — Melanosinapis  (M. 


FIG.  401. — Transverse  section  of  a  silicula  FIG.  405.— Transverse  section  of  ft 

with  broad  replum  :  s  replum  j  fc  the  valves.  silicala  with  narrow  replum. 

nigra,  Black-mustard)  ;  the  style  is  compressed,  two-edged;  the 
valves  of  the  siliqua  are  one  ribbed. — Sinapis  (Mustard)  ;  quad- 
rangular or  flat  style  (in  which  in  most  cases  there  is  a  seed)  and 
3-5  strong,  longitudinal  ribs  on  the  valves. — Eruca  differs  from 
Brassica  by  the  shorter  siliqua,  broad,  sword-like  "  beak "  and 
seeds  in  two  rows. 

O=  (Fig.  399) :  Cardamine  (Bitter  Cress)  has  a  long,  linear 
siliqua,  with  flat,  unribbed,  elastic  valves.  The  leaves  are  most 
frequently  pinnatifid  or  pinnate.  C.  pratensis  reproduces  by  buds 
formed  in  the  axils  of  the  leaves. — Arabis  (Rock  Cress) ;  Matthiola 
(Stock);  Cheiranthus  cheiri  (Wallflower);  Barbarea  (Winter  Cress) 


FIG.  406.— Thlaspi  arvense.  FIG.  407.— Silicula  of  Capsella  lursa-pastoris. 

(double-edged,  quadrangular  siliqua)  ;  Nasturtium  (N.  officinaley 
Water-cress) ;  the  siliqua  of  the  latter  genus  is  in  some  species 
short,  in  others  long. 

O  ||  (400)  :  Sisymbrium  (Hedge  Mustard)  the  valves  of  the 
siliqua  are  3-ribbed. — Erysimum  ;  Hesperis ;  Schizopetalum  (with 
fimbriate  petals). 


RHCEADINJE. 


403 


4.  Fruit  jointed  (Lomentaceaa).  The  fruit  is  divided  by 
transverse  walls  into  as  many  spaces  as  there  are  seeds,  and  de- 
hisces at  maturity,  generally  transversely,  into  a  corresponding 
number  of  nut-like  joints  ("  articulate-siliqua.") 

O=:  Crambe  (Kale,  Fig.  408).  The  fruit  has  only  2  joints. 
The  lower  one  resembles  a  short,  thick  stalk,  and  is  barren,  the 
upper  one  is  spherical,  and  has  1  seed. — Cakile  (C.  maritima,  Sea- 
kale)  ;  the  lower  node  is  triangular,  1-locular,  the  upper  one  more 
ensiform,  1-locular  (Fig.  409). 

O  ^>^>  :  Raphanus  has  a  long  siliqua,  which,  in  the  garden 
Radish  (E.  sativus'),  is  spongy  and  slightly  abstricted  (Fig.  411),  but 


FIG.  408.— Fruit  of 
Crambe  maritima. 


FIG.  109.— Caldle        FiG.410.--Kap7iemus        FIG  411.—  Raplianus 
maritima.   Fruit  (f).  raphanistrum.  sativus. 


neither  opens  nor  divides  transversely  (a  kind  of  dry  berry),  and 
which  in  the  Wild  Radish  (R.  raphanistrum)  (Fig.  410)  is  abstricted 
in  the  form  of  a  string  of  pearls,  and  separates  into  many  joints. 
R.  sativus  ;  the  "  Kadish  "  is  formed  by  the  bypocotyl,  after  the  bursting  of 
its  external,  cortical  portions  (of  which  there  are  generally  two  patches  at  the 
top  of  the  Kadish). 

5.  Siliqua  indehiscent  (Nueumentaceee) .  The  fruit  is  a 
short,  unjointed,  unilocular  and  1-seeded  nut,  and  the  fruit-stalks 
are  often  long,  slender,  and  drooping.  (Sometimes  a  thin  endo- 
sperm is  present). — Isatis  (Woad)  has  most  frequently  an  oblong, 
small-winged  nut;  O  ||  (Figs.  412,  413). — Bunias ;  Neslia. 


404 


PICOTYLEDONES. 


[The  systematic  division  of  this  order  given  above  is  founded  upon  that  of 
A.  P.  de  Candolle.  Prantl  (Engler  and  Prantl,  Nat.  Fam.),  1891,  adopts  a  some- 
what different  system,  which  may  briefly  be  summarised  as  follows  :  — 

A.  Hairs  unbranched  or  absent;  no  glandular  hairs. 

1.  THELYPODIE^E.     Stigma  equally  developed  on  all  sides  ;  style  undivided 

or  prolonged  above  the   middle  of  the   carpels,   or   turned   back.  — 
Stanley-itue  ;  Heliophilince. 

2.  SINAPE^E.     Stigma  strongly  developed  above  the  placenta  ;  style  beaked 

or  two-lobed. 

a.  Cotyledons  arising  behind  the  bend  of  the  embryo.  —  Lepidiince. 

b.  Cotyledons  arising  at  the  bend  of  the  embryo. 

a.  Only  lateral   nectaries.     Generally   a   silicula   or  indehiscent 

fruit.  —  Cochleariince. 
/3.  Generally  a   siliqua,  more   rarely  a   silicula   or   transversely- 

divided  or  indehiscent  fruit.      Nectaries  generally  lateral  and 

median.  —  Alliariince  ;     Sisymbriince  ;     Vellina;     Brassicina  ; 

Cardaminince. 

B.  Hairs  collectively  or  partially  branched,   very  rarely    entirely  absent  ; 
glandular  hairs  are  sometimes  also  present. 

1.  SCHIZOPETALE^;. 

2.  HESPERIDE^E.     Stigma  strongly  developed  above  the  placenta;  style  un- 

divided or  prolonged  above  the  placenta  into  shorter  or  longer  lobes. 

a.  Surface  cells  of  the  replum,  not  divided  diagonally.  —  Capsellince  ; 
Turritince;  Erysimince  ;  AlyssiiKZ. 

b.  Surface  cells  of  the  replum  divided  diagonally.  —  Malcolmiinee  ; 
Hesperidince  ;  Morieandiince.] 

POLLINATION.  Honey  is  secreted  by  the  nectaries  mentioned  above  ;  but  the 
position  of  the  stamens  is  not  always  the  most  favourable  for  pollination  by 

insects  (in  these  flowera  the  honey-seeking 
insect  must  touch  the  anthers  with  one  of  its 
sides  and  the  stigma  with  the  other),  and 
self-fertilisation  is  common.  In  some  species 
(Cardamine  praiensis)  the  long  stamens  turn 
their  anthers  outwards  towards  the  small 
ptamens,  so  that  3  anthers  surround  each  of 
the  two  large  entrances  to  the  nectaries. 

1200  species  (180  genera),  especially  in 
the  cold  and  temperate  parts  of  the  Old 
World  (Europe,  W.  Asia).  Many  are  ivceds 
in  this  country,  e.g.  Wild  Cabbage  (Brassica 
campestris),  Charlock  (Sinapis  aru<jns/s),Wild 
Badish  (Raphanus  raphaniatrum)  and  others. 
—  The  order  is  acrid  and  oleaginous.  Oil  is 
obtained  from  many  of  the  oil-containing 
seeds,  especially  of  the  Rape  (Bnmsica  nupus), 
Summer-Kape  (the  oil-yielding  cultivated  form 
°£  lhe  ^-Cabbage)  and  Ca,nclma.  Severa! 
tion  (Fig.  413).  (Mag.)  are  pot-herbs  or  fodder  plants,  e.g.  Cabbage 


413- 


405 


(Brasaica  oleracea)  with  its  numerous  varieties  :  Cauliflower  (var.  botry  tis  ;  the 
entire  inflorescence  is  abnormally  branched  and  fleshy),  Kohlrabi  (var.  gongy- 
lodes,  with  swollen,  tuberous  stem),  Kale,  Ked-Cabbage,  White-Cabbage,  etc. ; 
B.  campestris,  var.  rapifera  (Turnip) ;  B.  napus,  var.  rapifera  (Swede) ;  Rapha- 
nus  sativus  (Radish  from  W.  Asia),  R.  caudatus  (long  Radish) ;  Nasturtium 
officinale  (Water  Cress),  Lepidium  sativum  (Garden  Cress),  and  Barbarea  prcecox 
(Early  Cress) ;  Crambe  maritima  (Sea  Kale).  The  seeds  of  the  following  are 
especially  used  as  spices  :  (the  flour  of)  Melanosinapis  (Black-mustard),  and 
Sinapis  alba  (White-mustard),  which  are  officinal  like  the  root  of  Cochlearia 
armoracia  (Horse-radish,  E.  Eur).  The  herbaceous  parts  of  Coclilearia  officinalis 
and  danica  are  medicinal. — A  blue  dye  (woad)  is  extracted  from  Isatis. — Orna- 
mental plants:  Cheiranthus  cheiri  (Wallflower),  Matthiola  (Stock),  Iberis, 
Hesperi*,  Limaria,  and  others  (especially  from  S.  Eur.).  Sweet-scented  flowers 
are  rare. 


FIG.  414.— Gynandropsis  FIG.  415.— Cappavis  spinosa. 

pentaphylla. 

Order  4.  Capparidaceae  (Capers).  The  relationship  with  the  Cruciferae 
is  so  close  that  certain  forms  are  with  difficulty  distinguished  from  them.  The 
diagram  of  the  flower  is  the  same  in  the  number  and  position  of  its  parts,  but 
it  differs  in  the  modifications  which  occur  in  the  development  of  the  stamens. 
In  some  genera  all  4  stamens  are  undivided ;  in  others  both  the  2  median  ones 
are  divided  as  in  the  Cruciferas  (6  stamens,  but  not  tetradynamous)  (Fig.  414)  ; 
in  other  genera  only  1  of  these ;  in  other  instances  again  they  are  divided  into 
more  than  2  ;  and  finally  the  2  lateral  ones  also  may  be  found  divided,  so  that 
indefinite  stamens  occur  (Fig.  415).  The  bi-carpellate  gyno3ceum  is  unilocular 
(without  roplum),  but  more  than  2  carpels  may  occur.  The  ovary  is  elevated 


406  DICOTYLEDONES. 

on  a  stalk  (sometimes  as  much  as  1  foot  in  length)  ;  also  between  the  stamens 
and  corolla  a  similar  stalk  may  be  found  (Fig.  414).  The  fruit  is  long  and 
siliquose  (Cleome,  Polanisia,  Gynandropsis),  or  a  berry  (Capparis).  Endos- 
perm absent.  Some  have  zygomorphic  flowers.  Gamosepalous  calyx  and 
perigynous  flowers  also  occur. — 350  species ;  especially  in  the  Tropics.  The 
majority  are  trees  and  shrubs,  and  they  differ  also  from  the  Cruciferas  in  having 
distinct  stipules  present  in  some  species. 

"  Capers  "  are  the  flower  buds  of  the  climbing,  thorny  shrub,  Coppans  spinosa 
(Fig.  415),  which  grows  in  the  Mediterranean. 

Family  11.     Cistiflorae. 

The  flowers  in  this  family  are  perfect,  regular  (except  Resedacese, 
Violacese),  hypogynous,  the  perianth-leaves  free  (a  few  have  them 
slightly  united),  aestivation  most  frequently  imbricate  ;  they  are 
eucyclic  in  the  androecium,  and  most  frequently  in  the  other  parts, 
and  generally  5-merous  with  So,  P5,  A5  +  5,  Gr3,  but  other  numbers 
also  occur  ;  several  have  indefinite  stamens,  but  the  stamens  arise 
(where  the  development  is  known)  in  centrifugal  order  and  are 
arranged,  often  very  distinctly,  in  bundles;  in  other  words,  the 
large  number  of  stamens  is  formed  by  the  splitting  of  a  small 
number  (most  frequently  5)  ;  a  true  spiral  arrangement  is  never 
found.  Grynceceum  syncarpous,  multicarpellary  (Dilleniacese  and 
a  few  Resedacese  are  apocarpous),  most  frequently  the  number  of 
carpels  is  3,  forming  a  unilocular  ovary  with  parietal  placentse, 
but  parallel  with  this,  multilocular  ovaries,  with  the  ovules  placed 
in  the  inner  angle  of  the  loculi,  are  also  found,  and  a  few  genera 
have  a  free,  centrally-placed  placenta.  The  fruit  is  most  frequently 
a  capsule.  The  dehiscence  is  never  with  a  "  replum,"  i.e.  the 
persistent  frame  of  the  placenta,  as  in  the  family  Rhceadinae.  One 
half  of  the  orders  has  endosperm  (Violacese,  Gistacese,  Droseracese, 
Bixaceee,  Ternstrosmiaceas,  etc.),  the  other  has  no  endosperm 
(Resedacese,  Hypericacese,  Elatinacete,  Tatnaricacese,  etc.)  ;  some 
have  a  curved,  the  majority  a  straight  embryo.  The  family  is 
scarcely  quite  natural;  in  the  future  the  orders  will  probably  be  arranged 
differently. 

Order  ].  Resedacese  (Mignonettes).  —  Herbs  or  small 
shrubs  with  spirally-placed  leaves  and  very  small,  gland-like 
stipules  (as  in  Cruciferae)  ;  the  §,  hypogynous  flowers  are  zygo- 
morphic, and  arranged  in  racemes  or  spikes  typically  without 
bracteoles.  The  zygomorphic  structure  is  produced  by  the  greater 
development  of  the  posterior  side  of  the  flower,  especially  the*  petals 
and  the  nectary  ("  disc,"  in  Fig.  416  d)  which  is  situated  between 


CISTIFLOK^E.  407 

the   petals   and    stamens;  in  general   there  are    5-8    free    sepals 
and  petals,  the  latter  consisting  of  a  large  scale-like  sheath  with 
a  fimbriated  blade  (see  Fig.);  stamens  numer- 
ous ;  carpels  6-2  united  together ;  ovary  uni- 
locular  with  parietal  placentae,  but  the  cavity 
of  the  ovary  is  not  closed  at  the  top.     In  Astro- 
carpus  the  gynoeceum  is  apocarpous.     The  fruit 
is  most  frequently  a  capsule  ;    the  seeds  are 
reniforrn,  without  endosperm,  and  the  embryo 
is  curved. 

This  order  connects  the  Bhceadinae  with  the   Cisti- 

a  T,     .        ,       ,        ,,.    ,    ,       ,.      T,,         ,.          ,       .,  FIG.  416.— Diagram  of 

floras.     It   is  closely  allied   to   the  Rhoeadmae    by  its  Reseda  odorata 

external  appearance,  even  by  the  smell  and  taste,  the 

parietal  placentation,  structure  of  the  seeds,  the  inflorescences,  etc.,  whilst  by 
the  irregular  flowers  and  the  disc  placed  at  the  posterior  side  of  the  flower, 
it  is  allied  to  Capparidacece,  but  differs  from  this  order  in  not  having  its 
characteristic  number  (2-4)  and  by  the  very  different  mode  of  dehiscence  of 
the  fruit,  etc.  It  differs  from  the  other  orders  of  this  family  chiefly  in  the 
fact  that  the  number  of  the  perianth-leaves  is  not  constantly  5.  In  Eeseda 
luteola.  both  the  calyx  and  corolla  appear  to  be  4-leaved,  because  the  posterior 
sepal  is  suppressed,  and  the  2  posterior  petals  are  united.  Where  there  are  10 
stamens,  they  stand  in  2  whorls,  i.e.  in  front  of  the  sepals  and  petals  ;  if  there 
are  several,  their  position  depends  upon  the  splitting.— Astrocarpus  is  remark- 
able for  its  apocarpous  fruit  and  the  position  of  the  ovules  on  the  dorsal  suture 
of  the  carpel. 

The  yellow,  flat  disc  at  the  back  of  tbe  flower  serves  as  a  nectary,  the  honey 
being  protected  by  the  lobes  of  tbe  petals.  If  pollination  by  insects  is  not 
effected,  then  self-pollination  may  take  place,  at  all  events  in  It.  odorata. 

45  species ;  the  majority  in  the  Mediterranean  and  in  Persia.  Reseda  odorata 
(from  Egypt)  is  cultivated  on  account  of  its  sweet  scent ;  R.  luteola  ("  Dyer's 
Weed  ")  yields  a  yellow  dye. 

Order  2.  Droseracese  (Sundews).  Herbs,  chiefly  living  on 
moors  or  in  water,  and  whose  leaves  are  adapted  to  catch  and 
digest  small  animals.  With  regard  to  the  flower,  they  are  closely 
allied  to  the  Violacea3,  especially  to  those  with  regular  flowers. 
Drosera  (Sundew)  has  a  long-stalked  scorpioid  cyme  with  regular, 
^ ,  hypogynous  flowers,  5-merous  as  in  Viola.  S5,  P5,  A5,  G3 
(in  a  syncarpous  gynoeceum,  with  free,  bifid  styles  and  basal  or 
parietally-placed  ovules  in  the  unilocular  ovaries).  The  capsule 
opens  also  as  in  Viola,  but,  among  other  differences,  the  styles  are 
free,  the  seeds  very  small,  and  surrounded  by  a  loosely  lying,  thin 
shell.  Droser.i  has  radical,  long-stalked  leaves  with  the  blade  (Fig.  417) 
covered  by  numerous  strong  glandular  hairs,  placed  on  the  edge  and  in  the 
middle ;  when  small  animals  are  caught  by  these  hairs,  the  latter  and  the  entire 


408 


DICOTYLEDONES. 


blade  close  slowly  over  them  dissolving  and  absorbing  all  the  digestible  matter 
as  nourishment. 

Dioncea  muscipula  (Fly-trap  ;  N.  Am.)  has  the  same  appearance  as  Drosera, 
but  the  leaves  are  constructed  as  in  Fig.  418.  The  stalk  is  flat  and  winged,  the 
blade  small,  circular,  with  powerful,  pointed  teeth  along  the  edge,  and  on  its 


FIG.  417.— Leaf-rosette  of  Drosera  rotundifolia  <nat.  size),  and  a  leaf  (magnified). 

surface  are  6  small  bristles  (A),  which  are  very  sensitive.  When  these  are 
touched  the  blade  quickly  closes,  folding  along  the  midrib  (B,  C)  and  im- 
prisoning the  irritating  object,  the  teeth  round  the  edges  fitting  like  the  teeth 
of  a  trap.  If  it  happens  to  be  an  insect  or  similar  body,  a  digestive  fluid  is 
secreted  which,  like  the  gastric  juice,  dissolves  the  digestible  portions.  Aldro- 
vandia  vesicuiosa  (Central  and  S.  Europe)  captures  small  aquatic  animals 
in  a  similar  manner  ;  it  is  a  floating,  aquatic  plant,  the  two  halves  of  its  leaves 
also  close  together  when  irritated  (Fig.  419). — Drosophyllum. 
About  110  species  ;  most  of  them  in  the  temperate  regions. 


FIG.  4'18. — Dioncea  muscipula.    Leaves  (nat.  size). 

Orders  3  and  4.  Sarraceniaceae  and  Nepenthaceae.  These  two  orders 
are  perhaps  most  closely  allied  to  the  Droseracese  and  agree  with  these,  among 
other  things,  in  the  manner  of  taking  nourishment.  Like  the  Droseraceae  they 
absorb  nitrogenous  food  from  dissolved  animal  matter  by  means  of  their  leaves, 
which  are  specially  constructed  both  to  catch,  to  retain,  and  to  digest  any  small 


animals  which  may  be  caught.  The  SARRACENIACE^;  are  North  American 
marsh-plants  (10  species)  which  have  pitcher-like  leaf-stalks,  in  the  cavity  of 
which  a  fluid  (with  properties  approaching  those  of  gastric  juice)  is  secreted,. 


FIG.  419.— AUrovandia  vesiculosa .  A  a  plant  (nat.  size).  B  Leaf 
(mag.);  the  blade  is  closed;  the  winged  stalk  is  prolonged  into 
4-6  irritable  bristles. 


and  which  bear  at  the  apex  a  small,  lid-like  blade ;  these  leaf-stalks  are  tha 
catching  and  digestive  organs. — Sarracenia,  Darlingtonia. 


FIG.  420.— Nepenthes  (reduced). 

NEPENTHACE.E  has  only  1  genus,  Nepenthes  (the  Pitcher-plant;  about  35 
species),  especially  found  in  tropical  E.  Asia;  the  majority  are  climbing  shrubs. 
The  leaf -stalks  are  twining  organs,  and  terminate  either  simply  in  a  tendril, 
or  in  addition  to  this,  with  a  pitcher- shaped  body  (which  in  some  species  maybe 


410 


DICOTYLEDONES. 


as  much  as  a  foot  in  length)  on  whose  upper  edge  a  lid-like  structure  is  found 
(Fig.  420).  In  this  pitcher,  as  among  the  Sarraceniace«,  a  fluid  is  secreted 
which  is  able  to  digest  the  animals  captured  (sometimes  rather  large)  and  which 
corresponds  in  some  degree  to  the  gastric  juice. 

Order  5.  Violacese  (Violets).  The  flowers  are  $  ,  and  gener- 
ally zygomorphic,  hypogynous,  with  S5,  P5,  A5,  Gr3  (Fig.  421). 
The  stamens  are  closely  applied  to  the  ovary,  they  have  a  very 
short  filament,  and  at  their  summit  generally  a  membranous 
appendage  formed  by  the  prolongation  of  the  connective  (Fig. 
422  g).  The  ovary  is  unilocular  with  3  parietal  placentae;  style 


FIG.  421. — Diagram  of 
Viola. 


FIG.  422.— The  large-flowered  form  of  Viola  tricolor  •.  A  the 
flower  in  median  longitudinal  section ;  B  the  gynceceum. 


undivided  (Fig.  422  B).  The  fruit  is  usually  a  3-valved  capsule, 
opening  along  the  dorsal  sutures  (Fig.  423).  Embryo  straight ; 
endosperm  fleshy  (Fig.  425). — Many  are  herbaceous  plants  (e.g. 
Viola),  but  in  the  Tropics  shrubs  are  also  found  (e.g.  Tonidium) ; 
a  few  are  lianes ;  the  leaves  are  scattered,  with  stipules,  and 
involute  in  the  bud. 

Viola.  The  sepals  are  prolonged  backwards  beyond  the  point  of 
insertion  (appendiculate)  ;  the  corolla  is  polypetalous,  descending 
imbricate,  and  zygomorphic,  its  anterior  petal  being  larger  than  the 
others  and  provided  with  a  spur  (Fig.  421).  The  2  anterior  of  the 
5  almost  sessile  stamens  are  provided  with  a  spur-like  nectary, 
which  protrudes  a  considerable  distance  into  the  petaloid  spur 
(Figs.  421,  422  n,  sp).  The  style  is  club-like,  and  bears  the 


CISTIFLOR^. 


411 


stigma  in  a  groove  on  the  anterior  side  (Fig.  422  st).  Herbs 
with  rhizomes,  or  annuals ;  flowers  solitary.  V.  odorata,  canina,  etc.,  have 
cleistogamic  flowers  which  produce  fruit  in  addition  to  the  large,  coloured 
(violet)  flowers.  The  Pansy  (V.  tricolor)  has  large  flowers  adapted  for  insect - 
pollination,  and  also  smaller,  less  conspicuous  ones  designed  for  self-pollina- 
tion. The  stigma,  as  in  Fig.  422  A,  st,  and  B,  is  situated  on  the  anterior 
side  of  the  stylar-head,  immediately  in  front  of  the  channel  leading  down  to 
the  spur  (sp) ;  below  it  is  situated  a  valve,  easily  covered  with  pollen  when  the 
proboscis  of  an  insect  is  introduced  into  the  spur,  but  which  closes  upon  its 
withdrawal ;  cross-pollination  is  thus  secured. — The  sweet-scented  V.  odorata 
is  visited  by  the  honey-bee,  which  insures  cross-pollination,  and  in  the  absence 
of  insect  visits  it  effects  self-fertilisation  by  cleistogamic  flowers.  The  con- 
spicuous but  scentless  V.  tricolor,  var.  vuJgaris,  is  less  frequently  visited  by 
insects  (humble-bees).  In  V.  silvatica  and  V.  canina  the  pollen  is  carried  on 


42i.  .  425. 

FIGS.  423-425.— Viola  tricolor. 

FIG.  423.— Capsule  after  dehiscence  (nat.  size).  FIG.  421.— External  view  of  the  seed. 

FIG.  425. — Seed  in  longitudinal  section. 

the  head  or  proboscis  of  the  honey-sucking  bee. — The  fruits  of  V.  odorata  bury 
themselves  slightly  in  the  soil.  In  the  others  the  fruits  are  raised  above  the 
ground;  the  3  boat-shaped  valves  close  together  along  the  central  line,  and 
eject  the  seeds,  one  by  one,  with  much  violence,  so  that  they  are  thrown  to  a 
great  distance. 

The  Alsodeia-group  has  regular  or  almost  regular  flowers.  Gamopetalous 
corollas  are  found  in  Paypayrolece.  Sauvagesiea  differs  the  most  by  its  regular 
corolla,  and  5-  oo  free  or  united  staminodes. 

250  species  ;  especially  in  the  Tropics. — The  Zo/u'dmra-species  are  used  as 
ipecacuanha.  A  number  of  FioZa-species  are  cultivated  as  garden  plants, 
especially  V.  odorata  (sweet-scented  Violet)  and  V.  tricolor,  which  have  a  large 
number  of  varieties. 

Order  6.  Frankeniacese.  A  small  order  with  doubtful  relationships. 
Perennial  herbs  or  shrubs  ;  beach  plants  with  nodose  stem.  Sepals  united, 
petals  free.  Unilocular  ovary,  with  3-4  parietal  placentas.  Fruit  a  capsule. 
Embryo  straight,  endospermous.  Especially  in  S.  Europe,  Africa,  on  the 
shores  of  the  Mediterranean  and  Atlantic. 

Order  7.  Tamaricaceae  (Tamarisks).  To  this  order  belong  only  Tamarix 
and  Myricaria.  They  are  shrubs  of  a  cypress-  or  heather-like  appearance,  as 


412  DICOTYLEDONES. 

the  scattered  leaves  are  very  small,  sessile,  scale-like  or  linear,  adpressed, 
entire,  and  usually  glaucous,  and  the  branches  are  slender  and  whip-like.  The 
flowers  are  borne  in  small  spikes  or  racemes,  and  are  small,  reddish  or  whitish, 
regular,  £,  hypogynous  and  polypetalous  ;  formula  So,  P5,  A5  +  0  (Tamarix, 
which  often  has  stipular  teeth  at  the  base  of  the  filaments),  or  A5  +  o  (Myricaria, 
in  which  the  stamens  ar.e  united  at  the  base) ;  the  number  4  may  appear  instead 
of  5,  but  in  either  case  there  is  usually  a  tricarpeliate  gynosceum,  which  is 
^m^locular  and  has  either  parietal  placentae  (Myricaria)  or  a  small  basal 
placenta  (Tamarix) ;  1  trifid  style,  or  3  styles.  Capsule  dehiscing  along  the 
dorsal  suture,  and  resembling  the  Willows  in  having  a  unilocular  ovary  with 
numerous  woolly  seeds ;  but  the  seed-wool  in  this  case  is  borne  on  the  chalaza, 
and  may  be  attached  to  a  long  stalk. — Some  Tamarix-species  shed  part  of  their 
branches  in  the  winter. — 40  species ;  North  Temperate,  on  the  sea-shores  or 
steppes,  especially  in  Asia.  Ornamental  shrubs :  Myricaria  gennanica,  and 
Tamarix  gallica. 

•  Order  8.  Cistaceae.  Shrubs  or  herbs,  natives  especially  of 
the  Mediterranean  region.  Flowers  generally  in  raceme-like 
scorpioid  cymes,  regular,  ^ ,  hypogynous ;  sepals  5,  free,  tivisted 
in  the  bud,  of  which  the  two  outer  are  generally  much  smaller 
than  the  others;  petals  5,  free,  twisted  in  the  bud  (in  the  direction 
opposite  to  the  sepals),  fugacious;  stamens  numerous;  gynceceum 
syncarpous,  carpels  usually  3-5,  style  simple,  ovary  unilocular,  with 
parietal  placentation  (seldom  divided  into  loculi,  with  axile  pla- 
centation).  The  ovules  are  orthotropous  in  opposition  to  some  of 
the  other  orders  of  this  family.  The  capsule  dehisces  along  the 
dorsal  sutures  ;  embyro  curved.  The  leaves  are  simple,  undivided, 
generally  opposite  and  stipulate. — They  are  Violaceas  with  regular  flowers, 
numerous  stamens,  and  curved  embryo.  The  numerous  stamens  are  in  reality 
only  one  or  two  5-merous  whorls,  divided  into  a  large  number  of  stamens ;  these 
are  formed,  therefore,  in  descending  order,  like  the  lobes  of  many  compound 
foliage-leaves. 

Helianthemum  (Rock-Rose),  has  3  carpels. — Cistus  has  5  (-10) 
carpels. 

About  70  species ;  temperate  climates,  especially  about  the  Mediterranean. 
The  resin  of  the  Cistus-species  has  been  used  medicinally  (ladanum). 

Order  9.  Bixaceae.  This  order  is  closely  allied  to  the  Cistacese  and 
Ternstroemiacea3 ;  like  these  it  has  regular,  5-merous,  hypogynous  flowers  with 
numerous  stamens,  unilocular  ovary  and  parietal  placentze ;  sometimes  uni- 
sexual flowers  ;  it  differs  in  having  anatrcpous  ovules,  in  the  aestivation  of 
the  sepals,  etc.  All  species  (about  180)  are  trees  or  shrubs,  with  scattered, 
simple  leaves,  which  usually  have  stipules,  and  are  occasionally  dotted  with 
pellucid  oil-glands. — Bixa  orellana  (Trop.  Am.)  is  the  best  known  species;  it 
has  a  2-valved  capsule;  the  seeds  are  enclosed  in  a  shiny  red,  fleshy  testa, 
which  contains  the  well-known  orange  or  yellow  dye,  annatto. 


CISTIFLORJE.  413 

Order  10.  Dilleniaceae.  Gynoeceum  usually  apocarpous,  seed  arillate.  The 
flower  has  most  frequently  S5,  P5,  and  compound  stamens  (one  or  more 
bundles) ;  sometimes  irregular.  200  species  ;  Tropical ;  woody  plants,  many 
lianes. — Dillema,  Candollea,  Pleurandra,  Davilla,  etc. 

Order  11.  Elatinaceae  (Water-worts).  About  25  species  belong  to 
this  order ;  especially  in  temperate  climates.  They  are  small,  creeping,  rooted, 
aquatic  plants,  with  opposite  or  verticillate  leaves  and  stipules.  The  flowers 
are  solitary  or  situated  in  small  dichasia  in  the  leaf-axils,  they  are  small, 
regular,  $  ,  hypogynous,  with  free  petals,  the  same  number  in  all  5  whorls 
(Sn,  Pn,  An  +  n,  Gn),  3-merous  (e.g.  Elatine  hexandra),  4-merous  (e.g.E.  hydro- 
piper),  or  5-merous  (Bergia) ;  the  corolla-stamens  are  sometimes  suppressed ; 
petals  imbricate  without  being  twisted ;  the  ovary  is  3-4-5 -locular,  with  3-4-5 
free  styles  •  the  capsule  dehisces  septicidally.  The  seeds  are  orthotropous  or 
curved,  often  transversely  ribbed,  endosperm  wanting.  The  order  is  most 
nearly  allied  to  Hypericaceffl,  whose  primitive  form  it  appears  to  represent. 


FIG.  426.  -Diagram  of  Hypericum  quadran-  FIG.  427.— Hypericum.    Flower  with 

gulum:  S  indicates  the  bud  of  the  helicoid  three  bundles  of  stamens. 

cyme  in  the  axil  of  the  bracteole  /3. 

Order  12.  Hypericacese  (St.  John's-worts).  This  order 
is  recognised  by  its  always  opposite  or  verticillate,  simple,  and 
entire,  penninerved  leaves,  without  stipules,  and  usually  dotted 
with  pellucid  glands  ;  by  the  always  J  ,  regular,  hypogynous  flowers 
in  a  cymose  inflorescence ;  the  generally  5-merous  calyx  and 
corolla,  with  sepals  and  petals  free  ;  the  stamens  3-5,  numerously 
branched  (Figs.  426,  427)  ;  and  the  gynoeceum,  3-5-carpellate, 
styles  usually  free.  The  ovary  is  3-5-locular,  or  unilocular  with 
3-5  parietal  placentae.  Fruit  a  capsule  (dehiscing  septicidally)  or 
berry.  Endosperm  absent. 

The  inflorescence  is  a  dichasiurn  or  helicoid  cyme.  The  structure  of  the 
flowers  is  the  same  as  that  of  the  foregoing  orders  :  S5 ,  P5  ;  succeeding  these 
in  some  cases  are  t',vo  5-merous  whorls  of  stamens  in  regular  alternation,  of 


414  DICOTYLEDONES. 

which  the  inner  is  epipetalous  ;  but  the  outer  whorl  is  only  represented  by  5  small 
scales  (Fig.  427),  or  is  altogether  absent  (Hypericum  calycinum,  H.  hircinum), 
and  the  inner  divided  into  numerous  stamens,  that  is,  these  5  stamens  are  so 
deeply  divided  that  5  epipetalous  groups  bearing  anthers  are  found  (as  in  the 
Cistacese) ;  in  other  cases  the  flower  becomes  3-merous  after  the  petals,  stamens 
3  +  3  following  in  regular  alternation  (Figg.  426,  427),  the  outer  whorl  of 
stamens  in  these  cases  is  also  present  as  staminodes  (Fig.  427),  or  may  be  al- 
together suppressed.  Carpels  3-5.  The  petals  are  often  twitted,  in  the  bud,  and 
are  then  oblique. 

Hypericum.  Some  species  have  a  square  stem ;  in  these  cases 
the  leaves  are  placed  opposite  the  edges.  Fruit  a  capsule. — Vismia 
has  a  berry. — The  flowers  of  Hypericum  have  no  honey,  and  supply  only 
pollen  ;  self-pollination  often  takes  place. 

About  240  species  ;  the  tropical  ones  being  often  shrubs  or  trees  ;  the  others 
generally  perennial  shrubs. — Hypericum,  St.  John's-wort,  contains  a  resinous, 
red  matter,  which  can  be  extracted  with  alcohol.  The  American  gamboge  is 
the  dried  sap  of  species  of  Vismia. 

Order  13.  Guttiferse,  or  Clusiaceae.  Closely  allied  to  the  Hypericaceae 
and  Ternstroemiacea}.  Leaves  opposite  or  verticillate.  The  flowers  are  often 
unisexual ;  stamens  united ;  the  gynoeceum  has  most  frequently  a  sessile, 
radiating  or  shield-like  stigma. 

370  species;  chiefly  in  the  Tropics  (Am.).  They  are  principally  woody  plants 
and  their  bark  contains  a  yellow  gum  resin,  "  gamboge,"  which  is  extracted 
from  Garcinia  morella  (E.  Ind.)  and  others.  Mangosteen  (Garcinia  mango- 
stana,  S.E.  Asia),  and  Mammea  americana  (W.  Ind.),  have  very  delicious  fruits. 
To  this  order  also  belong  Platonia  insignia,  Pentadesma  butyracea  (the  Butter- 
tree),  Clusia,  Calophyllum,  Cataba,  etc. 

Order  14.  Ternstrcemiaceae.  Trees  and  shrubs  with 
scattered,  simple,  and  often  more  or  less  leathery,  evergreen,  pen- 
ninerved  leaves,  without  stipules  (Fig.  428).  The  two  most 
important  genera  are  :  Camellia  and  the  closely  allied  Thea  (by 
some  authorities  these  are  united  into  one  genus).  The  flowers 
are  regular,  hypogynous,  and  situated  singly  oa  very  short  stalks. 
A  number  of  green  floral-leaves  are  placed  below  the  calyx  and 
gradually  pass  over  into  the  sepals,  and  the  leaves  (5-6)  of  the 
calyx  again  gradually  pass  over  into  the  corolla  (this  being  especi- 
ally marked  in  Camellia),  of  which  the  number  of  leaves  varies  (5,  6, 
7  and  upwards)  ;  the  calyx  and  the  corolla  are  acyclic  or  eucyclic  ; 
the  petals  are  slightly  united  at  the  base  ;  stamens  numerous  in 
many  whorls,  the  external  ones  are  arranged  in  bundles  and 
united  with  the  petals  as  in  the  Columniferse  ;  gynoeceum  syn- 
carpous  ;  styles  often  free  nearly  to  the  base ;  ovary  3-5-locular, 
ovules  numerous  in  each  loculus.  The  fruit  is  a  woody  capsule. 


CISTIFLORS:. 


415 


— Other  genera  show  more  distinctly  than  these  the  same  structure  as  in  the 
preceding  orders,  namely  :  S5,  P5,  A5  +  5,  of  which  the  calyx-stamens  are  often 
suppressed,  and  the  petal-stamens  divided  into  numerous  stamens. — Kielmeyera 
(S.  Am.) 

260  species  ;  especially  in  the  Tropics  (E.  Asia,  Am.)  The  leaves  of  Thea 
chinensis  (or  Camellia  thea),  the  Tea-tree  (E.  Asia),  are  cultivated  for  the 
well-known  "tea,"  and  contain  thein  :  the  best  are  the  young,  still  hairy 
leaves,  of  greyish  colour;  there  are  many  varieties.  Ornamental  plants, 
Camellia  japonica  and  Actinidia. 


FIG.  428.— Thea  chinensis  (reduced). 

Closely  allied  to  this  order  are  :  Order  15.  Rhizoboleae  (with  enormously 
large  hypocotyl — hence  the  name),  and  Order  16.  Marcgraviaceae  (partly 
epiphytes,  with  dimorphic  leaves  and  cup-  or  helmet-like,  coloured,  honey- 
secreting  floral-leaves,  which  serve  to  attract  insects). 

Order  17.  Dipterocarpaceae.  This  order  has  taken  its  name  from  the  large 
wings  attached  to  the  fruits  in  Dipterocarpus  (the  wings  being  largely  developed 
sepals) ;  trees  and  shrubs  from  Trop.  Asia.  180  species.  Camphor  ready 
prepared  is  found  in  the  stem  of  Dryobalanops  camphor  a.  Hopea  ;  Valeria. 


W.  B. 


EE 


-116 


DICOTYLEDONES. 


Family  12.     Gruinales. 

The  flowers  are  hypogynous,  g  ,  polypetalous,  usually  regular 
(except  Pelargonium,  Trop&eolaceze,  BalsaminacedB)  and  throughout 
5-merous  :  S5,  P5,  A5  +  5,  or  5  +  0,  G5  (epipetalous) .  The  stamens 
soon  fall  off  and  are  oMiplostemonous,  often  united  at  the  base 
(monadelphous)  ;  the  corolla- stamens  are  in  some  completely  sup- 
pressed (e.g.  Balsaminacese,  Fig.  438),  in  others  reduced  to  teeth 
(Linum,  Fig.  431;  Erodium),  The  Tropseolacede  have  3  carpels 
and  only  8  stamens  (Fig.  437).  Ring-like  nectaries  are  not  pre- 
sent, but  at  most  only  glandular  bodies,  borne  outside  the  base  of 
the  stamens.  Ovaries  many-locular.  The  ovules  as  a  rule  are 
pendulous,  with  the  micropyle  directed  outwards  (Fig.  431,  5), 
and  the  radicle  therefore  also  points  outwards.  Usually  herbs. 
Related  to  the  Columniferce. 

Order  1.  Oxalidacese.  Most  of  the  species  are  herbs  with 
rhizomes ;  the  leaves  are  stalked,  compound,  with  entire  leaflets 
which  are  folded  and  bent  backwards  in 
the  bud  (and  in  the  sleep  position),  ex- 
stipulate  ;  some  species  have  sensitive 
leaves.  The  flowers  (Fig.  429)  are  re- 
gular, and  have  S5,  P5,  which  are  twisted 
to  the  left  or  right  in  aestivation,  A5  +  5, 
all  united  at  the  base  (monadelphous), 
gynceceum  5-car  pell  ate,  styles  5  free,  stig- 
mas capitate,  ovary  5-locular,  ovules 
numerous.  The  fruit  is  a  capsule  opening 
with  clefts  on  the  dorsal  sutures  through 
which  the  seeds  are  ejected,  while  the 

fleshy,  external  layer  of  the  testa  springs  off  elastically.     Embryo 
straight.     Endosperm. 

Oxalis  (Wood-Sorrel).  Leaves  digitate.  Species  also  occur  with  phyllodia,  i.e. 
leaf-like  petioles  placed  vertically  without  lamina  ;  a  few  have  pinnate  leaves. 
The  flowers  are  situated  singly  or  in  dichasia,  and  unipared  scorpioid  cymes. 
The  pollination  is  effected  by  insects.  Some  species  are  trimorphic  (long-, 
short-,  medium-styled  flowers)  and  some,  e.g.  O.  acetosella,  have  cleistogamic 
flowers  in  addition  to  the  ordinary  ones.  Glands  are  found  on  the  outer  side  of 
the  corolla-stamens  or  of  all  the  stamens.  0.  tetraphylla  and  others  have 
adventitious  edible  roots,  resembling  tap-roots. — Averrhoa  is  a  tropical  tree,  with 
berries  and  pinnate  leaves. 

235  species  (205  belong  to  Oxalis)',  chiefly  in  S.  Africa  and  Trop.  America. 
— Oxalate  of  potash  is  contained  in  the  leaves  of  Oxalis. 


FIG.  429.— Diagram  of  Oxalis 
acetosella. 


GRUINALES. 


417 


Order  2.  Linaceae.  Herbs  with  scattered  or  opposite,  sessile, 
simple,  small,  entire  leaves,  without  (rarely  with  small)  stipules. 
The  flowers  (Fig.  430)  are  regular,  5-  or  4-merous.  Petals  are 
free,  twisted,  quickly  falling  off.  Stamens  united  at  the  base ;  the 
petal-stamens  are  either  reduced  to  teeth  (Fig.  431  A,  in)  or  entirely 
suppressed.  Styles  free.  The  (5-4)  epipetalous  loculi  of  the 
ovary  are  incompletely  halved  by  false  divisional  walls,  each  half 


431. 


430. 


432. 
FIG.  430-433. — Linum  usitatissimum. 


433. 


Fio.  430.— The  Flax  plant. 

FIG.  431.— A  Flower  after  removal  of  sepals  and  petals;  m  petal-stamens  reduced  to 
teeth.*'  B  Longitudinal  section  of  ovary.  C  Transverse  section  of  capsule. 

FIG.  432.— Capsule  (nat.  r>ize). 

FIG.  433. — Transverse  and  longitudinal  section  of  seed  :  bl  the  cotyledons  j  k  the 
plumule;  R  the  radicle  ;  fr  the  endosperm  ;  si:  the  testa. 

contains  one  ovule  (Fig.  431  C).  The  fruit  is  a  spherical 
capsule,  dehiscing  along  the  divisional  wall  (Fig.  432)  ;  the  10  (-8) 
seeds  kave  a  straight  embryo  and  very  slight  endosperm  (Fig. 
433). 


418  DICOTYLEDONES. 

Linum  (Flax)  has  5-merous  flowers.  The  main  axis  terminates  in 
a  flower;  and  the  succeeding  branching  is  cymose,  or  unipared  scorpioid 
branching  by  unilateral  developnieut,  and  the  flowers  in  consequence  of  the 
vigorous  sympodial  development  of  the  lateral  axis  (and  also  by  the  leaves 
being  displaced  and  pushed  aside),  assume  a  position  apparently  lateral  (i.e. 
racemose)  without  bracts;  each  brauch  of  the  sympodium  generally  has  2 
leaves.  The  testa  is  shining  and  smooth  when  dry,  but  its  external  cellular 
layer  becomes  mucilaginous  in  water. — Hadiola  has  a  4-merous  flower. 
It  is  a  small  herb  with  opposite  leaves,  and  regular,  dichasial 
branching. 

The  anthers  and  stigmas  in  L.  catharticum  and  witatinsinmm  develop 
simultaneously,  and  cross-pollination  as  well  as  self-pollination  takes  place. 
L.  grandiflorum,  perenne,  and  others,  are  dimorphic  (short-  and  long-styled). 
There  are  5  nectaries  outside  the  stamens. 

130  species ;  Linum  and  Radiola  are  native  genera. — L.  usitatissimumis  exten- 
sively cultivated  in  Europe  (especially  in  Kussia  and  Belgium),  N.  America  and 
elsewhere  (its  home  no  doubt  being  Asia),  partly  on  account  of  the  oil  (linseed 
oil)  which  is  extracted  from  the  seeds,  and  partly  on  account  of  the  bast  of  the 
stem,  which  has  very  thick-walled  cells.  The  seeds  and  oil  are  OFFICINAL.  The 
species  cultivated  in  ancient  times  was  L.  angustifolium.  Several  species  are 
cultivated  as  ornamental  plants. 

Order  3.  Geraniaceae.  The  majority  are  herbs  with  dichasial 
branching,  and  scattered  or  opposite,  stalked,  palminerved  (rarely 
penninerved)  leaves  with  small  stipules.  The  flowers  are  regular 
(except  Pelargonium)  and  5-merous,  with  10  or  5  stamens,  which 
are  slightly  united  at  the  base.  Nectaries  alternate  with  the 
corolla-stamens.  The  ovary  is  most  frequently  5-locular,  deeply 
5-grooved,  and  bears  1  well  developed  style  ("  beak  "),  which  towards- 
the  apex  divides  into  5  branches  bearing  stigmas ;  ovules  1  in  each 
loculus,  pendulous  or  ascending.  The  5  carpels  become  detached 
from  one  another  when  ripe,  and  bend  or  roll  back  (Fig.  434)  or 
become  spirally  twisted  in  the  upper  "  beak-like "  part  (Figs. 
435,  436),  whilst  a  central  column  (septal  column)  persists  ;  each 
carpel,  in  consequence,  remains  either  closed,  and  the  fruit  is  a> 
5-merous  schizocarp  whose  nut-like  lower  portion,  containing  the 
seed,  is  forced  into  the  ground,  thus  burying  the  seed  by  the 
movements  of  the  spirally-twisted,  hygroscopic  "  beak "  (Figs. 
435,  436)  ;  or  it  opens  along  the  ventral  suture,  so  that  the  seeds- 
may  fall  out,  and  it  is  then  a  5-valved  capsule,  with  septicidal  de- 
hiscence  (Fig.  434)  and  the  rolling  up  of  ten  takes  place  so  suddenly 
and  violently  that  the  seeds  are  shot  out  to  considerable  distances. 
The  embryo  is  usually  green  and  curved,  and  the  cotyledons  are 
folded  ;  endosperm  is  wanting. 


GRU1NALES. 


419 


Geranium  (Crane's-bill)  has 5  +  5  stamens,  and  a  septicidal  capsule ; 
the  carpels  most  frequently  remain  suspended  from  the  apex  of  the 
column  (Fig.  434).  The  leaves  are  most  frequently  palminerved. 
The  flowers  are  situated  solitarily  or  2  together  (2-flowered  scor- 
pioid  cyme).  —  Erodium  (Stork's-bill)  ;  inflorescence  a  many- 
flowered  unipared  scorpioid  cyme,  stamens  5  +  0  (petal-stamens  are 
wanting),  and  fruit  a  schizocarp  whose  carpels  become  detached; 
their  beaks  are  hairy  on  the  internal  surface  and  twist  themselves 
spirally  (Fig.  436).  The  umbellate  inflorescences  are  composed  of 
rnultiflowered  scorpioid  cymes.  The  leaves  are  often  penninervcd. 
-r-The  most  primitive  type  is  represented  by  Biebersteinia:  So,  P5,  A5  +  5,  G5 
(ovaries  free,  and  styles  united  above) ;  fruit  5  small  nuts.  The  most  advanced 
type  is  Pelargonium,  which  has  zygomorphic  flowers,  the  posterior  sepal  being 


FIG.  434. — Geranium  san- 
guineum.    Fruit  (j). 


Pi&.  435. — Pelargonium. 


FIG.  436.— Erodium  cicu- 
tarium,  detached  carpel. 


prolonged  into  a  spur  which  becomes  adnate  to  the  peduncle ;  the  petals  are 
unequal  in  size  ;  some  of  the  petal-stamens  are  often  wanting.  (Erodium  may 
be  slightly  zygomorphic). 

POLLINATION.  The  large-flowered  Geranium-species  are  protandrous,  e.g. 
G.  pratense  (one  whorl  of  stamens  opens  first,  and  then  the  other,  and  suc- 
ceeding .these  the  stigmas,  after  shedding  the  pollen  the  stamens  bend  out- 
wards) ;  the  small-flowered  are  also  adapted,  with  various  modifications,  for 
self-pollination.— 470  species  ;  moderately  hot  climates,  especially  S.  Africa. 
— Several  Pelargonium-species,  with  numerous  varieties,  are  ornamental  plants 
(from  S.  Africa). 

Order  4.  Tropaeolaceae.  Herbaceous,  juicy  plants  which  have 
scattered,  long-stalked,  peltate  leaves  without  stipules,  and  often 


420 


DICOTYLEDONE&. 


FIG.  437. — Diagram  of 

Tropceolum, :  sp,  spur. 

sperm    is    wanting. 


climb  by  their  sensitive  petioles.  The  flowers  are  situated 
singly  in  the  axils  of  the  foliage-leaves  on  long  stalks,  arid  are 
zygmnorpliic,  the  receptacle  under  the  posterior  sepal  being  pro- 
longed into  a  spur ;  there  are  also  differences  between  the  posterior 
and  anterior  petals,  the  2  posterior  petals  situated  on  the  border 
of  the  spur  being  perigynous,  and  the  edge 
of  the  anterior  petals  adjoining  the  claw 
fringed.  After  the  5  sepals  (which  are 
more  or  less  coloured)  and  the  5  petals, 
follow  8  stamens  (as  the  2  median  ones  are 
suppressed,  one  from  each  whorl)  and  a 
gynceceum  formed  of  3  carpels ;  in  each  of 
the  3  loculi  of  the  3-grooved  ovary  is  1 
ovule.  The  fruit  is  a  schizocarp  and  divides 
into  3  1-seeded,  drupe-like  fruitlets,  which 
do  not  (as  in  the  Geraniacese)  leave  any 
pronounced  column  between  them.  Endo- 
The  cotyledons  are  thick  and  sometimes 
slightly  coalescent.  Tubers  often  occur. 

Tropceolum. — About  40  species ;  all  from  America. 

POLLINATION. — The  spur  is  the  receptacle  for  the  nectar;  the  flowers  are 
protandrous  ;  the  anthers  open  first,  and  one  by  one  take  up  a  position  in  front 
of  the  entrance  to  the  ppur,  resuming  their  original  position  when  the  pollen  is 
shed ;  the  stigma  finally  takes  their  place  after  the  filaments  have  bent  back- 
wards.— These  plants  have  an  acrid  taste  (hence  the  name  "  Nasturtium," 
"Indian  Cress"),  on  which  account  the  flower-buds  and  young  fruits  of  2'. 
majus  are  used  as  capers.  Some  species  are  ornamental  plants. 

Order  5.  Balsaminaceae.  Herbaceous,  chiefly  annual  plants 
with  juicy,  brittle  stems,  so  transparent  that  the  vascular  bundles 
may  be  distinctly  seen.  The  leaves  are  simple,  usually  scattered, 
penninerved  and  dentate;  stipules  are  wanting,  but  sometimes 
large  glands  are  present  in  their  place  at  the  base  of  the  petioles. 
The  flowers  are  strongly  zygomorphic;  of  their  five  5-merous 
whorls  the  petal-stamens  are  suppressed  (S5,  P5,  A5  +  0,  Go); 
the  sepals  are  coloured,  the  2  anterior  ones  (Fig.  438  j,  j)  are 
very  small  or  entirely  suppressed,  the  posterior  one  is  very  large 
and  elongated  into  a  spur,  and  the  2  lateral  ones  pushed  forward  ; 
sometimes  the  weight  of  the  spur  turns  the  flower  completely 
sound,  so  that  the  posterior  leaves  assume  an  anterior  posi- 
tion -  apparently  only  3  petals,  since  the  lateral  and  the  pos- 
terior petals  become  united  in  pairs,  and  the  anterior  is  larger 


COLUMNIFER^].  421 

and  differently  shaped ;  the  5  stamens  have  very  short  and  thick 
filaments  united  at  the  base,  and  their  anthers  finally  adhere  to- 
gether and  remain  in  this  condition,  covering  over  the  gyriceceum ; 
the .  filaments  ultimately  rupture  at  the  base,  and  the  entire 
anthers  are  raised  on  the  apex  of  the  gynceceum  as  it  grows  up. 
The  gynceceum  has  a  sessile  stigma  and  a  5-locular  ovary.  The 
fruit  is  a  capsule  which,  on  maturity,  opens  suddenly  when 
irritated,  dividing  into  valves  from  the  base  upwards,  and  as  the 
5  valves  roll  up  elastically,  the  seeds  are  shot  out  on  all  sides  to 
considerable  distances  ;  a  central  column  persists  (Fig.  439).  The 
embryo  is  straight,  and  without  endosperm. 

Impatiem ;  in  Europe  only  I.  noli-rne-tanyere.     225  species  ;  especially  from 
Asia.  Several  species  have  two  kinds  of  flowers:  small,  cleistogamic,  but  fertile; 


FIG.  438.— Diagram  of  Impatiens  glanduligera.  FIG.  439.— Fruit  of  Impatiens. 

and  large,  coloured  flowers,  which  in  I.  balsamine  (ornamental  plant,  E.  Ind.) 
are  protandrous  and  pollinated  by  hive-  and  humble-bees,  as  they  suck  the  honey 
from  the  spur. 

Order  6.  Limnanthaceae.  The  flowers  are  regular  and  differ  from  all  the 
other  orders  in  the  family  by  having  the  carpels  not  in  front  of  the  petals,  but 
in  front  of  the  sepals  (which  are  valvate),  and  further,  the  loculi  are  nearly  free 
individually,  but  with  a  common  yynobastic  style  ;  the  ovules  are  ascending  and 
apotropom  (anatropous  with  ventral  raphe).  The  fruit  is  a  schizocarp*  with 
nut-like  cocci. — Limnanthes  (4  species;  N.  Am.)  perhaps  belongs  to  another 
family. 

Order  7.     Humiriaceae.     Trees  and  shrubs;  about  20  species;  Trop.  Am. 

Family  13.     Columniferae. 

The  chief  characteristics  of  the  orders  belonging  to  this  family 
are  the  ^  >  regular,  generally  5-merous,  liypogynous  flowers  with 


422  DiCOTYLEPONES. 

5-merons  calyx,  sepals  united  and  valvate  in  the  bud;  petals  5, 
free  (often  twisted  in  the  bud) ;  stamens  GO  e.g. :  10,  in  two  whorls, 
but  one  of  these  is  more  or  less  suppressed,  often  altogether  want- 
ing, or  replaced  by  5  staminodes,  while  the  other  (inner  whorl)  is 
generally  divided  more  or  less  deeply  into  a  large  number  of  anther- 
bearing  filaments.  The  filaments  too  (except  Tiliacese}  are  united 
into  a  tube,  which,  especially  in  the  Malvaceae,  forms  a  long  column 
in  the  centre  of  the  flower,  surrounding*  the  gynoeceum  (Figs.  445, 
448)  ;  in  this  case,  which  is  the  most  pronounced,  the  filaments  are 
united  into  one  bundle  \monadelplious),  in  other  instances,  poly- 
adelphous. The  number  of  carpels  varies  greatly  (2  to  about  50),  but 
they  are  nearly  always  united  and  form  a  syncarpous  multilocular 
gynoeceum. — The  vegetative  characters  also  closely  agree,  the  leaves 
are  always  scattered  and  generally  stipulate ;  all  the  green  portions 
very  often  bear  stellate  hairs,  and  the  bark  in  all  the  3  orders  is  rich 
in  tough  bast.  Mucilage  is  often  present  in  cells  or  passages. — 
This  family  is  connected  with  the  Ternstroemiacese,  from  which  it 
is  very  hard  to  draw  a  sharp  line  of  demarcation,  and  it  is  also 
allied  to  the  Cistacese  and  to  the  Gruinales. 

Order  1.  Sterculiaceae  (including  Biittneriacege).  This  is,  no 
doubt,  the  least  modified  order,  and  one  in  which  the  stamens  occur 
undivided.  Obdiplostemonous.  The  10  stamens  in  two  whorls 
are  most  frequently  united  at  the  base  into  a  short  tube,  and  have 
4<-locular,  extrorse  anthers.  The  calyx-stamens  are  nearly  always 
simple,  tooth -like  staminodes,  situated  on  the  edge  of  the  tube,  or 
are  entirely  suppressed.  The  same  relation  is  found,  for  instance, 
in  the  AmpelidaceeD  and  Rhamnaceae,  namely  5  stamens  in  front  of 
the  5  petals ;  not  infrequently  the  5  stamens  are  doubled  (Fig. 
441).  Unisexual  flowers  are  found  in  Sterculia,  Cola,  Heritiera. 
The  corolla  is  often  wanting,  or  developed  in  an  unusual  manner. 
Each  loculus  of  the  ovary  (generally  5)  always  contains  more 
than  one  ovule.  Fruit  a  capsule.  Androgynophore  often  present 
(Helicteres;  Sterculia,  etc.). 

Herpiannia,  Mahernia,  Melochia,  etc.,  have  flat  petals  with  twisted  estivation  ; 
5  undivided  stamens,  which  usually  are  but  slightly  united  at  the  base,  and 
most  frequently  without  staminodes.  Thoma»ia ;  Helicteres;  Sterculia  (free 
follicles). — Ihtobroina,  Rulingia,  Biittneria,  Commersonia,  Gnaziinia,  etc.,  have 
petals  concave  at  the  base,  and  terminating  in  a  limb  abruptly  bent  back,  and 
at  the  boundary  between  them  most  frequently  ligular  outgrowths,  as  in 
certain-  genera  of  the  Caryophyllacese  ;  stamens  5-15-  <x,  anthers  at  the  edge  of 
a  short  tube  and  5  linear  staminodes  (Fig.  441). — The  Cocoa-tree  (Theobroma, 
Fig.  440  bears  large,  reddish-yellow,  berry-like  fruits,  resembling  short  cucum- 


COLUMNIFERJI. 


423 


bers,  but  ultimately  becoming  leathery  to  woody  ;  in  each  of  the  5  loculi  are 
2  (apparently  only  1)  rows  of  horizontal,  oily  seeds,  as  large  as  almonds, 
Cotyledons  large,  thick,  and  irregularly  folded.  Endosperm  absent  (Fig.  442). 
49  genera,  with  about  750  species  ;  almost  entirely  confined  to  the  Tropics  ; 
none  in  Europe  or  in  N.  Asia.— The  seeds  of  the  Cocoa- tree  (T.  cacao,  licolor, 
glaucum,  etc.,  natives  of  Trop.  Am.,  especially  north  of  the  Equator)  are 


FIG.  410.— Theobroma  cacao.    Branch  with  flowers  and  fruits  (-J-). 


411. 


442. 


FIGS.  441-442.— Theobroma  cacao. 


FIG.  441. — Diagram  of  the  flower:  si  barren  stamens. 

FIG.  442.— J?  Seed  in  transverse  section  :  nhilum.    A  Embryo  after  the  removal  of  one 
of  the  cotyledons. 

used  for    chocolate  and   are  also  officinal  ("Cocoa-beans,"    "Cocoa-butter," 
"  Oil  of  Theobroma").     Theobromine.     Cola  acuminata,  Africa. 

Order  2.  Tiliaceae.  This  differs  from  the  other  orders  of  the 
Columniferas  chiefly  in  the  stamens  being  entirely  free  from  each 
other,  and  also  divided  into  many  filaments,  as  far  as  the  6ase,  or  at 
all  events  very  far  down,  so  that  the  flower  appears  to  have  numer- 


424 


DICOTYLEDONES. 


otis  stamens  or  to  be  slightly  polyadelphous  (Fig.  443) ;  in  addition 
to  this,  it  may  be  observed  that  the  anthers  are  4-locular  and 
introrse.  In  Liihea  the  groups  of  stamens  alternate  with  the 
petals.  In  a  few  genera  (Corchorus,  Triumfetta)  10  free  and 
single  stamens  are  found  in  2  whorls ;  but,  in  the  majority, 
groups  of  free  stamens  in  separate  bundles.  The  stamens  are 
more  or  less  united  in  Apeiba,  Liihea.  Style  simple.  Ovary  2- 

5  locular.  The  ovules  are  pen- 
dulous ;  raphe  turned  inwards. 
The  calyx  readily  falls  off ;  the 
aestivation  of  the  entirely  free 
petals  is  slightly  imbricate  (not 
twisted}. 

Tilla  (Pigs.  443,  444).  Calyx 
and  corolla  5-merous  ;  the  5 
staminal  leaves  (opposite  the 
petals)  divided  as  far  as  the  base 
into  a  large  number  of  stamens 
which  are  free  or  united  into 
groups ;  gynoeceum  with  5 
loculi  in  the  ovary  (opposite  the 
sepals)  ;  there  are  2  ovules  in 
each  loculus,  though  the  ovary 
ripens  into  a  1-seeded  nut,  which 
is  not  detached  from  the  axis  of 
the  inflorescence,  but  is  carried 
away  by  the  wind,  whirling 
round  and  round,  its  large- 
winged  bracteole  serving  as  a 

r~  h  parachute  (Fig.  443).— Only  trees, 

with  alternate,  obliquely  heart-shaped 
and  dentate  leaves;  stellate  hairs,  as 
in  the  other  Colurnniferse,  are  often 
present.  The  terminal  bud  of  the 
branch  always  fails  to  develop,  and 
the  growth  is  then  continued  sym- 
podially  by  the  uppermost  axillary 
buds.  The  INFLORESCENCE  (Figs.  443,  444)  is  a  3-7-flowered  dischasium  (Fig. 
444  t,  d,  e),  which  is  developed  in  the  axil  of  a  foliage-leaf  (Fig.  444).  The 
first  of  its  2  bracteoles  (a)  is  large,  thin,  leaf -like,  and  united  with  the  inflor- 
escence, the  lower  portion  of  which  forms  a  broad  wing,  its  so-called  "bract  "  ; 
the  second  bracteole  (ft),  on  the  other  hand,  remains  scale-like,  and  supports 
a  winter  foliage-bud  covered  with  bud-scales  which  thus  is  situated  at  the  base 


FIG.  443.  -Inflorescence  of  Tilia,  with  its 
winged  bracteole  (h);  a,  a  axis  of  the  shoot; 
the  vegetative  bud  is  seen  between  the  in- 
florescence and  the  axis  of  the  shoot ;  b 
petiole  of  foliage-leaf. 


COLUMNIFER.E. 


425 


of  the  inflorescence,  and  is  a  bud  of  the  2nd  order,  in  relation  to  the  vegetative 
shoot.  This  bud  is  always  found  beneath  the  inflorescence  on  the  branch 
placed  horizontally,  and  the  winged  bracteole  is  always  found  above  it,  a  relation 
which  is  connected  with  the  fact  that  the  2  rows  of  shoots  on  the  sides  of  a 
branch  are  antidromous  with  regard  to  each  other. — The  dichasium  itself  (Fig. 
444)  terminates  with  the  flower  (t) ;  it  has  3  floral-leaves  (c,  d,  e),  which  soon 
fall  off  ;  c  is  barren :  the  other  two  bear  flowers,  or  few  flowered  dichasia,  or 
unipared  scorpioid  cymes  (indicated  in  the  figure). — The  foliage-leaves  are 
folded  in  the  bud  upon  the  median  line  (1,  2,  3  in  Fig.  444  are  foliage-leaves 
with  their  2  stipules),  the  inner  half  is  broader  than  the  outer,  and  after  unfold- 
ing is  turned  away  from  the  mother-axis  (the  position  of  the  utw  inflorescences 
and  vegetative  buds  is  indicated  in  their  axils  on  the  figure). — The  cotyledons 
on  germination  appear  above  the  ground  as  large,  lobed  leaves. 

Of  the  other  genera  some  have  a  bell-shaped,  gamosepalous  calyx,  some  have 
no  corolla,  the  anthers  of  some  open  at  the  apex  (Aristotelia,  Elaocarpus,  etc.), 
the  majority  have  a  capsule,  some  have  berries,  or  drupes,  some  separate  into 
fruitlets,  etc. — Corchorus,  Triunifetta  (nut,  with  hooked  bristles),  Luhea, 
Apeiba,  etc.  Spannannia  is  an  African 
genus  ;  4-merous  flowers  ;  fruit  a 
warted  capsule ;  filaments  numerous 
and  sensitive  to  touch,  the  external 
ones  are  without  anthers  and  rnonili- 
form  above.  The  plant  is  covered 
with  numerous  soft  and  stellate  hairs, 
and  at  the  apex  of  the  branches  bears 
several  cyrnose  umbels. 

POLLINATION  in  Tilia  is  effected  by 
insects,  especially  bees  and  Diptera, 
which  swarm  round  the  tree  tops, 
allured  by  the  numerous  strongly- 
scented  flowers  and  the  easily  ac- 
cessible honey  (formed  in  the  hollow 
sepals).  As  the  flowers  are  pendu- 
lous, the  nectar  is  protected  from 
rain ;  and,  iu  addition,  the  inflor- 
escence is  more  or  less  concealed 
beneath  the  foliage-leaf.  Self-pollination  is  impossible,  on  account  of  pro- 
tandry. — About  470  species  (nearly  all  trees  and  shrubs)  ;  especially  in  the 
Tropics,  only  a  few  being  found  in  the  temperate,  none  in  the  polar  regions,  or 
in  high  mountainous  districts. — The  inflorescence  of  the  native  species  of  Tilia 
is  medicinal.  The  wood  is  used  for  charcoal.  — The  majority  are  used  for  timber, 
and  for  the  sake  of  the  bast  ("  Bast,"  "  Jute,"  the  bast  of  Corchorus  textilis, 
Lilhea,  and  others). 

OrderS.  Malvaceae  (Mallows).  The  plants  are  easily  re- 
cognised by  the  scattered,  simple,  palminerved,  most  frequently 
lobed,  stipulate  leaves,  folded  in  the  bud  ;  the  perfect,  regular, 
hypogynous  flowers,  with  gamosepalous ,  persistent,  5-merous  calyx 


FIG.  444. — Diagram  of  the  inflorescence  of 
Tilia  and  the  vegetative  bud  ;  the  position 
of  the  leaves  is  indicated,  and  also  the 
position  of  the  inflorescences,  which  de- 
velop from  their  axils  in  the  following 
year. 


426 


DICOTYLEDONES. 


with  valvate  aestivation ;  the  5  petals  twisted  in  the  bud  and  united 
V7ith  one  another  at  the  base,  and  by  the  5  apparently  numerous 
stamens  (Figs.  445,  448),  with  the  filaments  united  into  a  tube,  with 
reniform  bilocular  anthers  opening  by  a  crescentic  slit  (in  2  valves). 
Carpels  3 — oo  united  into  onegynceceum  ;  the  embryo  is  curved  and 
the  cotyledons  are  folded  (Figs.  447,  451)  ;  endosperm  scanty,  often 
mucilaginous. — Most  of  the  plants  belonging  to  this  order  are' 
herbs,  often  closely  studded  with  stellate  hairs.  The  leaves  are 
most  frequently  palmatifid  or  palmatisect. 

An  epicalyx  is  often  found  formed  by  floral-leaves  placed  close  beneath  the 
calyx,  in  some  3,  in  others  several.  The  median  sepal  is  posterior  in  the 
species  without  epicalyx,  often  anterior  in  those  which  have  an  epicalyx. — The 
petals  are  twisted  either  to  the  right  or  to  the  left  in  accordance  with  the  spiral  of 
the  calyx  ;  they  are  most  frequently  oblique,  as  in  the  other  plants  with  twisted 
corollas,  so  that  the  portion  covered  in  the  aestivation  is  the  most  developed. 
The  corolla  drops  off  as  a  whole,  united  with  the  staminal  tube. — Only  the  5 
petal-stamens  are  developed,  but  they  are  divided  into  a  number  of  stamens, 
placed  in  2  rows,  and  provided  only  with  half- anthers  (leaf-segments,  see  Fig. 
446;  the  sepal- stamens  are  completely  suppressed) ;  these  5  staminal  leaves 
are  then  united  into  a  tube,  frequently  5-dentate  at  the  top,  and  bearing  the 


FIG.  415. — Longitudinal  section  throu3a  the 
flower  of  Malva  silvestris. 


FIG.  446. — Diagram  of  Althcea  rosea  ; 
i  the  epicalyx. 


anthers  on  its  external  side.  The  pollen-grains  are  specially  large,  spherical 
and  spiny.  There  are  from  3  to  about  50  carpels  united  into  one  gynoeceum  and 
placed  round  the  summit  of  the  axis  which  most  frequently  projects  between 
them.  There  is  only  1  style,  which  is  generally  divided  into  as  many  stigma- 
bearing  branches  as  there  are  carpels  (Figs.  445,  448).  The  fruit  is  a  schizo- 
carp  or  capsule.  Endosperm  (Figs.  447  A,  451)  scanty,  often  mucilaginous 
round  the  embryo,  which  is  rich  in  oil. 

The  order  is  the  most  advanced  type  of  Columniferas ;  it  stands  especially 
near  to  the  Sterculiaceae,  but  is  separated  from  these  and  from  the  Tiliaceae, 
among  other  characters,  by  its  2-locular  (ultimately  1-chambered)  anthers. 

The  sub-orders  may  be  arranged  as  follows : — 


COLUMNIFEli^. 


427 


I.  Carpels  in  one  whorl. 

A.  The  fruit  a  capsule,  most  frequently  with  loculicidal 
dehiscence,  and  many  seeds  in  each  loculus. 

I.  GrOSSYPiEj;.  The  staminal-column  is  naked  at  the  apex, 
blunted,  or5-dentate. —  Gossypium  (the  Cotton  plant)  has  an  epicalyx 
of  3  large  ovate-cordate  leaves,  an  almost  entire,  low  and  compressed 
calyx.  Solitary  flowers.  Large,  most  frequently  yellow,  corollas. 
A  3-5-valved  capsule  with  many  spherical  seeds.  "  Cotton  "  is  the 
seed-hairs  developed  upon  the  entire  surface  of  the  seeds  (Fig. 
447),  and  consists  of  long,  1-cellular  hairs,  filled  with  air  (and 
therefore  white)  ;  these  are  thin-walled,  with  a  Jarge  lumen,  and 
during  drying  twist  spirally,  and  come  together  more  or  less  in 
the  form  of  bands.  They  consist  of  cellulose,  and  have  a  cuticle. 
— Hibiscus  has  several,  most  frequently  narrow,  epicalyx-leaves,  a 
distinct  5-toothed  or  5-partite  calyx. — Abutilon ;  Modiola. 


A  B 

FIG.  447. — A  Seed  of  Gossypium  with  hairs ;  B  the  same  in  longitudinal  section. 

2.  BOMBACE^;.  The  staminal  tube  is  more  or  less  deeply  cleft  into  bundles, 
sometimes  almost  to  the  base  ;  pollen  smooth,  style  simple  with  capitate,  lobed 
stigma.  Almost  all  plants  belonging  to  this  group  are  trees,  and  in  many 
instances  have  large  barrel-shaped  stems,  that  is,  swollen  in  the  centre,  and 
sometimes  covered  with  large  warts.  The  wood  is  exceptionally  light  and  soft. 
The  flowers  are  often  enormously  large,  and  have  beautiful  petals ;  in  some  they 
unfold  before  the  leaves.  The  capsule-wall  is  sometimes  closely  covered  on  its 
inner  service  with  long,  silky,  woolly  hairs,  while  the  seeds  themselves  are  gener- 
ally without  hairs.  These  hairs,  however,  on  account  of  their  brittle  nature, 
cannot  be  used  like  those  of  the  Cotton-plant.  Digitate  leaves  are  found  in  the 
Baobab-tree  (Adamonia)  from  Africa,  noted  for  its  enormously  thick,  but  short 
stem,  and  in  the  American  Silk-cotton  trees  (Bombax,  Eriodendron,  Chorisia). 
Ochroma,  CIieiro*temon,  Durio,  and  others  also  belong  to  this  group.  Durio  is 
noted  for  its  delicious  fruits,  which  have  a  most  unpleasant  smell. 

[Bombax  malabaricum  is  diplostemonous ;  the  five  sepal-stamens  repeatedly 


428 


DICOTYLEDONS. 


branch,  and  the  filaments  bear  unilocular  anthers  ;  the  five  petal-stamens  bear 
bilocular  anthers.] 

B.  Schizocarps,  with  1-seeded  fruitlets,  most  frequently  nut- 
like  and  reniform  (Figs.  449,  451). 

3.  MALVE^,  MALLOW  GROUP.  The  carpels  are  arranged  in  one 
whorl  (Fig.  449) ;  the  number  of  stylar-branches  equals  that  of 
the  carpels  ;  fruitlets  1-seeded,  reniform,  iridehiscent,  but  detach- 
ing themselves  from  one  another  and  from  the  persistent  central 
column  (Figs.  450,  451). — Malva  has  an  epi  calyx  of  3  free  leaves. 
A  flower  with  2  suppressed  bracteoles  is  situated  in  the  axil  of  the  foliage- 
leaves  ;  one  of  these  supports  a  homodromous  foliage-shoot  which  forms  a 
repetition  of  the  main  axis,  the  other  an  a,ntidromous  flower  which  continues 
the  branching  as  a  unipared  scorpioid  cyme.— Alt/leva,  Rose  Mallow,  has 
an  epicaJyx  of  6—9  leaves  united  at  the  base.— Lav atera,  Sida,  Anoda, 
Bastardia,  etc.,  have  no  epicalyx. 


418.  450.  451. 

FIGS.  448- 1 51.— Malv a  silvestris. 

FIG.  448.— The  flower  after  removal  of  the  perianth  (|).  FIG.  449.— The  fruit  (*). 

FIG.  450. — A  fruiilet  (f).  FIG.  451. — The  same  in  longitudinal  section. 

4.  URENK^;,  have  always  only  5  carpels  arranged  in   1  whorl,  with  1  ovule 
in  each  loculus,  and  the  fruit  a  schizocarp,  generally  with  nut-like  fruitlets 
provided  with  warts  and  hooks  ;  but  in  some  they  dehisce  by  2  valves  (capsule). 
They  differ  principally  from  the  other  groups  in  having  .twice  as  many  stt/htr- 
branches  as  carpels  ;  the  staminal  tube  is  naked  at  the  point,  blunt  or  5 -toothed. 
— The  genera  Urena,  Pavonia,  Malachra,   Malvaviscus  (with  berry-like  fruits) 
belong  to  this  group. 

II.  Carpels  arranged  in  a  spherical  head  in  five  groups  opposite 
to  the"  petals. 

5,  MALOPE^E,  differ  from  all  the  others  in  having  a  large  number  of  fruitlets 
arranged  irregularly  in  a  round  head,  and  separating  considerably  from  each 


COLUMNIFERj;. 


429 


other  even  before  maturity  ;  there  is,  however,  only  1  style,  divided  into  a 
corresponding  number  of  branches  (this  condition  may  be  considered  to  have 
arisen  from  the  branching  [dedoublement]  of  5  carpels).  Malope  has  3  large, 
heart-shaped  (Kitaibelia  6-9)  epicalyx-leaves,  united  at  the  base.  Palava  has  no 
epicalyx. 

POLLINATION.  The  majority  have  protandrous  flowers,  and  are  pollinated 
by  insects.  Between  the  basal  portions  of  the  5  petals,  there  are  5  nectaries, 
protected  from  the  rain  by  hairs,  e.g.  in  Malva  silvestris.  When  the  flower  first 
opens  the  numerous  anthers  occupy  the  centre  of  the  flower,  and  the  still 
undeveloped  stigmas  are  concealed  in  the  staminal  tube  ;  in  the  next  stage  the 
anthers  are  withered  and  empty,  and  the  stigtnas  protrude  and  assume  their 
places  (Fig.  452).  The  large-flowered  forms,  it  appears,  are  pollinated  only  by 
insects  ;  but  self-pollination  takes  place  in  small-flowered  forms,  as,  for  example, 
in  Malva  rotnndifolia,  in  which  the  stylar-branches,  twisting  themselves,  place 
the  stigmas  in  between  the  undeveloped  anthers. 


FIG.  452. — Anoda  hastata  :  a  the  bud  just  opened,  the  stigmas  are  concealed  by  the 
nnthers;  6  fully  opened  flower  in  tf -stage;  the  upper  stamens  are  developed  first,  and 
then  the  others  in  descending  order ;  the  stylar-branches  are  now  visible,  and  lie  bent  back 
011  the  staminal  column ;  c  all  the  stamens  project  upwards,  and  all  the  anthers  are  open, 
but  the  stylar-branches  are  still  bent  back  j  d  the  anthers  are  emptied  and  the  filaments 
shrunk  together,  but  the  styles  have  now  straightened  themselves  upwards,  and  the 
stigmas  are  in  the  receptive  condition. 

DISTKIBUTION.  800  species  (63  genera),  most  of  which  are  natives  of  the 
Tropics,  especially  America.  Althaea  and  ^some  of  the  species  of  Malva  are 
natives  of  the  temperate  regions  of  the  Old  World,  the  latter  is  also  found  in 
North  America.  Gossypium  is  tropical,  no  doubt  especially  Asiatic  (G.  herbaceum 
from  India ;  G.  arboreum  from  Upper  Egypt).  Cotton  was  introduced  into 
Greece  in  the  time  of  Herodotus,  and  was  cultivated  in  America  before  the 
arrival  of  the  Europeans. 

USES.  Pungent  and  poisonous  properties  are  entirely  wanting  ;  mucilage, 
on  the  other  hand,  is  found  in  abundance  in  all  parts  of  the  plant.  Medicinal : 
the  root  of  Althcea  officinalis,  leaves  and  flowers  of  Malva-species  (M.  silvestris 


430  DICOTYLEDONES. 

vulgaris  and  borealis)  and  Gossypium. — The  seeds  contain  a  large  quantity  of  fa  tty 
oil,  which  is  in  some  cases  extracted  (Cotton-seeds  and  others).  The  seed-hairs 
of  the  Cotton  plant  are  the  most  important  product  of  the  order.  The  cultivated 
forms  of  Cotton  belong  to  several  species  :  G.  barbadense,  herbaceum,  religiosum, 
arboreum  (Nankin),  hirsutum,  and  others.  According  to  other  botanists,  there 
are  only  3  species.  Bast  is  obtained  from  e.g.  Hibiscus  cannabinus  (Gambo-herap, 
Africa),  Paritium  tiliaceum  and  Sida  retusa.  The  fruits  of  certain  species  of 
Hibiscus  (e.g.  H.  esculentua,  from  Tropical  Africa)  are  used  in  tropical  countries 
as  a  vegetable  before  they  are  ripe. — The  colouring  matter  in  the  flowers  of 
Altlicca  rosea,  var.  nigra,  is  used  for  colouring  wines,  and  hence  is  extensively 
cultivated  in  certain  parts  of  Europe.— Ethereal  oils  and  siveet-scented  .floiver* 
are  rare ;  but  several  species  possess  a  peculiar  musk-like  odour  (Malva  moschata, 
Hibiscus  abelmoschus,  and  others). — Many  are  cultivated  as  ornamental  plant* 
on  account  of  the  large  flowers,  e.g.  Hollyhock  (A.  roaea,  etc.),  Laratera 
trimestris,  Malope  grandiflora  and  trifida,  Malca-species,  Hibiscus  rosa  ttinentit) 
syriaca ;  Sphceralcea,  etc. 

Family  14.     Tricoccae. 

The  very  large  order  Euphorbiacece  and  three  smaller  ones 
belong  to  this  family.  They  have  in  common  :  unisexual,  hypo- 
gynous,  frequently  regular  flowers,  the  perianth  most  frequently 
single,  rarely  double,  or  entirely  wanting  ;  there  is  such  a  great 
variety  in  the  structure  and  parts  of  the  flower  that  one  only  can 
be  cited  as  the  rule :  viz.  the  simple  gynceceum  composed  of  3 
carpels  forming  a  3-locular  ovary,  which  is  frequently  more  or  less 
deeply  grooved  (hence  the  name,  Tricoccce}  ;  in  the  inner  angles 
of  the  loculi  are  found  1  or  2  (never  several)  pendulous  (except 
Empetracece) ,  anatropous  ovules,  with  upward  and  outwardly  turned, 
frequently  swollen,  micropyle  (Fig.  455).  The  seed  most  frequently 
has  a  large  endosperm  and  a  straight  embryo  (Figs.  455  B,  464). — 
The  family  approaches  the  nearest  to  the  Gruinales  and  Columniferae  ;  it  may 
perhaps  be  regarded  as  an  offshoot  from  the  Sterculiaceae. 

Order  1.  Euphorbiacese.  Flowers  unisexual.  In  each  of 
the  loculi  of  the  ovary,  generally  3,  there  are  1  or  2  pendulous 
ovules  with  upward  and  outwardly  turned  micropyle.  The  placenta 
protudes  above  the  ovules  (Figs.  454,  461  13).  On  the  ripening 
of  the  capsule  the  3  carpels  separate  septicidally,  frequently  with 
great  violence,  ejecting  the  seeds  and  leaving  a  central  column. 
Endosperm  copious. — For  the  rest,  the  flowers  present  all  stages, 
from  genera  with  calyx  and  corolla,  to  those  which  are  the  most 
reduped  in  Nature,  namely  the  naked,  1-stamened  flowers  of 
Euphorbia. 

The  same  variety  which  is  found  in  the  flower  is  also  present  in 


TEICOCC2E. 


431 


the  vegetative  parts.  Some  are  herbs,  as  onr  Spurges,  others  are 
shrubs  and  trees ;  some  African  Euphorbia-species  even  resemble 
the  habit  of  a  Cactus.  Leaf -like  branches  with  rudimentary  leaves 
are  found  in  Phyllanthus  (sub-genus  Xylophylla)  (Fig.  456).  The 
leaves  are  scattered  or  opposite,  often  stipulate  ;  they  are  nearly 
always  simple.  Large,  highly- branched  cells  containing  a  great 
quantity  of  pungent  latex  are  found  in  many,  and  watery  juice  in 
others.  Glands  and  glandular  hairs  are  general. — Only  a  few 
genera  can  be  considered  in  this  book. 

As  an  example  of  the  most  perfect  flowers  (which  partly 
reproduce  the  Geraniaceous  type)  may  be  mentioned,  Croton, 
Manihot,  and  Jatropha ;  5  sepals,  5  petals,  sometimes  gamopeta- 
lous,  andrcecium  diplostemonous,  or  many-stamened,  often  mono- 
delphous. 


453. 


454 


FIGS.  453-455.— Ricinus  communis. 

FIG.  453.  —  d  -flower  (magnified).  FIG.  454.  —  $  -flower  in  longitudinal  section. 

FIG.  455.— A  seed  entire;   B  in  longitudinal  section. 

Kicinus  (Castor-oil)  (Figs.  453-455)  ;  monoecious;  the <$ -flowers,, 
situated  in  the  lower  portion  of  the  inflorescence,  have  5  perianth- 
leaves  and  a  large  number  of  branched  stamens  ;  the  $  -flower 
has  3-5  perianth-leaves  ;  3  bifid  styles.  Leaves  peltate,  palmately 
lobed.  The  seeds  (Fig.  455)  contain  an  abundance  of  fatty  oil 
and  large  aleurone  grains. — Mercurialis  (Mercury)  :  the  perianth  is 
most  frequently  3-merous  ;  in  the  <$  -flowers  9-12  stamens  ;  in  the 
$  -flowers  most  frequently  a  2-locular  gynoeceum. — Phyllanthus  : 
Pr3  +  3,  A3,  united  in  some  and  forming  a  column  in  the  centre  of 
the  flower  (Figs.  457,  458)  ;  Xylophylla  is  a  section  of  this  genus. — 
W.B.  FF 


432 


DICOTYLEDONES. 


Hura  crepitans  (Sand-box  ti-ee)  has  a  many-carpellate  gynceceum, 
which  separates  with  great  violence  when  ripe. — A.  drupe  is  found 
in  Eippomane  mancinella  (the  Mancinil-tree,  W.  Ind.). — Alchornea 
(Coelebogyne)  ilicifolia  is  well  known  on  account  of  its  "partheno- 
genesis ";  only  the  $  -plant  has  been  introduced  into  Europe,  but  it 
nevertheless  produces  seeds  capable  of  germination;  these  have 
generally  several  embryos. 

Euphorbia  (Spurge)  has  the   most  reduced  flowers,  which   are 

borne  in  a  very  complicated 
inflorescence.  Each  ^-flower 
(Fig.  460  B)  is  naked,  and  con- 
sists of  one  stamen  only  (ter- 
minal on  the  axis).  In  the 
closely  allied  genus  Anthostema, 
a  small  perianth  is  situated  at 
the  place  where,  in  Euphorbia, 
there  is  a  joint  in  the  "fila- 
ment," Fig.  461  .4).  The  ?- 
flowers  (Fig.  460)  are  naked, 
with  a  3-locular  ovary  and  3 
bifid  styles.  (Anthostema  has  a 
distinct  perianth  (Fig.  461  E)  ; 
in  a  few  Euphorbias  traces  of  a 
perianth  are  present).  In  Eu- 
phorbia the  <^-  and  $ -flowers 
are  grouped  into  flower-like  in- 
florescences termed  "  cyathia." 
T^ach  oyathium  consists  of  a 
centrally  placed  $  -flower  which 
is  first  developed,  surrounded 
by  5  groups  of  £  -flowers  (sta- 
mens) placed  in  a  zig-zag,  with 
a  centrifugal  order  of  develop- 
ment (Figs.  459,  460  B),  that 
is,  in  unipared  scorpioid  cymes ; 
these  flowers  are  surrounded  by 
an  involucre  of  5  leaves  united 
into  a,  bell-shaped  structure  (Fig.  459,  1-5)  (resembling  a  calyx)  ; 
on  its  edge  are  placed  4,  generally  crescent-like,  yellow  glands, 
one  in  each  of  the  intervals,  except  one,  between  the  lobes  of  the 
involucre '(shaded  in  Fig.  459;  see  also  Fig.  460  A).  Scale-like 


456.  458. 

FIGS.  456-458.—  Phyllanthus  (XylopTiylla) 
angustifoliua. 

FIG.  456.— Leaf-like  branch  with  flowers 
(nat.  size).  FIG.  467. —  <j'-1lower  ;  and 
FIG.  468,  ?  -flower  (mag). 


TRICOC(LE. 


433 


thin  structures  (floral-leaves  ?)are  situated  between  the  ^-flowers. 
The  ?  -flower  has  a  long  stalk,  and  finally  bends  down  on  one  side, 
namely  to  the  place  on  the  edge  of  the  involucre  where  the  gland 
is  not  developed.  These  cyathia  are  again  arranged  in  an  inflor- 
escence which  commences  as  a  3-5-rayed  umbellate  cyme  (pleio- 
chasium),  the  branches  of  which  ramify  dichasially  and  finally  as 


A  B 

FIG.  460. — Euphorbia  latinjris:  A  an  (entire)  inflorescence  (cyathiu:u) ;  B  the  same  after 
the  removal  of  the  involucre. 


FIG.  461.—  Anthostema  : 
(4)  and  ?-(B)  flowers;  p  the 
perianth;  or  the  node;  o  the 
ovule. 


FIG.  459.— Diagram  of  nn  inflorescence  (cyathium)  of 
Euphorbia  with  3  floral-leaves,  m,n,  o,  supporting  other 
cyathia  which  are  subtended  by  2  floral-leaves  (brac- 
teoles ;  m,  n).  1-5,  the  ihvolucral  leaves  in  their  order 
of  development ;  the  shaded  portions  are  the  crescentic 
•glands. 

scorpioid  cymes. — Latex,  with  peculiar-shaped  starch -grains,  is 
found  in  laticiferous  cells  (especially  in  the  Cactus-like,  leafless 
species.) 

205  genera;  more  than  3,000  species;  especially  in  the  Tropics.— Many  are 
used  on  account  of  the  oil,  and  of  the  pungent  (aperient,  poisonous,  anthelmintic, 


434 


DICOTYLEDONES. 


etc.)  properties  in  the  latex  or  the  seeds.  OFFICINAL  :  "  Cascarilla-bark "  of 
Croton  eluteria\  the  fatty  oil  of  the  seeds  of  Croton  tiglium  (Trop.  Asia) ;  "  Cas- 
tor oil "  from  Ricinus  communis  (Africa,  and  cultivated  in  all  warm  climates 
throughout  the  world);  the  glandular  hairs  of  Mallotus  philippinensis  ("  Kamala  ") ; 
this  also  yields  a  red  dye.  Gum  "Euphorbium"  is  the  hardened  (resinous) 
latex  of  the  Cactus-like  Euphorbia  resinifera  (Morocco). — NUTRITIVE  plants: 
Manihot  utilissima  and  other  species  (Maniok,  Am.).  Their  large,  farinaceous 
roots  form  a  very  important  article  of  food  in  the  Tropics  (Cassava-flour, 
Tapioca  or  Brazilian  arrow-root).  The  fresh  latex  of  the  root  in  some  species 
is  a  powerful  poison ;  but  the  poisonous  properties  are  diminished  by  roasting 
or  cooking.  Caoutchouc  is  obtained  from  Siphonia  elastica  (Trop.  S.  Am.). 
The  vegetable  tallow  of  the  Chinese  tallow-tree  (Stillingia  sebifera)  is  used  in 
large  qantities  in  soap  factories.  AD  indigo-like  dye  is  obtained  from  Crozo- 
phora  tinctoria,  and  is  also  found  in  Mercurialis  perennis.  Shellac  is  obtained 
from  Aleurites  laccifera.  ORNAMENTAL  plants  :  Acalypha,  Croton,  Dalechampia.. 
— Hippomane  is  poisonous. 

Order  2.  Buxaceae.  This  order  differs  from  the  Euphorbiacete  in  having 
the  micropyle  turned  inwards  ;  the  $  -flower  has  a  4-partite  perianth  and  4 
stamens ;  the  ?  -flower  a  6-partite  perianth  and  3  carpels.  Capsule  with, 
loculicidal  debiscence,  the  inner  layer  being  detached  elastically  from  the  outer. 
— 30  species.  Shrubs  without  latex  and  with  evergreen  leaves.—  Buxus  semper  - 
virens  (Box)  is  an  ornamental  shrub  (poisonous) ;  it  has  a  very  hard  and  valuable: 
wood  which  is  used  for  wood  engraving  and  carving. 


463.  4';4 

FIGS.  462-164.     Caliitrich}  stagnates. 

FIG.  462.—  <J -flower  with  the  2  bracteoles  and  the  solitary  stamen.        FIG.  4G3. —  ?  -flower.. 
FIG.  464. — Longitudinal  section  of  the  ripe  fruit. 

Order  3.  Callitrichaceae.  Aquatic  plants,  growing  at  the  bottom  of  shallow 
water,  with  opposite,  simple,  undivided,  entire,  exstipulate  leaves,  which  are 
generally  crowded  and  form  a  rosette  in  the  apex  of  the  branches.  The  flowers 
are  unisexual  (monoecious)  and  borne  singly  in  the  leaf-axils;  they  have  no 
perianth,  but  are  provided  with  two  delicate  bracteoles ;  the  $  -flowers  consist . 
of  only  1  terminal  stamen  (Fig.  462) ;  the  $  -flowers  of  a  bicarpellate  gynoeceum 
(Fig.  463)  which  is  originally  2-locular,  but  later  on  becomes  4-locular,  as  in  the 
case  of  the  gynoeceum  of  the  Labiata?,  by  the  formation  of  a  false  partition- 
wall  ;  in  each  loculus  there  is  1  pendulous  ovule  with  the  micropyle  turned 
outwards.  Fruit  a  ^-partite  schizocarp  (Fig.  464).  25  species.—  Callitriche. 

Order  4  (?).     Empetraceae.     4  species.     Empetrum;  E.  nigrum  (Crowberry)/ 


TEREBINTHIN^].  435 

is  a  heather-like,  moorland,  evergreen  undershrub  with  linear  leaves,  having  a 
deep  groove  closed  with  hairs,  on  the  under  side.  The  erect  ovules  show  the 
greatest  deviation  from  the  Euphorbiaceae.  Dioecious  (and  £  ) ;  S3,  P3  ;  in  the 
<?  -flower,  3  stamens ;  in  the  ?  -flower,  a  6-9-locular  ovary.  Fruit  a  drupe. 

Family  15.     Terebinthinse. 

The  diagram  of  the  flower  (Figs.  465-467)  is  the  same  as  in  the 
Gruinales,  namely  S,  P,  A2  and  Gr  in  whorls  of  5  (less  frequently 
3,  4,  6,  8),  and  the  same  modifications  also  occur  with  the  suppres- 
sion of  the  petal- stamens,  etc.  But  a  ring  or  sometimes  cup-like 
glandular  structure  (disc)  is  found  between  the  androecium  and  the 
gynceceum  (Figs.  465,  466).  The  flowers  similarly  are  regular, 
hypogynous,  £  and  polypetalous,  though  exceptions  are  found  to  all 
these  characters  :  thus,  for  example,  united  sepals  and  petals  fre- 
quently occur,  and,  in  some  orders,  unisexual  flowers  by  the  sup- 
pression of  one  sex.  In  most  cases  the  flowers  are  small,  greenish- 
yellow,  and  arranged  in  paniculate  inflorescences.  The  carpels  (most 
frequently  5)  are  free  in  a  few,  but  generally  united  into  a  rnulti- 
locular  gynceceum ;  rarely  more  than  1  or  2  ovules  in  each  loculus. 
The  gynceceum  in  the  AriacardiacesB  is  so  reduced  that  it  has  only 

1  fertile  loculus  with  1  ovule. — The  ovules  are  epitropous,  i.e.  ana- 
tropous  with  outward-turned  raphe  (except  the  Anacardiaceoe). — 
The  majority  of  the  species  are  trees  and  shrubs  with  scattered, 
often  compound  (pinnate)  leaves  without  stipules,  and  as  in  addition 
they  frequently  contain  aromatic,  especially  turpentine-like  substances, 
they  assume  a  certain  resemblance  to  the  Walnut  trees,  and  were 
formerly  classed  with  them  mainly  on  this  account.     In  a  series 
of  genera  the  volatile,  scented  oils  are  found  in  special  glands  in 
the  bark  of  the  branches  and  in  the  leaves,  in  the  latter  case 
appearing  as  pellucid  dots.     This  family  includes   several  orders 
which  are  somewhat  difficult  to  distinguish  from  each  other. 

Order  1.  Connaracese.  This  order  forms  the  connecting  link  between 
Terebinthinae  and  Rosifloras  (Spircea)  as  well  as  Leguminosae,  with  which  they 
are  sometimes  classed.  The  flowers  have  5  5-merous  whorls;  2  ovules  in  each 
loculus;  micropyle  turned  upwards.  Fruit  a  follicle,  rarely  a  collection  of 
follicles.  Seed  with  aril.  Shrubs  with  scattered  (most  frequently  pinnate) 
leaves,  without  stipules.  170  species.  Tropical. . 

Order  2.  Meliacese.  Trees  and  shrubs  with  scattered,  often  pinnate  leaves 
without  pellucid  dots  and  exstipulate  ;  the  leaflets  are  nearly  always  entire. 
Flowers  email  in  paniculate  inflorescences.  Calyx  and  corolla  4-5-merous  ; 

2  whorls  of  stamens  ;  3-5  carpels   in   the   gynoaceum.     A  Tery  characteristic 
feature  is  the  union  of  the  filaments  into  a  tube,  on  the  edge  of  which  stipule- 
like  teeth  are  often  found.     There  are  most  frequently  2  ovules  in  the  loculi ; 


4S6 


DICOTTLEDONKB. 


fruit  a  capsule  with  many  winged  seeds  in  Swletenia  (Mahogany  tree  ;  Trop. 
Am.),  Cedrela,  etc. ;  berries  iu  others.  The  wood  of  Cedrela  is  used  for  making 
cigar  boxes.  550  species  ;  tropical. 

Order  3.  Rutaceae.  Leaves  glandular  with  pellucid  dots. 
The  type  is  the  same  as  that  of  the  family.  Flowers  4-5-merous. 
The  ovary  is  most  frequently  4-5-grooved.  Disc  well  pronounced, 
often  appearing  as  a  "  gynophore."  The  majority  are  shrubs  with 
alternate  or  opposite,  compound,  more  rarely  simple,  leaves. 

A.  The  ovary  is  deeply  2-5-cleft  with  basal  styles  which  are 
more  or  less  united ;  the  carpels  in  some  genera  are  entirely  free 
(groups  1,  2).  The  fruit  is  capsular  and  most  frequently  dehisces 
like  follicles  along  the  ventral  suture  or  septicidally,  so  that  a 
horn-like  internal  layer  (endocarp)  separates  elastically  from  the 
external  layer. 

1.  ZANTHOXYLE.E.     Zanthoxylum ;  Choisya ;  Ecodia. 

2.  BORONIE^E.     Australia. — Correa. 

3.  DIOSMEJE.     Heather-like  shrubs;  Africa. — Diosma,  Coleonema,  Empleurum 
and  Barosma.     OFFICINAL  :   Baroitma  crenulata  and  betulina,    "  broad  Buchu 
leaves"  (B.  serratifolia  and  Empleurum  serrulatum,  "narrow  Buchu -leaves"). 


FIG.  465,-Rutj..    Flower  (m-ig.). 


Fie.  46P.— Euia.     Longitudinal  s<  c.ion  of  flower. 


FJG.  457.— Ruta.    Floml diagram. 


4.  RUTEJ;.  lluta  (Figs.  465-467)  graveolens  is  an  herbaceous, 
glaucous,  strongly  smelling  plant  with  bipinnate  leaves  and  yellow 
flowers ;  the  terminal  flower  is  5-merous,  the  others  4-merous 
(S.  Eur.). — Dictamnus;  zygomorphic  flower.  The  individual  carpels 


TEREBINTHIN2E. 


437 


of  the  fruit  separate  from  each  other,  and  dehisce  like  follicles,  upon  which  the 
internal  layer  is  detached  elastically  and  springs  out,  carrying  the  seeds  with  it. 
Several  species  are  ornamental  plants. 

5.  CUSPARIE^:.    American.    Flowers  often  zygomorphic  with  gamopetalous 
corolla;    stamens  5. — Ticorea  ;  Galipea  (G.  qfficinalis ;    S.  Am. ;  "Cortex  an- 
gosturas  ")  ;  Cusparia  ;  Almeidea. 

B.  The  ovary  is  entire  or  only  slightly  grooved ;  the  style  is 
terminal,  undivided.  The  fruit  is  most  frequently  a  drupe  or 
berry. 

6.  TODDALIE.E.     Ptelea ;   winged  fruit.     The  buds  are  enclosed  in  the  leaf- 
sheath.     Skimmia  :  Phellodendron. 


4(59. 


FIGS.  468-470.— Cirrus  vulgaria. 


FIG.  468. — Branch  with  compound  leaves.  FIG.  469. — Transverse  section  of  fruit. 

FIG.  470.— Flowers  (after  the  removal  of  the  petals). 

7.  AURANTIE^E,  ORANGE  GROUP.  BVuit  a  berry  with  a  leathery 
external  layer. — The  most  typical  flower  is  found  for  example  in 
Limonia:  S5,  P5,  A5  +  5,  Go  (2-5). — Citrus  has  4-5-8-merous 


438  DICOTYLEDONES. 

flowers,  a  gamosepalous,  dentate  calyx,  free  petals,  one  whorl  of 
stamens  which  are  split  irregularly  into  several  bundles  (Fig.  470). 
The  fruit  is  a  multilocular  berry  provided  with  a  thick,  tough,  outer 
layer.  The  juicy  pulp,  which  fills  up  the  loculi  and  envelopes 
the  seeds,  is  formed  from  many  large-celled,  juicy  hair-structures 
which  arise  on  the  inner  side  of  the  walls  of  the  loculi  and  by 
degrees  entirely  fill  them  up  ;  the  dissepiments  remain  thin,  and 
form  the  partitions  so  easily  separating  from  each  other  (Fig. 
469).  The  seeds  in  many  instances  are  remarkable  for  containing 
several  embryos.  The  blade  of  the  leaf  is  separated  from  the 
frequently  winged  stalk  by  a  node  (and  hence  is  a  compound  leaf 
with  only  the  terminal  leaflet  developed  ?)  (Fig.  468)  ;  in  other 
genera,  as  Triphasia,  there  is  a  fully  developed  trifoliate  leaf. 
Thorns  are  frequently  developed. — The  species  of  this  genus,  which  is 
a  native  of  the  warmer  parts  of  S.  E.  Asia,  are  very  hard  to  separate.  The 
differences  are  found  in  the  forms  of  the  fruit,  the  leaves  and  the  leaf-stalks, 
and  in  the  number  of  stamens.  Citrus  medica,  "  Cedrat "  (Ind.) ;  C.  Liinonum, 
"Citron,"  "Lemon"  (introduced  into  Italy  in  the  3rd  to  4th  century). 
OFFICINAL  :  the  fruits  and  essential  oil  of  Lemon.  C.  aurantium  from  E.  Asia, 
the  Orange  (introduced  into  Italy  in  the  14th  century).  C.  vulgaris  (Fig.  468), 
Bitter  Orange  (introduced  into  Europe  at  the  time  of  the  Crusades) ;  the  unripe 
Bitter  Oranges,  and  peel  of  the  Bitter  Orange  is  officinal ;  it  is  from  the  flowers 
of  this  species  especially  that  the  essence  of  Neroli  is  made.  C.  limetta, 
C.  bergamia,  Bergamot ;  essence  of  Bergamot  is  officinal.  C.  decumana,  Pomalo, 
a  native  of  the  Islands  of  the  Pacific.  About  780  species  ;  chiefly  tropical. 

Order  4.  Burseraceae.  Fruit  a  drupe  ;  1-5  stones.  The  bark,  as  well  as 
the  other  parts,  contain  strong  aromatic  resins  and  balsams,  and  hence 
several  species  are  used :  the  Myrrh  tree,  Commiphora  (Balsamodendron)  from 
Arabia  and  Africa;  OFFICINAL:  Myrrha  (Commipliora  my rrha).  Mecca-balsam 
from  C.  opobalsamum,  Arabia  ;  E.  Africa.  The  Incense-tree  (Boswellia)  from  the 
same  parts  of  the  globe  and  E.India.  The  incense  of  B.  carteri  is  medicinal 
(Frankincense).  The  resin  (Elemi)  of  Protium-species  is  officinal,  and  is  used 
technically  for  varnish  (S.  Am.).  Takamahaka- resin  from  Elaphrium  (S.  Am.) 
Protium  (Idea)  ;  Amyris  (1  carpel).  270  species ;  tropical. 

Order  5.  Zygophyllaceae.  The  majority  have  opposite,  pinnate  leaves  with 
stipules.  Leava:  without  pellucid  dots.  The  filaments  have  a  scale  on  the 
inner  side.  1'ht  most  important  is  Guaiacum  qfficinale  (West  India),  the  wood 
{Lignum  Vitae)  of  which  is  very  hard  and  beavy,  this  wood  and  Gum-guaiacum 
are  officinal.  Others  have  a  peculiar  repulsive  smell  and  taste  :  the  Creosote 
shrub  (Larrea  mexicana)  and  Zygoplnjllum  simplex.  Tribulus  teirester  is  a 
common  weed  in  S.  Europe.  Fagonia.  Peganum  harmala  (South  of  Russia) 
yields  a  red  dye. — 110  species  ;  especially  in  the  Tropics  ;  several  species  in 
sandy  deserts.  Nitraria. 

Order  6.  Simarubaceae.  This  order  is  distinguished  by  the  abundance  of 
Utter  substances  which  it  contains  (Quassine)  especially  in  the  bark  and  the 
•wood.  The  wood  of  Quassia  amara  (Guiana,  Antilles)  is  officinal ;  Picracna 


AESCULIN^:.  439 

excelsa  yields  Jamaica  Quassia;  the  bark  of  Simaruba,  Simaba-species  and 
others  is  used.  AOanthw  glandulosa  is  a  garden  plant  (pinnate  leaves,  winged 
fruit). — 110  species.  Tropical. 

Order  7.  Ochnacese.  Flowers  diplostemonous,  5-merous.  The  unilocular 
ovaries,  which  are  individually  free,  project  considerably  into  the  air  around  the 
gynobasic  style  ;  1  ovule  in  each  loculus ;  the  fruitlets  are  drupes.  Shrubs ; 
leaves  alternate,  with  stipules.  Ochna ;  Ouratea.  — 160  species;  tropical; 
especially  American. 

Order  8.  Anacardiaceae.  The  ovary  rarely  contains  more  than  1  ovule, 
even  though  there  be  several  loculi  and  several  carpels ;  in  Anacardinm  all  the 
10  stamens  except  one  become  suppressed.  Eesin  passages. — Anacardium.  The 
most  peculiar  feature  is  the  development  of  the  flower-stalk  into  a  fleshy  body 
about  the  form  and  size  of  a  pear  (A.  occidental  from  Trop.  Am.  and  A. 
orientate  from  E.  Ind.)  which  bears  the  kidney-shaped  nut  (the  so-called 
"  Cashew-nut ")  on  its  apex.  Mangifera  indl-a  (the  Mango-tree,  from  E. 
lad.)  is  cultivated  in  several  tropical  countries  on  account  of  its  delicious 
drupe.  Similarly,  species  of  Spmdias  (S.  dulcis,  Pacific  Islands,  S.  luted). 
Several  species  of  Ehus  are  ornamental  shrubs  in  this  country,  for  instance, 
R.  typhina  (N.  Am.),  R.  c-ttinus  (the  Wig-tree,  the  barren  flower-stalks  of  the 
panicles  being  feather-like  and  hairy) ;  R.  toxicodendron  (Poisonous  Sumach,  from 
N.  Am.)  is  poisonous.  Chinese  galls  are  produced  by  the  sting  of  a  leaf-louse 
(Aphis  chinensis)  on  R.  semialata  (China),  and  Japanese  wax  is  from  the  the 
seeds  of  R.  succedanea  (Japan).  Considerable  quantities  of  Sumach  (R.  coriaria) 
are  used  in  tanning  and  as  a  black  dye.  OFFICINAL:  the  mastic  resin  of 
Pistacia  lentiscus  (the  Mastic-tree,  from  the  Mediterranean).  The  fruits  of 
Pistacia  vera  (Syria)  are  edible ;  P.  terebinthus  and  others  yield  turpentine. — 
450  species ;  tropical. 

Order  9.  Icacinaceae.  Flowers  4-5-merous  ;  haplostemonous  ;  receptacle 
convex  or  cup-like  surrounding  the  gynoeceum ;  in  the  (single)  loculus  of  the 
ovary,  2  anatropous,  pendulous  ovules. — 200  species ;  tropical. 

Family  16.     Aesculinae. 

The  essential  characters  of  this  family  are  in  the  main  the 
same  as  those  of  the  Terebinthinae  and  Gruinales.  The  flowers  are 
hypogynous,  perfect,  with  free  petals,  5-merous  (S5,  Po,  typically 
A5  +  5,  all  of  which,  however,  are  not  generally  developed  ;  in  our 
native  orders  there  are  only  7-8  stamens),  and  most  frequently 
a  3-merous,  3-locular  gynoeceum  (less  frequently  2  or  5  carpels  with 
as  many  loculi).  In  each  loculus  there  are  usually  only  1-2 
ovules.  A  deviation  from  the  preceding  families  is  the  frequent 
zygomorphy  of  the  flower,  with,  as  a  rule  an  oblique  plane  of 
symmetry  (Fig.  471).  When  a  disc  is  developed  it  is  placed 
outside  the  stamens.  The  majority  have  no  endosperm  (Fig.  473). 
— The  members  of  the  family  are  nearly  all  trees. 

The  family  is  closely  allied  to  the  Terebinthinae,  but  unlike  this  it  never  has 


440 


DICOTYLEDONES. 


aromatic  properties,  and  differs  also  in  the  position  of  the  nectary,  in  the 
flowers,  which  are  often  irregular  with  a  reduction  in  the  number  of  stamens, 
and  in  the  ovule  which  is  usually  ascending  with  micropyle  pointing  downwards 
(the  Terebinthinfe  having  the  micropyle  turned  upwards),  etc.  It  is  also  related 
to  Frangulinae,  the  Staphyleaceae  being  the  chief  connecting  link  ;  but  the 
vEsculinze  generally  have  compound  leaves. 

Order  1.  Staphyleacese.  Leaves  opposite,  often  compound.  Flowers 
regular,  £  ,  5-merous  in  calyx  and  corolla,  5-stamened.  The  stamens  are  placed 
outside  the  nectary.  Ovary  syncarpous  or  2-3-partite  with  free  styles.  The 
capsule  is  thin,  bladder-like,  2-3-locular,  opening  at  the  apex,  and  has  several 
very  hard  seeds  with  a  shining  testa  without  aril.  Endosperm.  Staphylea 
pinnata  (^.Europe)  and  trifoliata  (N.  Am.)  are  cultivated  in  gardens;  they 
have  white  flowers  in  pendulous,  axillary  racemes  or  panicles. — 16  species. — 
Staphylea  is  found  in  the  Tertiary  of  N.  America. 

Order  2.  Melianthaceae.  Glaucous  shrubs  with  scattered,  pinnate  leaves, 
and  large  stipules.  Meliaitfhu*, — 8  species ;  S.  Africa. 

Order  3.  Sapindacese.  Trees  or  shrubs,  often  climbing  by 
tendrils  (lianes  with  anomalous  structure  of  the  stem)  and  with 
compound  leaves.  The  flowers,  in  most  cases,  are  small,  insigni- 


471.  472.  473. 

FIGS.  471-473. — JEsculus  hippocastanum. 

FIG.  471.— Diagram  of  the  flower  and  of  a  scorpioid  cyme.         FIG.  472. — Flower  in 
longitudinal  section.        BIG.  473. — Seed  in  longitudinal  section. 

ficant,  and  without  scent,  and  in  some  polygamous  and  zygo- 
morphic.  S4-5,  P4-5,  A8  (less  frequently  5-10)  inside  the  nectary 
(disc)  ;  ovary  generally  3-locular,  with  1-2  ovules  in  each  loculus 
(raphe  ventral,  micropyle  turned  downwards).  Seed  without 
endosperm,  often  with  an  aril.  The  embryo  is  often  thick  and 
curved  (Fig.  473). 

jffisculus  (Horse-Chestnut).  Trees  with  opposite,  digitate,  den- 
tate leaves  without  stipules ;  the  inflorescence  is  composed  of 
unipared  scorpioid  cymes  arranged  in  a  pyramidal  panicle  (termed 
a  thyrsus).  The  flowers  are  irregular,  with  an  oblique  plane  of 


AESCULIN^. 


441 


sy mmetry (th rough  the  4th  sepal,  Fig.  471);  there  are  5  sepals,  5  free 
petals,  of  which  the  one  lying  between  S3  and  S5  is  the  smallest 
(see  Fig.  471)  and  may  be  absent  ;  stamens  7  (5  +  2),  three  being 
suppressed ;  gynoecenm  simple,  3-carpellary  and  3-locular,  with 
single  style  ;  of  the  two  ovules  one  is  ascending,  the  other  descend- 
ing (Fig.  472). — The  fruit  is  a  3-valvate,  sometimes  spiny,  capsule, 
with  loculicidal  dehiscence,  the  seed  having  a  large  hilum,  a 
curved  embryo  without  endosperm  and  united  cotyledons  (the 
radicle  lies  in  a  fold  of  the  testa,  Fig.  473).  JE.  hippocastanum 
(Greece,  Asia),  introduced  into  cultivation  about  300  years  ago; 
the  majority  of  the  other  species,  e.g.  JE.  pavia,  etc.,  several  of 
which  are  frequently  cultivated  in  gardens,  are  from  N.  America. 
The  flower  of  the  Horse-Chestnut  is  adapted  for  bees,  whose  abdomen  touches 
the  anthers  or  style  when  visiting  the  flower.  The  flowers  are  protogynous. 

The  other  Sapindaceaa  have  most  frequently  4  sepals,  8  stamens,  various 
fruits  (septicidal  capsule,  nuts  with  or  without  wings,  schizocarp),  etc.  Ser- 
jania,  Cardiospermum,  Sapindus,  Koclrc.uti-ria,  etc.  (about  118  genera,  970 
species).  The  seeds  of  Paullinia  sorbili.s  contain  caffeine,  and  are  used  as 
"Pasta  guarana,"  in  the  North  Western  Brazils  in  the  manufacture  of  a 
common  drink.  Neplieliuin  (or  Euphoria)  litchi  (with  edible  aril),  and  other 
species,  from  Asia. 

Order  4.  Aceracese.  This  order  is  so  closely  allied  to  the 
Sapindacece,  that  some  authorities  have  classed  it  with  them. 
The  main  difference  is 
in  the  regularity  of  the 
flowers,  and  the  2-mer- 
ous  gynoeceum  (in  ab- 
normal cases  several 
carpels  occur). — They 
are  trees,  and,  like  the 
Horse-Chestnuts,  have 
opposite  leaves  without 
stipules ;  in  Acer  the 
leaves  are  palminerved, 
but  imparipinnate  in 
Negundo,  a  plant  fre- 
quently cultivated  in 
gardens.  The  flowers 
are  often  unisexual,  polygamous  (some  species  have  g -,  £-  and 
?  -flowers) ;  sepals  5,  petals  5  free,  stamens  8  (that  is,  5  +  5,  but 
the  two  median  ones  are  absent)  inside  a  large  disc.  Fruit  a 
samara  (schizocarp)  with  2  winged,  nut-like  fruitlets  (Fig.  474). 


FIG.  474.  — Samnra  of  Acer  platanoides. 


442 


DICOTYLEDONES. 


In  each  of  the  2  loculi  of  the  ovary  are  2  ovules.  Embryo  curved, 
with  thin,  folded  cotyledons.  Endosperm  absent. — The  inflorescences 
are  racemes  with  a  more  or  less  elongated  main  axis  and  terminal  flower  (which 
sometimes  has  10  stamens);  when  the  lateral  branches  are  developed  they  are 
similar  to  the  main  axis.  In  some  species  both  corolla  and  petal- stamens  are 
suppressed.  Acer  is  pollinated  by  insects,  Negundo  by  the  wind. — 88  species  ; 
North  Temperate  zone.  Acer  in  the  Tertiary  from  the  Oligocene.  The  follow- 
ing are  native  plants:  Maple  (Acer  campestre),  Sycamore  (A.  pseudoplatanus, 
doubtful  native).  Important  as  avenue  trees  and  timber.  Sugar  is  obtained 
from  the  spring  sap  of  the  Sugar  Maple  (N.  Am.). 

Order  5.  Malpighiaceae.  A  tropical  (especially  American)  order  closely 
related  to  the  Aceracese,  having  often  the  same  form  of  fruit  (but  3-partite). 
Some  species  are  lianes  with  anomalous  stem-structure.  Leaves  opposite.  The 
flowers  are  regular  or  obliquely  zygomorphio  (the  plane  of  symmetry  passing 
through  sepal  3,  with  S5,  P5,  A5  +  5,  G3;  1  pendulous  ovule  in  each  loculus. 
Important  characteristics  for  identification  are  the  numerous  grandular  struc- 
tures on  the  sepals.  Peculiar  2-spined  hairs  are  found  in  some.  Malpighia, 
Bunchosia,  Galphimia,  Tetrapteris,  Heteropteris,  etc.— About  600  species. 

Order  6.  Erythroxylacese.  Sepals  5,  petals  5  (with  a  ligular  corona),  10 
stamens  in  one  bundle.  Gynoeceum  3-locular.  Fruit  a  drupe.  Tropical 
(especially  American)  trees  and  shrubs,  the  Coca-plant  (Erythroxylon  coca) 
being  best  known.  Its  leaves  are  considered  by  the  inhabitants  of  Chili  and 
^Peru  to  be  one  of  the  indispensable  necessaries  of  life  ;  they  are  chewed,  and 
possess  intoxicating,  exhilarating  properties,  and  contain  the  alkaloid  cocaine, 
which  is  frequently  employed  as  a  local  anaesthetic. — 103  species  ;  chiefly  in 
America. 

Order  7.     Vochysiaceas.     Trees  ;  Trop.  Am.     1  stamen. — 140  species. 

Order  8.     Trigoniaceae.     Shrubs;  Trop.  Am. — 30  species. 

Order  9.  Tremandraceae.  Polygalacese  with  regular  flowers. — 27  species. 
Australia. 

Order    10.     Polygalacese.      Herbs  or  shrubs   (some   tropical 
species  are  lianes)  with    scattered   (rarely  opposite),  simple  and 
most  frequently  quite  entire  leaves,  without  stipules.     The  flowers 
are  usually  borne  in  terminal  spikes  or 
racemes,  and  are   strongly  zygomorphic 
(the  plane  of  symmetry    being    median)  ; 
they  have    5   free  sepals,    the   2  lateral 
ones  of  which  (4  and  5  in  Figs.  475,  476) 
are  very  large,  petaloid,  and  frequently 
project  on  each  side   like  the  "wings" 
of  a  Pea- flower  ;  petals  5,  of  which  the 
two    lateral  ones    are  wanting  or  rudi- 
FIG.  ,475.  — Diagram  of  Poly-      mentary   (dotted  on   Fig.  475),  and  the 

gala:  d  a  gland  in  the  posterior  anterior  "  the  farf  "  (Fif?.  tfQ  c)  ig  la™ 
side  of  the  flower ;  a  and  /3  the 

two  caducous  bracteoies.  hollow  and  boat-shaped,  and  frequently 


FRANGULIN.E. 


443; 


with  a  lobed  or  fimbriated  edge  (Fig.  476  A  and  B,  c)  ;  stamens  8, 
the  two  median  ones  being  absent,  all  united  into  a  tube  split 
along  the  back,  which  is  also  slightly  united  to  the  keel  (the 
anthers,  often  2  locular,  open  by  pores,  Fig.  476  B,  st)  ;  the  2  median 
carpels  form  a  bilocular  ovary.  1  pendulous  ovule  in  each  loculus 
(Figs.  476  C,  475)  ;  capsule  compressed  with  loculicidal  dehis- 
ceiice,  rarely  a  nut.  Polygala  (Milk- wort). 

470  species  ;  distributed  over  the  whole  globe  (none  Arctic).  OFFICINAL  :  the 
root  of  P.  senega,  from  N.  Am.  Some  are  used  as  ornamental  plants. 

POLLINATION.  The  flowers  of  Polygala  are  pollinated  by  insects  (chiefly  bees). 
The  fimbriated  processes  of  the  anterior  petal  support  the  insect  when  it  alights. 
The  anthers  lie  on  each  side  of  the  stigma  in  the  pouch  of  the  anterior  petal ; 
the  apex  of  the  style  is  spoon-shaped,  and  immediately  bebind  it  is  a  viscid 
stigmatic  lobe.  In  reaching  the  honey  the  proboscis  of  the  insect  must  come 
in  contact  with  the  pollen  and  the  viscid  stigma,  by  winch  it  is  rendered  sticky  ; 
this  ensures  the  pollen  adhering  to  the  proboscis  and  so  being  carried  to  other 
flowers. 


FIG.  470.— Polygala  amava.  Parts  of  the  flower  (mag.)  A  Flower  from  side,  1-5  sepals  t 
c  keel;  B  flower  from  above  spread  out :  st  the  8  stamens;  c  fimbriated  edge  of  "keel"; 
C  ovary  with  style  and  stigma. 

Family  17.      Frangulinse. 

The  plants  belonging  to  this  family,  with  very  few  exceptions, 
are  trees  or  shrubs.  The  leaves  are  usually  simple  ;  stipules  may 
be  absent  or  present.  The  flowers  in  almost  all  the  orders  are 
small,  green  or  whitish ;  they  are  always  regular,  4-  or  5-merous  with 
2-5  carpels,  but  never  have  more  than  1  whorl  of  stamens,  which 
in  JRhamnacece  and  Ampelidacece  are  placed  opposite  the  petals 
(typically  5  +  5  or  4-f-4  stamens,  of  which  however  either  the  ex- 
ternal or  internal  whorl  is  always  wanting)  ;  hypogynous  or 
slightly  perigynous,  in  Rhamnacece  only  strongly  perigynous  or 
epigynous ;  generally  ^  ;  the  calyx  is  inconspicuous  ;  petals  free  or 


444  DICOIYLEDONES. 

slightly  united.  Gyrioaceum  simple  ;  ovary  generally  multilocular  • 
style  short  or  entirely  wanting.  A  disc  is  nearly  always  developed 
in  the  flower,  but  is  found  sometimes  inside  the  staminal  whorl, 
sometimes  outside  it  or  between  the  stamens.  The  ovules  are 
apotropous  (anatropous  with  dorsal  or  ventral  raphe). 

Order  1.  Celastraceae.  Euonymus  europcea  (Spindle-tree)  may 
be  chosen  as  a  type.  It  is  a  shrub  with  simple,  opposite  leaves 
and  small  caducous  stipules.  The  small,  greenish-yellow  flowers, 
borne  in  regularly-branched  dichasia,  are  regular,  £  ,  with  4  whorls, 
4-  (or  5-)  merous  in  regular  alternation.  There  is  a  thick  disc 
upon  which  the  polypetalous  corolla  (imbricate  "in  the  bud)  and 
the  stamens  are  borne,  with  a  slightly  perigynous  insertion.  The 
style  is  short  and  thick  ;  the  ovary  has  2  erect  ovules  in  each 
loculus.  The  fruit  is  a  red,  4-valvate  capsule  with  loculicidal 
dehiscence  ;  the  seeds  are  few  in  number,  and  have  a  large,  red- 
yellow  aril  (developed  from  the  micropyle).  Embryo  green,  in  a 
large,  fleshy,  white  endosperm.  The  dingy  yellow  flowers  are  generally 
visited  only  by  flies  and  ants  for  the  sake  of  the  honey  secreted  by  the  disc,  and 
while  they  run  about  on  the  flowers  they  touch  the  anthers  and  stigmas,  now 
with  one  part  of  the  body,  now  with  another.  The  flower  is  protandrous.  The 
stigmas  are  not  developed  till  several  days  after  the  opening  of  the  anthers. — 
Celastrus,  Cassine,  Catha,  etc. 

38  genera ;  300  species.  Distributed  over  the  entire  globe,  with  the  exception 
of  the  colder  districts,  and  especially  in  the  Tropics.  Some  are  ornamental 
bushes  (Kuonymu*  japonica).  The  leaves  of  Catha  edulis  are  used  by  the 
Arabs  and  Abyssinians  in  the  same  way  as  those  of  Coca  by  the  Peruvians. 

Order  2.  Hippocrateaceae.  150  species;  tropical;  chiefly  lianes.  So, 
P5,  A3,  G3.  Anthers  extrorse. 

Order  3.  Aquifoliacese  (Hollies).  The  genus  Ilex  forms 
almost  the  entire  order.  (175  species  out  of  180  ;  especially  from 

S.  Am.)  They  are  shrubs  or  trees 
with  scattered,  leathery,  simple 
leaves  (in  Hex  aquifolium,  spiny) 
with  very  small  stipules.  The 
flowers  are  small,  white,  and  borne 
in  few-flowered  inflorescences  in  the 
axils  of  the  foliage-leaves  ;  they  are 
most  frequently  unisexual  and  dioe- 
cious. There  are  4-5  sepals,  petals, 
stamens  and  carpels  in  regular  al- 
ternation :  the  calyx  and  corolla 

FIG.  477.— Ilex  aquifolium. :  '  t/< 

magnified  flower  have    their  leaves  slightly  connate  \ 


FRANGULINJB. 


44,5 


stamens  slightly  adnate  to  the  corolla ;  the  ovary  is  generally 
almost  spherical  with  a  thick,  sessile  stigma  (Fig.  477).  This 
order  deviates  especially  from  Celastraceze  in  the  absence  of  the 
disc  and  in  having  only  1  (pendulous)  ovule  in  each  of  the  4 
loculi  of  the  ovary,  and  in  having  a  drupe  with  generally  4  stones. 
Embryo  extremely  small,  at  the  apex  of  .the  large  endosperm,  with 
the  radicle  directed  upwards. — 3  genera. — Z".  aquifolium  (Holly)  principally 
on  the  coasts  of  European  countries :  from  Norway  to  W.  Denmark,  and  further 
westward.  It  is  a  common  garden  shrub  with  stiff,  shining  leaves  and  red  fruits. 
Several  South  American  species  contain  so  much  caffeine  that  they  may  be  used 
as  a  beverage  in  the  place  of  tea  (I.  paraguayensis,  Paraguay  tea,  or  Mate). 
The  Holly  does  not  contain  caffeine. 

Order  4.  Ampelidacese  (Vines).  Shrubs  with  the  stem 
swollen  at  the  insertion  of  the  petioles  and  climbing  by  tendrils 
borne  opposite  the  leaves  (Figs.  478,  479).  The  leaves  are  scattered 
(generally  J),  stalked,  stipulate,  frequently  palminerved  and  lobed, 
divided  or  compound.  The  small,  greenish  flowers  are  generally 
borne  in  paniculate  inflorescences,  ivhose  position  is  the  same  as  that 
of  the  tendrils  (Fig.  478)  ;  they  are  hypogynous  or  slightly  peri- 
gynous,  5,  with  4-5  sepals,  petals,  stamens  (which,  as  in  the 
Ehamneae,  are  opposite  the  petals ;  Fig.  480  A,  B}  and  2  carpels.  The 
calyx  is  very  small,  entire,  or  slightly  dentate  ;  corolla  valvate,  and 
in  some  falling  off  as  a  hood,  since  the  individual  parts  remain 
united  at  the  summit  (Fig.  480  A}.  Between  the  stamens  and 
gynceceum  is  situated  an  hypogynous  disc,  with  5  lobes  alternating 
with  the  stamens  (Fig.  480  A,  B,  E).  In  each  loculus  of  the 
2-locular  ovary  there  are  2  erect  ovules  (E)  ;  .the  style  is  short  or 
wanting.  The  fruit  is  a  berry.  The  embryo  is  small  and  lies  in 
a  horny,  sometimes  slightly  folded  (ruminate)  endosperm  (Fig. 
480  0,  D). 

Vitis  and  Ampelopsis  (5-merous  flowers) ;  Cissus  (4-merous  flower) ;  Leea 
(without  stipules,  corolla  gamopetalous).  The  inflorescence  in  Pterisanthes 
(E.  Ind.)  has  a  peculiar,  flat,  leaf-like  axis,  on  the  edges  of  which  <? -flowers 
are  borne,  and  on  the  surface  $  -flowers. 

The  TENDRILS  in  AmpelidaceaB  are  modified  branches,  since  they  bear  leaves 
and  may  be  abnormally  developed  as  branches  with  foliage-leaves,  and  Anally 
the  inflorescences  are  borne  in  the  position  of  the  tendrils,  and  tendrils  ave  met 
with  which  are  partly  inflorescences.  The  explanation  of  the  position  Of  the 
tendril,  namely,  right  opposite  the  foliage-leaf  but  without  a  subtending-leaf, 
has  been  much  disputed.  The  relative  positions  are  as  follows  :  in  Vitis  vinifera 
the  following  two  kinds  of  shoots  and  relative  positions  are  found  (the  other 
species  deviate  in  one  or  other  particular),  (a)  LONG-BKANCHES,  which  have 
2  scale-leaves  and  a  large  number  of  foliage-leaves  with  a  divergence  of  ^  ; 


446 


DICOTYLEDONES. 


opposite  the  lowest  3-5  foliage-leaves  no  tendrils  are  found,  then  follow :  2 
foliage-leaves  with  tendrils,  1  without  a  tendril,  2  with  and  1  without,  etc., 
with  great  regularity.  Buds  are  developed  in  the  axils  of  the  foliage-leaves 


478 


4SO. 


4~9.  481. 

FIGS.  478-481.— Fitts  vinifera. 

FIG.  478. — Branch  with  bunch  of  grapes. 

FIG.  479. — Diagram  of  the  position  of  leaf  and  tendrils.  The  branch  is  divided  into 
sections  on  the  sympodial  theory  (the  successive  generations,  I,  II,  III,  IV,  are  alternately 
white  and  shaded) ;  Ic  buds. 

FIG.  430. — A  Flower  throwing  off  the  corolla  ;  B  flower  after  the  removal  of  the  corolla ; 
C,  D  longitudinal  and  transverse  section  of  seed  ;  E  longitudinal  section  of  gynceceum  ;  » 
calyx. 

FIG.  481. — Diagram  of  branch  and  position  of  leaves  ;  si  tendril ;  It  the  main  axis  ;  ax 
stipules  of  the  foliage-leaf  shown  below ;  g  axillary -bud  (the  dwarf-branch) ;  v  its  fore- 
leaf  j  I,  12  its  first  two  foliage-leaves  with  their  stipules ;  ltt  long-branch  in  the  axil  of  v 
(everything  appertaining  to  this  branch  is  entirely  black)  j  vt  the  first  leaf  of  this  branch. 


FRANGUL1NJ1.  447 

(Fig.  479) :  these  develop  into  (/>)  DWARF-BRANCHES,  which  commence  with 
1  laterally-placed  scale-leaf  (fore-leaf  ;  Fig.  481  v)  succeeded  by  several 
foliage-leaves  with  a  divergence  of  £  (in  a  plane  at  right  angles  to  that  of  the 
mother-shoot),  but  the  whole  shoot  is  extremely  small,  and  often  dries  up  and 
drops  off  in  the  autumn,  so  that  only  the  scale-leaf,  v,  with  the  bud  (Fig.  481 
Itj}  in  its  axil  remains.  This  bud  in  the  following  year  developes  into  a  new 
long-branch,  and  since  its  leaves  lie  in  a  plane  at  right  angles  to  that  of  the 
dwarf-branch,  their  plane  coincides  with  that  of  the  long-branch  from  which  it 
is  developed  (the  grandmother  axis). — The  tendrils  no  doubt  may  most  correctly 
be  regarded  as  the  modified  main  axis  which  has  been  pushed  aside  by  a  lateral 
branch.  The  branches  are  then  sympodia,  whose  successive  shoots  bear  al- 
ternately 1  and  2  foliage-leaves :  thus,  on  the  figure  there  are  portions  altogether 
of  5  shoots  (I.-V.),  the  1-leaved  ones  are  shaded,  the  2-leaved  ones  are  white. 
The  following  facts  however  are  adverse  to  this  theory :  (1)  the  first  leaf  on  an 
axillary  bud  is  then  situated  180°  from  the  subtending-leaf  (e.g.  the  lowermost 
shaded  leaf,  Fig.  479,  180°  from  the  lowermost  white  leaf),  whilst  the  rule  in 
the  Dicotyledons  is  that  it  is  placed  only  about  90°  to  one  side.  (2)  The  buds 
(Fig.  479  K)  from  which  the  dwarf-branches  develop,  must  then  be  accessory 
and  sister-buds  to  the  sympoiial  shoots,  but  their  first  leaves  have  a  different 
relative  position  to  this,  which  is  very  peculiar,  and  a  still  more  remarkable 
fact  is  that  the  buds,  K,  etc.  are  similar  in  structure  and  present  in  all  the  axils  ; 
thus  we  only  find  accessory  buds  in  the  cases  where  no  tendrils  are  opposite 
to  the  leaves,  and  the  main  bud  must  then  be  considered  to  be  suppressed. 
(3)  The  development  proves  that  the  tendrils  arise  on  the  side  of  a  vigorous 
growing-point  of  the  stem  or  by  its  division,  and  do  not  develop,  as  might  ba 
expected,  from  the  apex  of  the  shoot.  But  these  relations,  however,  find  their 
analogues  and  are  all  capable  of  explanation,  whereas  other  less  natural  modes 
of  explanation  are  opposed  to  them. 

435  species ;  especially  in  the  Tropics ;  they  are  rarer  in  America.  In 
N.  Am.  some  Fim-species  and  Ampelopsis  quinquefolia  are  found.  Vitis 
vinifera  is  supposed  to  have  originated  in  the  districts  East  and  South  of  the 
Caspian  Sea.  Wine  is  obtained  from  Vitis -species,  especially  V.  vinifera,  and 
"raisins," — (the  name  "  currants,"  given  to  a  special  variety  with  small,  seed- 
less fruits,  is  derived  from  Corinth). — The  species  of  Ampelopsis  (Virginian 
Creeper)  are  cultivated  as  ornamental  plants. 

Order  5.  Rhamnaceae.  The  stamens  are  placed  opposite  the 
petals  as  in  the  Ampelidacese  (Fig.  482),  but  the  flowers  are  much 
more  perigynous  or  entirely  epigynous.  The  trees  and  shrubs  be- 
longing to  this  order  have  simple,  most  frequently  penninerved 
leaves  with  stipules  ;  frequently  thorny  (modified  branches).  The 
flowers  are  inconspicuous,  sometimes  unisexual  (Fig.  482),  and 
have  5  (-4)  sepals,  petals,  stamens,  and  generally  3  (2-5)  carpels. 
The  calyx  has  valvate  aestivation.  The  petals  are  very  small 
(generally  less  than  the  sepals),  often  spoon-like,  hollow,  and  em- 
bracing the  stamens  ;  a  disc  covers  the  inner  surface  of  the  thalamus 
or  the  base  of  the  style  in  the  epigynous  flower  ;  gynoeceum  simple, 
w.  B.  GG 


448  D1COTYLEDONES. 

with  one  style  and  one  erect  ovule  in  each  loculus.  The  fruit  is 
most  frequently  a  drupe.  The  embryo  is  large,  often  green  or 
yellow,  with  endosperm. 

Bhamnus  (Buckthorn)  has  a  juicy  drupe  with  3  (2-4)  stones, 
surrounded  at  the  base  by  the  persistent  portion  of  the  receptacle ; 
the  disc  is  thin.  R.  cathartica  (common  Buckthorn)  :  dioecious, 
with  opposite,  serrate  leaves.  R.  frangula  (Alder  Buckthorn)  : 
flowers  $  ,  with  scattered,  entire  leaves. — Ceanothus  (N.  Am.,  with 
richly-flowered  inflorescences  and  a  fruit  closely  resembling  that  of  the 
Euphorbias).  Phylica,  Pomaderris  (Austr.,  fruit  a  capsule).  Zityphu8,FaUuru<t, 
Colletia  (S.  Am.)  are  thorny  sbrubs  ;  C.  spinosa  has  thorny  shoots  with  small, 
caducous  leaves ;  the  seedling  has  normal  foliage-leaves.  Others  climb  by 
tendrils  as  in  the  Ampelidacese,  e.g.  Gonania. 


FIG.  482. — Rhamnus  cathartica:  A  long-styled  <J -flower;  pet  petals;  B  short-styled 
<J -flower  ;  C  long-styled  ? -flower;  D  short-styled  ? -flower  (after  Darwin). 

475  species,  40  genera ;  chiefly  in  temperate  and  tropical  climes.  Some  are 
medicinal  plants,  the  bark  and  fruit  having  purgative  properties  (the  bark  of 
Rhamnus  frangula  and  "  Cascara  Sagrada  "  from  the  bark  of  B.  purshiana  are 
officinal).  The  fruits  and  seeds  of  others  are  edible,  for  example,  the  fruits  of 
Zizyphus  lotus,  Z.  vulgaris,  Z.  spina  Christi,  etc.  Green  and  yellow  dyes  are 
obtained  from  the  fruit  of  jR  cathartica,  infectoria  and  others  (Avignon  grain). 
Ceanothus-,  Rhamnus-  and  evergreen  Phylica-syeciea  are  ornamental  shrubs. 

Family  18.     Thymelaeinae. 

Exclusively  trees  or  shrubs  with  simple,  entire,  scattered  leaves 
without  stipules.  They  have  a  strongly  perigynous,  regular,  4- 
merous  flower.  The  receptacle  (often  coloured)  envelopes  a  simple 
gynoeceum  formed  of  1  carpel  and  with,  in  most  cases,  1  ovule,  bear- 
ing on  its  edge  4  (or  5)  petaloid  sepals  and,  but  rarely  at  the  same 
time,  small,  scale-like  petals.  The  corolla  is  most  frequently 
entirely  wanting  (and  hence  these  plants  were  formerly  reckoned 
among  the  Monochlamydeee)  ;  frequently  only  one  of  the  2  whorls 
of  stamens,  which  are  situated  on  the  inner  side  of  the  edge  of  the 
receptacle,  is  developed.  The  fruit  is  most  frequently  a  \-seedea 


THYMEL2EIK&.  449 

berry  or  drupe,  or  a  nut  which  may  be   falsely  berry-like,  the 
partly  persistent  receptacle  being  fleshy  and  enveloping  it. 

This  family  appears  the  most  nearly  allied  to  the  Frangulinse,  especially  the 
Rhamnaceae,  and  may  be  considered  as  a  further  development  of  these  in  the 
direction  of  the  petaloid  development  of  the  receptacle  and  reduction  of  the 
corolla  and  gynoeceum,  which  in  this  instance  only  consists  of  one  carpel. 
Another  deviation  is  that  both  the  whorls  of  stamens  are  present,  while  one 
of  these  is  always  wanting  in  Frangulinas.  They  also  appear  to  be  related  to 
the  Lauracese  (see  page  391). 

Order  1.  Thymelseaceae.  The  flowers  are  most  frequently  § 
(Fig.  483).  The  receptacle  is  high,  generally  tubular,  coloured, 
and  bears  on  its  edge  the  4- (or  5)-merous  calyx,  with  imbricate 
aestivation.  The  corolla  is  wanting  or  is  represented  by  small 
scales.  The  stamens  are  situated  on  the  inside  of  the  receptacle, 
and  number  4  +  4  (or  5  +  5)  ;  stigma  capitate.  1  pendulous  ovule 


FIG.  483.— Daphne  mezereum:  A  flower;  B  longitudinal  section  of  pistil. 

(Fig.  483  B),  the  radicle  pointing  upwards.  The  fruit  is  most 
frequently  a  berry.  A  disc  is  sometimes  developed.  Endosperm  wanting 
or  very  slight. 

Daphne  (Spurge-Laurel,  Fig.  483)  has  a  deciduous  receptacle, 
often  coloured  ;  sepals  4  ;  petals  absent ;  stamens  4  +  4.  Berry. — 
Gnidia  (corolla)  ;  Pimelea  (2  stamens)  ;  Thymel&a ;  Passerina  and 
others. 

400  species  ;  chiefly  in  the  warm,  sub-tropical  zone,  especially  the  Cape  and 
Australia.  Only  Daphne  and  Thymelcea  in  Europe.  In  the  fruit  and  bark  of 
some,  for  example  Daphne,  pungent,  burning  and  poisonous  properties  are  found. 
The  bark  of  D.  mezereum  (native  and  cultivated)  andD.  laureola  is  officinal.  A 
specially  tough  bast  is  found  in  some  species,  for  example  Lagetta  lintearia 
(Lace-tree,  Jamaica),  which  is  used  in  weaving.  Some  are  cultivated  in  gardens 
as  ornamental  shrubs,  especially  species  of  Daphne. 

Order  2.  Elseagnaceae.  Shrubs  or  trees,  which  are  easily 
recognised  by  the  covering  of  peltate  hairs  found  upon  almost  all 


450 


DICOTYLEDONES. 


parts  of  the  plant,  causing  them  to  assume  a  silvery  or  rusty-browii 
appearance.  Stipules  are  absent ;  the  leaves  are  simple,  most  fre- 
quently scattered.  Flowers  (Figs.  484,  485)  frequently  unisexual. 
The  sepals  are  valvate,  2-4  ;  the  corolla  is  wanting  ;  stamens  4  +  4  or 
0  +  4.  The  ovule  is  erect  and  the  radicle  turned  downwards  (Fig.  486). 
The  fruit  is  a  nut,  but  becomes  a  false  fruit,  being  surrounded  by 
the  persistent  receptacle  or  the  lower  part  of  it,  and  thus  assuming 
a  berry-  or  drupe-like  appearance  (Fig.  486).  Endosperm  insig- 
nificant.— Shepherdia  (opposite  leaves)  has  4  sepals,  4  +  4  stamens, 
as  in  Daphne.  Dioecious. — Elseagnus  (Silver-leaf)  is  ^ ,  has  4-6 
sepals,  and  4-6  stamens  alternating  with  them.  Hippophae  is 
dioecious;  it  has  2  sepals  and  4  stamens  in  the  ^-flower  (perhaps 
properly  speaking  2  +  2  stamens) ;  thorny  (stem-structures). 

16  species ;   especially  ornamental  shrubs,  e.g.  Elceagnus   argentea,  angusti- 
folia ;  Hippophae   rhainnoidts  and  Shepherdia  canadensis.     Northern  Temp. 


4S4.  485.  483. 

FIGS.  484-486. — Elceagnu*  angustijolia* 

FIG.  481.— Floral  diagram.         FIG.  485. — Longitudinal  section  through  the  flower. 
FIG.  486.— Longitudinal  section  through  the  fruit. 

Order  3  (?).  Proteaceae.  This  order  has  its. chief  centre  in  the  dry  regions- 
of  Australia  (3%-^,  of  about  1,000  species),  a  smaller  number  in  S.  Africa  (^-^K 
a  few  species  in  S.  Am.  Trees  or  shrubs,  leaves  generally  scattered,  without 
stipules,  and  more  or  less  dry,  leathery,  evergreen,  and  often  of  very  different 
forms  on  the  same  plant  (undivided,  compound,  etc.)  The  flowers  are  $ 
(rarely  unisexual),  and  4-merous  in  the  single,  petaloid  perianth  and  in  the 
staininal  whorl ;  1  carpel ;  sometimes  zygomorphic.  The  perianth-leaves  are 
generally  almost  free,  with  valvate  aestivation,  often  leathery.  Small  scales 
alternating  with  the  perianth  are  often  found  at  the  base  of  the  ovary.  The 
stamens  generally  have  extremely  short  filaments,  and  are  situated  opposite, 
Bometimes  quite  on  the  tip  of  the  perianth-leaves,  in  a  spoon-like  groove.  The 
gyno2ceum  is  1-locular,  has  1-several  ovules,  and  is  often  raised  on  a  stalk -like 
internode.  The  fruit  is  a  follicle  or  nut.  The  seeds,  most  frequently  winged, 
have  no  endosperm. — Protea,  Hanglesia,  Hakea,  Banksia,  Grevillea,  etc. 


SAXIFRAGIKE.  451 

50  genera  ;  about  1,000  species.  Several  species  are  cultivated  in  our  con- 
servatories for  the  sake  of  the  flowers,  which  are  beautifully  coloured  and 
arranged  in  crowded  inflorescences.  Protandrous.  It  is  doubtful  whether  they 
were  existent  in  Europe  in  the  Tertiary  Period.  The  true  systematic  position 
of  the  order  is  doubtful.  They  are  related  to  the  Leguminosae  and  Eosiflorae, 
but  more  closely  no  doubt  to  the  two  preceding  orders. 

Family  19.     Saxifraginae. 

The  flower  is  generally  perfect,  regular  and  polypetalous, 
usually  perigynous  or  epigi/nous,  encyclic  and  5-merous  ;  most  fre- 
quently S5,  P5,  A5  +  5  or  5  +  0  and  G2  —  5,  but  other  numbers 
are  found,  especially  4  ;  the  flowers  are  very  frequently  obdi- 
plostemonous.  The  calyx  is  sometimes  large  and  the  corolla 
small  ;.  the  carpels  in  some  are  entirely  free,  in  others  more  or 
less  united.  Endosperm  is  found  in  the  majority.  The  hypogynous 
forms  approach  the  Cistiflorae,  the  others  the  following  families,  especially 
the  Bosifloraa.  This  family  is  not,  upon  the  whole,  so  well  defined  and  natural 
as  most  of  the  others.  The  Saxifragacese  proper,  approach  very  near  to  the 
Rosacese,  especially  Spircea,  and  form  a  transition  to  it.  The  forms  with  op- 
posite leaves,  as  Philadelphia,  etc.,  approach  the  Myrtiflorse,  just  as  the  Escal- 
lonise  appear  to  be  closely  allied  to  Bicornes,  especially  Vacciniacea.  Finally 
through  Pittosporacea,  they  pass  over  to  the  Frangulinas.  The  family  ter- 
minates in  very  reduced  forms,  on  the  one  hand  in  the  arborescent  orders  with 
crowded  inflorescences,  on  the  other  perhaps  in  the  very  remarkable  order 
Podostemacece. 

Order  1.     Crassulacese.    Nearly  all  are  herbs  or  small  shrubs 
with  round,  succulent  branches  and  scattered,  fleshy,  often  more 
or  less  round  leaves,  which  are  very  rarely  incised,  and  never 
have  stipules.     The  flowers  are  generally 
borne   in   dichasia  or  unipared  scorpioid  • 

cymes,  which  again  may  be  arranged  in 
racemes,  umbels,  etc.  ;  they  are  regular, 

^  ,  hypogynous  or  peryginous,  and  most 
frequently  have  free  sepals  and  petals 

(gamopetalous  corollas  with  sessile  sta- 
mens are  found  in  Cotyledon,  Bryophyllum, 
Echeveria,  and  others)  ;  the  floral  formula 

,  1  FIG.  487.  —  Diagram    of    a 

is  Sn,  Pn,  An  +  n,  Gn,  where  n  may  have  6_merous  flower  (Sedum  his. 
very  different  values,  partly  depending  panicum)  •.  w  branch  of  scor- 
upon  the  size  of  the  flower  (e.g.  4-7  in  ^** 


Sedum,  Fig.  487;  6-30  in  Sempervivum  ; 
4  in  Bhodiola,  Bryophyllum,  and  Kalanchoe  ;  5  in  Echeveria,  Um- 
bilicus, Cotyledon).  The  carpels  are/ree  and  are  placed  opposite  the 
petals  (Fig.  487).  Fruit  a  syncarp  composed  of  follicles  containing 


452  DICOTYLEDONES. 

many,  small  seeds  without  endosperm.  Outside  each  carpel  is 
found  a  small,  nectariferous  scale  (Fig.  487).  The  northern  genus, 
Rlwdiola,  is  dioecious.  The  petal-stamens  are  wanting  in  some  (Crassula, 
Bulliarda,  and  others).  The  floral-leaves  are  very  often  displaced  upon  their 
axillary  branches.  A  multitfarpellary  gynreceum  also  occurs. 

Sedum  (Stonecrop)  is  generally  5-merous  with  10  stamens  ;  Sem- 
pervlvum  tectorum  (House-leek),  12-merous,  and  with  24  stamens. 
The  leaves  of  JBryophyllum  cahjcinum  very  readily  form  buds,  and  also  fre- 
quently exude  water  from  the  edges. 

485  species  ;  especially  Temp.  (Cape,  Europe).  Principally  used  as  orna- 
mental plants. 

Order  2.  Saxifragaceae.  The  flowers  are  4-5-merous  with  £ 
(-3)  carpels,  most  frequently:  S5,  P5,  A5  +  5  (obdiplostemonous), 
G2.  They  are  regular,  £  ,  polypetalous,  hypogynous,  perigynous 
or  most  frequently  more  or  less  epigynous  (Fig.  488) .  The  carpels 
may  be  individually  quite  free,  but  are  more  frequently  united  at 
the  base,  or  the  entire  portion  enclosing  the  ovules  is  united  into  a 
1-  or  2-locular  ovary,  the  styles,  however,  are  always  free.  Fruit 
a  capsule  with  many  seeds ;  endosperm  present. — They  are  herbs, 
most  frequently  with  scattered  leaves  without  stipules  ;  but  the 
leaf-base  is  broad.  The  inflorescences  are  most  frequently  cymose, 
and  a  displacement  of  the  floral-leaves  is  frequent  (e.g.  Ghrysos- 
plenium). — Some  Saxifraga-species,  e.g.  S.  sarmentosa,  have  irregular  flower 
with  an  oblique  plane  of  symmetry.  The  petal-stamens  in  some  may  be 
wanting  :  Heuchera,  species  of  Saxifraga  and  Mitella.  The  corolla  is  wanting 
in  others. 

Saxifraga  (Saxifrage)  :  S5,  P5,  A5  +  5,  G2  (Fig.  488)  ;  capsule 
bilocular,  opening  along  the  ventral   suture  between   the  2  per- 
*v        t<\  Q\  sistent  styles.      S.  granulata  has  small 

tubers  at  the  base  of  the  stem. — Cliry- 
sosplenium  (Golden  Saxifrage)  :  4 
sepals,  no  corolla,  4  +  4  stamens ; 
1-locular  capsule. 


\ 


Protandry  is  most  frequently  found  in 
Saxifraga,  with  the  stamens  successively 
bending  towards  the  gynoeceurn;  protogyny 
is  more  rare.    In  other  genera   there   is 
Fm.  483.-So*;/raga  granulata.    Longi-       protogyny  without  any  movement  of  the 
tudinal  section  of  flower. 

stamens ;  Chrysosplemum  is  homoganmus. 

— About  300  species;  mostly  in  temperate  climates.  Saxifraga  is  especially 
Alpine.  S.  crassifolia  and  other  species,  Hoteia  japonica,  Tellima,  etc.,  are 
ornamental  plants. 


SAXI*  RAGING.  453 

The  following  genera  are  allied  to  the  Saxifragaceaa  :  — 

1.  Parnassia   (about   14  species  ;    P.  palustris,  Grass  of    Par- 
nassus).    The  flower   is  slightly  perigynous,  and  has  S5,   P5,   5 
fertile  sepal-stamens,  and  5  petal-stamens,  which  are  developed  as 
barren  staminodes,  palmately-lobed,  and  (3-)  4  carpels  united  in  a 
1-locular  ovary  with  (3-)4  parietal  placentae.   Capsule. — Protandrous. 
The  flower  has   a   slightly  oblique  plane  of    symmetry,    which  is    especially 
shown   during  its  development  and  in  the  order  of  sequence  in  which   the 
anthers  dehisce:  originally  they  lie  closely  round  the  gynoeceum;  the  anthers 
dehisce  extrorsely,  first  the  one  which  is  placed  opposite  the  most  external 
sepal  (the  f  arrangement  is  very  distinct  in  the  calyx),  the  filament  elongating 
so  that  the  anther  lies  over  the  ovary,  and  this  is  followed  successively  by  the 
4  others  in  a  zig-zag  line ;  the  filaments  bend  backwards  after  the  pollen  is  shed 
and  the  anthers  drop  off,  and  the  stigmas  are  not  developed  until  this  is  com- 
pleted.    The  barren  stamens  are  palmately  divided  into  an  uneven  number 
(7,  9,  II)  of  lobes,  tapering  from  the  centre  towards  the   edge,  and  bearing 
apparently  glandular  tips;   their  gland-like  appearance  is  supposed   to  allure 
flies  to  visit  the  flower,  or  they  may  act  as  a  kind  of  fence  which  compels  the 
insects  to  enter  the  flower  in  a  certain  way,  and  thus  effect  pollination;  the 
honey  is  secreted  on  their  inner  side,  and  not  by  the  gland-like  tips. 

2.  Adoxa  moschatellina  (Moschatel).  This  is  a  perennial,  creeping 
herb ;  the  horizontal  rhizome  has  an  unlimited  growth,  and  bears, 
in  a  scattered  arrangement,  both  foliage-leaves,  and  white,  fleshy 


FIG.  489.— Portion  of  CephoZotiw  /ollicuZaris:    fc  pitcher-like  leaf  with  thick  corrugated 
edge  (m)  and  lid  (1) ;  b  foliage-leaf  of  the  ordinary  f orrn. 

scale-leaves.  The  aerial  stem  bears  2  opposite  foliage-leaves  and  a 
capitate  inflorescence  of  5  flowers,  4  placed  laterally  (in  opposite 
pairs)  and  1  terminally.  The  flower  is  semi- epigy nous,  the  calyx 


454  DICOTYLEDONES. 

gamosepalous,  corolla  absent.  The  stamens  are  divided  to  the 
base,  so  that  each  filament  bears  a  bilocular  anther.  The  style  is 
free,  deeply  cleft.  The  terminal  flower  has  2  bracteoles,  4  sepals, 
4  stamens,  cleft  to  the  base,  and  a  4-locular  ovary.  The  bracts 
of  the  lateral  flowers  are  displaced  on  the  flower-stalk,  as  in 
Chrysosplenium,  and  united  with  the  2  bracteoles  into  a  kind  of 
3-leaved  involucre;  these  flowers  have  5  sepals,  5  split  stamens 
with  2-locular  anthers,  and  a  5-locular  ovary.  1  pendulous  ovule 
in  each  loculus.  Fruit  a  drupe,  green-coloured,  with  1-5  stones. — 
This  plant,  which  would  perhaps  be  best  placed  in  a  special  order, 
has  also  been  classed  with  the  Araliaceaa  and  Caprifoliaceae. 

The  following  are  also  allied  to  this  order  :  E<calloniacece  (arborescent  plants 
with  simple,  scattered,  leathery  leaves),  Cunoniacece  (arborescent  with  opposite 
leaves),  CephaJotacetf  (with  pitcher-like,  insect-catching  leaves  ;  Australia ; 
Fig.  489)  and  JFrancoacece.  These  have  respectively  85,  107,  1  and  3  species. 

Order  3.  Ribesiaceae  (Currants).  5-stamened  Saxifragaceee 
with  epigynous  flowers. — Moderately  sized  shrubs  with  scattered, 


490.  491.  49?. 

FIGS.  490-492.— Ribes  rubrum. 

FIG.  4-90. — Floral  diagram.  FIG.  491. — Flower  in  longitudinal  section. 

FIG.  492. — Seeds  in  longitudinal  section. 

stalked  and  palminerved,  and  generally  palmilobed  leaves,  with  a 
large  leaf-sheath.  The  flowers  (Figs.  490,  491),  most  frequently 
borne  in  racemes,  are  regular,  epigynous,  and  have  often,  above  the 
ovary,  a  cup-  or  bell-shaped,  or  tubular  prolongation  of  the  recep- 
tacle, on  which  the  sepals,  petals  and  stamens  are  situated;  they 
have  5  sepals  (often  large,  coloured),  5  small,  free  petals,  only  5 
stamens  (opposite  the  sepals)  and  a  S-carpellate  gynoeceum  with  a 
unilocular  ovary  and  2  parietal  placentae  bearing  many  ovules.  The 
fruit  is  a  berry,  whose  seeds  have  a  fleshy  and  juicy  outer  covering 
(Fig.  492).  In  some  species,  for  example  Ribes  grossularia,  there  is  found 
an  inibrancued,  or  a  3-5-branched  spine,  very  closely  resembling  the  spiny 
leaves  of  the  Berberis,  but  which,  however,  are  emergences  springing  from  the 
base  of  the  petiole.  Ribes  has  two  kinds  of  branches  :  long-brauches  and 


SAXIFRAGINJ:.  455 

dwarf-branches,  the  latter  alone  bearing  the  flowers.  —  Ribes  (Figs.  490-492). 
The  blades  of  the  leaf  are  folded  or  rolled  together  in  vernation. 
.R.  alpinum  is  dioecious. 

75  species;  especially  from  the  N.  Temp,  regions  (especially  N.  Am  ).  —  The 
receptacle  secretes  honey  on  its  inner  surface.  The  Gooseberry  -flower  is 
slightly  protandrous,  others  are  hornogamous;  insect-  and  self-pollination  are 
found.  Tlie  following  are  FRUIT  BUSHES  :  R.  nig  rum  (Black  Currant),  R.  rubrnm 
(Red  Currant),  R.  grossularia  (Gooseberry),  originating  in  Northern  Europe  and 
Asia.  ORNAMENTAL  BUSHES  :  the  North  American  R.  aureum  (Golden  Currant) 
and  R.  sanguiiieum  (Blood-red  Currant),  etc. 

Order  4.  Hydrangeaceae.  Shrubs,  with  simple,  opposite  leaves,  without 
stipules;  flowers  generally  epigynous,  4-5-merous  (Fig.  493).  —  Hydrangea  (H. 
hortensia,  etc.).  Shrubs  from  N.  Am.  and  E.  Asia;  corolla  often  valvate.  The 
inflorescence,  as  in  the  case  of  the  inflorescence  of  Viburnum  opulus  (Guelder 
Eose),  has  often  irregular,  large,  but  barren  flowers  at 
the  circumference,  whilst  the  others  are  much  smaller, 
legular  and  £  ;  the  barren  flowers  are  mostly 
4-rnerous  ;  in  these  cases  it  is  the  calyx  which  is 
large  and  petaloid,  while  the  other  parts  of  the  flower 
are  more  or  less  suppressed.  The  branches  of  the 
inflorescence  appear  to  be  partially  devoid  of  floral- 
leaves,  since  they  are  displaced  upon  the  main  axis.  — 
Philadelphia;  racemes  (with  terminal  flower),  sepals 
4  (valvate),  petals  4  (twisted),  stamens  many,  and  car- 
pels 4  (opposite  the  petals),  forming  a  4-loculHr  ovary. 
The  numerous  stamens  (20-30)  occur  by  the  splitting  Fiff.493.—  Deufaia  crenata. 


of  the  sepal-stamens  and  are  often  therefore  placed  u<na     ' 


in  distinct  bundles.  Fruit  a  capsule.  Ph.  coronaria 
(Syringa.  Mock  Orange-blossom),  from  S.  Eur.,  is  a  common  ornamental  shrub, 
AS  also  is  Deutzia  (Fig.  493)  from  N.  Am.  and  E.  Asia.  The  latter  has  S5,  P5, 
A5  +  5,  G3.  —  About  70  species. 

Order  5  (?).  Pittosporacese.  This  order  has  its  home  especially  in  Australia 
(90  species).  The  flower  has  S5,  P5,  A5  (episepalous),  G2  (3-5),  most  frequently 
a  unilocular  ovary  with  many  ovules  in  2  rows,  borne  on  2  parietal  placentae,  or 
a  bilocular  ovary.  Some  have  berries,  others  capsules.  Pittosporum,  Citrio- 
batus,  Sollya,  Billardiera. 

Order  6.  Hamamelidaceae.  Flowers  more  or  less  epigyuous,  with  S4,  PO 
or  4,  4  fertile  sepal-  stamens,  and  4  barren  petal-stamens,  bilocular  ovary  with 
1-2  ovules  in  each  loculus.  Fruit  a  capsule.  Hamanielis  :  one  species  in  Japan 
and  one  in  N.  Am.  Fothergilla.  Liquidambar:  monoecious;  flowers  in  capitula 
or  spikes  ;  $  -flowers  without  perianth,  stamens  indefinite  ;  £  -flower  :  slight 
perianth,  2-locular  ovary  with  many  ovules.  OFFICINAL:  "  Styrax-balsam," 
which  is  obtained  by  boiling  the  bark  of  Liq.  orientalis,  from  Asia  Minor. 
Liquidambar  and  Parrotia  are  found  as  fossils  in  the  Upper  Oligocene; 
llamamelis  perhaps  in  the  Chalk. 

Finally  two  orders  with  very  reduced  flowers  are  included  in  this  family. 

Order  7.     Platanaceae.     Trees,  with  large,  scattered,  palminerved  and  lobed 


456  DICOTYLEDON  ES. 

leaves,  and  ochreate  stipules  ;  the  buds  are  concealed  in  a  hollow  at  the  base 
of  the  petiole.  The  bark  falls  off  in  large  scales.  $  -  and  ?  -flowers  (monoecious) 
in  crowded,  spherical  inflorescences  which  are  placed  at  wide  intervals  on  a 
terminal,  thin,  and  pendulous  axis.  The  flowers  have  an  insignificant  calyx 
and  corolla;  the  <£ -flower  has  few  stamens;  $ -flower,  perigynous,  with  4  free 
carpels,  1  pendulous,  orthotropous  ovule  in  each.  Fruit  a  nut ;  endosperm 
absent.  5  species;  frequently  grown  in  avenues  and  parks.  P.  occidentalis 
(N.  Am.) ;  P.  orientalis  (W.  Asia.). 

Order  8.  Podostemaceae.  Aquatic  plants,  especially  in  swiftly  running 
water,  with  somewhat  of  an  Alga-like,  Moss-like,  or  thalloid  appearance  ;  they 
show  themselves  iu  many  ways  to  be  adapted  to  their  mode  of  life  and  situations 
(having  a  dorsiventral  creeping  stem,  the  flowers  sunk  in  hollows,  a  formation 
of  haptera  upon  the  roots,  and  thalloid  assimilating  roots  and  thalloid  stems, 
etc.).  Tropical ;  100  species. 

Family  20.     Rosiflorae. 

The  leaves  are  scattered,  stipulate,  or  have  at  least  a  well  de- 
veloped sheath,  which  is  generally  prolonged  on  each  side  into  a 
free  portion  ("adnate  stipules").  The  flowers  are  regular,  peri- 
gynous or  epigynous.  Calyx  and  corolla  5  (-4)-wenm.s  with  the 
usual  position.  The  corolla  is  always  polypetalous.  The  stamens 
are  present  in  very  varying  numbers  (5-co)  and  position,  but 
always  placed  in  5-  or  10-merous  whorls ;  they  are  frequently  20  in 
3  whorls  (10  +  5  +  5  ;  see  "Pigs.  494,  502,  505)  ;  the  nearer  they  are 
placed  to  the  circumference,  the  longer  they  are ;  they  are  gene- 
rally incurved  in  the  bud,  or  even  rolled  up.  The  number  of  the 
carpels  is  from  1— GO  ;  in  most  cases  all  are  individually  free 
(syncarp),  and  when  they  are  united  it  is  in  every  case  with  the 
ovaries  only,  whilst  the  styles  remain  more  or  less  free  (Pomaceae, 
species  of  Spiraea).  The  seeds  have  a  straight  embryo,  and  usually 
no  endosperm. 

The  perianth  and  stamens  are  most  frequently  pcrigynous  on  the  edge  of  the 
widened  receptacle;  its  form  varies  between  a  flat  cupule  and  a  long  tube  or  a 
cup  (Figs.  495,  496,  498,  499,  500) ;  the  carpels  are  situated  on  its  base  or  inner 
surface,  in  some  instances  on  a  central  conical  elongation  of  the  floral  axis 
(Fig.  496).  The  carpels  in  Pomacea  also  unite  more  or  less  with  the  hollow 
receptacle,  or  this  grows  in  and  fills  up  the  space  between  the  carpels,  so  that  a 
more  or  less  epigynous  flower  is  formed  (Fig.  504). — The  following  numbers  of 
stamens  occur:  5,  10  (in  1  whorl),  15  (10  +  5),  20  (10  +  5  +  5),  25  (10  +  10  +  5), 
30-50  (in  10-merous  whorls)— compare  the  diagrams.  The  theoretical  explan- 
ation of  this  relation  of  the  10-merous  whorls  and  their  alternation  with  the 
5-merous  whorls  is  not  definitely  determined;  a  splitting  of  the  members  of  the 
5-merous  whorls  may  be  supposed,  but  the  development  shows  no  indication  of 
this,  and  it  is  not  supported  in  any  other  way.  Several  genera  have  "  gynobasic, " 
styles,  that  is,  the  style  springs  from  the  base  of  the  ovary  (Fig.  497  A,  B). 


EOSIFLORJ;. 


457 


The  Rosifloras  are  on  one  side  closely  related  to  the  Saxifragaceae  (especially 
through  Spiraea)  from  which  it  is  difficult  to  separate  them,  and  to  the  Myrti- 
florae  ;  on  the  other  side  they  are  allied,  through  the  Mimosacae  with  the  large 
number  of  stamens,  and  through  the  Amygdalacese  with  its  single  carpel,  to  the 
Leguminosas.  The  family  begins  with  forms  which  have  many-seeded 
follicles,  and  passes  on  the  one  side  to  forms  with  nuts  and  drupes  in  perigynous 
flowers,  and  on  the  other  side  to  the  Pomaceae. 

Order  1.  Rosaceae.  Herbs  or  shrubs,  generally  with  com- 
pound leaves  and  persistent  (adnate)  stipules,  flower  perigynous, 
gynceceum  formed  of  many  free  (therefore  oblique)  carpels,  syncarps 
with  fruitlets  of  various  kinds.  The  exceptions  are  noted  under 
the  genera. 

I.  SPIEJIE^:  (Fig.  495)  has  2-many  ovules  in  each  ovary,  while 
in  the  other  groups  there  is  generally  only  1,  and  never  more  than 
2  ovules  in  each  loculus.  There  are  generally  5  cyclic  carpels  and 
the  fruit  is  5  follicles,  which  are  not  enclosed  by  the  receptacle. 


FIG.  494.— Diagram  of  Comarum  FIG.  49f.— Flower  of  Spiraea  lanceolata. 

palustre. 

The  majority  are  shrubs.  Stipules  are  often  wanting. — Spiraea 
(Meadow-  Sweet).  The  flowers  are  generally  borne  in  richly 
flowered  inflorescences  of  various  forms.  The  carpels,  in  some 
species,  unite  together  and  form  a  simple  gynceceum  with  free 
styles  (an  approach  to  the  Pomacese). — Closely  allied  to  Spiraea  are  the 
East  Asiatic  shrubs  :  Kerria  japonica,  which  has  solitary  flowers,  in  this  country 
nearly  always  double  (the  fruit  a  nut),  and  Rhodotypus  kerrioides  which  has 
Opposite  leaves,  a  remarkable  feature  among  the  Rosifloras ;  it  has  a  4-merous 
flower,  a  well  developed  disc  inside  the  andrcecium,  and  a  drupe.  Closely 
allied  also  is  Gillenia  (N.  Am.)  differing  chiefly  in  the  ascending  ovules,  Spiraea 
having  pendulous  ovules,  and  a  more  tubular  receptacle. 

The  groups  Quillajece  and  Neuradece  form  a  transition  from  Spiraea  to 
Pomaceos,  In  the  first  group,  which  contains  only  trees  or  shrubs  with  generally 
simple  leaves,  the  carpels  are  either  free  or  united  (into  a  capsule) ;  in  the 
second  the  receptacle  unites  with  the  carpels,  which  are  themselves  often  united 
together  ;  in  this  case,  too,  the  fruit  is  a  capsule.  QuilJaja  (S.  Am.) ;  Exochorda 
(China). 


458  DICOTYLEDONES. 

2.  POTENTTLLE.E  (Figs.  494,  496,  497).  The  flower  has  an 
"  epicalyx  "  (Fig.  494  G)  alternating  with  the  sepals  and  formed  by 
their  stipules  which  are  united  in  pairs,  and  hence  its  leaves  are 
often  more  or  less  deeply  bifid.  The  receptacle  is  cupular  and 
often  quite  insignificant.  The  sepals  are  valvate  in  the  bud.  The 
large  number  of  fruitlets  are  achenes,  borne  on  a  well-developed 
convex  portion  of  the  receptacle  (the  Eanunculeaa  resemble  the  Poten- 
tillese,  but  have  no  epicalyx,  no  enlarged  receptacle,  and  spirally-placed  stamens). 
Most  of  the  species  are  herbs  with  dichasial  inflorescences,  often 
arranged  in  racemes. — Potentilla  (Cinquefoil),  The  achenes  are 
borne  on  a  dry,  hairy  receptacle  ;  the  style  is  situated  towards  the 
apex  of  the  ovary,  and  is  not  prolonged  after  flowering.  Herbs 
with  digitate,  in  some,  however,  pinnate  leaves,  and  generally  yellow 
flowers. — Comarum  (Fig.  494)  (Marsh  Cinquefoil)  forms,  by  its 
fleshy-spongy  receptacle,  a  transition  to  the  next  genus. — Fragaria 
(Strawberry)  (Fig.  496).  The  receptacle  becomes  finally  fleshy, 


FIG.  496,  497.— Fragaria  vesca. 

FIG.  496. — Longitudinal  section  of  flower. 

FIG.  497.— A  carpel,  entire,  and  in  longitudinal  section. 

coloured,  and  falls  off  (biologically  it  is  a  berry)  ;  the  numerous 
fruitlets  (drupes  with  thin  pericarp)  have  basal  styles  (Fig.  497)  ; 
leaves  trifoliate;  long,  creeping  runners. — Geum  (Avens)  has  a 
terminal  style  which  after  flowering  elongates  into  a  long  beak, 
with  the  apex  (after  the  uppermost  part  has  been  thrown  off)  bent 
back  into  a  hook,  thus  furnishing  a  means  of  distribution  for  the 
fruits.  Leaves  pinnate. — Dryas  comprises  3  Arctic  or  Alpine  species  with 
simple  leaves  and  solitary  flowers,  the  calyx  and  corolla  8-(J-merous,  the  fruit 
resembles  that  of  Geum,  but  the  styles  become  still  longer  and  feather-like 
(a  flying  apparatus). 

3.  RuBK2£.  Rubus  (Bramble)  has  the  same  form  of  receptacle 
as  the  P'otentillc'fe,  but  no  epicalyx;  the  fruitlets  are  drupes,  not  en- 
closed by  the  persistent  calyx.  Most  frequently  shrubs  or  under- 


ROSIFLORJ]. 


459 


shrubs  with  prickles  (emergences),  glandular  bristles  and  com- 
pound leaves.  In  the  Raspberry  (R.  idce.us)  the  fruitlets  unite 
together  and  detach  themselves  from  the  receptacle. 

4.  ROSBJ!.    Rosa  ;  the  receptacle  is  hollow,  ovoid  and  contracted 
beneath  the  insertion   of  the 

calyx  (Fig.  498),  ultimately 
fleshy  and  coloured  ;  it  encloses 
a  large  number  of  fruitlets 
which  are  achenes  as  hard  as 
stones  ("  hip,"  biologically  a 
berry). — Shrubs  with  impari- 
pinnate  leaves  and  adnate 
stipules.  The  sepals  show  clearly 
the  order  of  their  development  (a 
divergence  of  §),  the  two  outer  ones 
ou  both  sides  are  lobed,  the  third 
one  on  one  side  only,  and  the  two 
last,  whose  edges  are.  covered  by  the  others,  are  not  lobed  at  all.  Prickles 
(emergences)  are  generally  present  and  in  some  species  are  placed  in  regular 
order,  being  found  immediately  below  each  leaf  (usually  two)  although  at  some- 
what varying  heights. 

5.  AGRIMONIES.     The  receptacle  is   more  or  less  cup-  or  bell- 
shaped,  and  almost  closed  at  the  mouth  ;  it  is  persistent  and  en- 
velopes the  nut-like  fruitlets,  but  is  dry,  and  in  some  species  hard, 
the  fruitlets  being  firmly  attached  to  it.     In  biological  connection 


Fia.  498. — Longitudinal  secdon  of  flower 
of  Rosa. 


600. 


FIG.  499,  500.— Agrimonia  eupatoria. 


FIG.  499.— Flower  in  longitudinal  section. 

FIG.  500.— Fruit  and  receptacle  in  longitudinal  section. 

with  this  the  number  of  the  carpels  is  generally  only  1  or  2,  and 
the  whole  becomes  a  false  nut  (Fig.  500).  Herbs. — Agrimonia 
(Agrimony ;  Figs.  499,  500) ;  the  perianth  is  5-merous,  stamens 
5-20.  The  receptacle  bears  externally,  on  the  upper  surface,  a 


460  DICOTYLEDONES. 

number  of  hooked  bristles  which  serve  as  a  means  of  distribution 
for  the  1-2  achenes  which  are  enclosed  in  it,  and  hence  the 
entire  flower  finally  falls  off.  The  inflorescence  is  a  long  upright 
raceme.  These  bristles  are  arranged  in  whorls  of  5  and  10,  of  which  the 
uppermost  alternate  with  the  sepals. — Alchemilla  (Ladies-mantle ;  Fig. 

501)  has  8  green  perianth-leaves 
in  two  whorls  (some  authorities 
consider  the  four  outer  as  an 
epicalyx,  and  the  flower  therefore 
apetalous),  and  4  stamens  alter- 
nating with  the  innermost  whorl. 
There  is  only  one  carpel  with  a 
basal  style  and  capitate  stigma. 

TTm.  501.-Flower  of  AlcUemMa  in  longi-        The  flowerg  are    smaU  and  green- 
tudinal  section.  .-1,1        ^,  .    .     .     i         rn, 

ish,  the  filaments  ]omted.      The 

anthers  open  by  one  extrorse  cleft.  The  leaf -sheath  entirely 
envelops  the  stem ;  the  leaves  are  palminerved.  A.  aphanes  has 
often  only  1-2  stamens.  The  following  genera,  with  4-merous  flowers 
borne  in  short  spikes  or  capitula,  are  allied  to  this  group.  Sanguisorba  has 
entomophilous,  £  -flowers  with  4  (-20)  stamens,  1  carpel;  stigma  papillose. — 
Poterium ;  spike  or  capitulum,  the  uppermost  flowers  are  ?  ,  the  lowermost  $ , 
and  some  intermediate  ones  £  (the  order  of  opening  is  not  always  centripetal) ; 
S4,  PO,  A20-30,  G2,  the  long  styles  having  brush-like  stigmas  (wind-pollination). 
Leaves  imparipinnate. 

TOLLINATION.  A  yellow  ring  on  the  inner  side  of  the  receptacle,  inside  the 
stamens,  serves  as  a  nectary  when  any  honey  is  formed  ;  this,  for  instance,  is 
not  the  case  in  Rosa,  Agrimonia,  Spircea  ulmaria,  S.Jiiipendula,  S.  aruncus,  etc., 
to  which  the  insects  (especially  flies  and  bees)  are  allured  by  the  quantity  of 
pollen.  Hornogamy  and  slight  protogyny  are  frequent,  in  many  instances  self- 
pollination  also  is  finally  possible.  Poterium,  with  the  long-haired  stigma,  is 
wind-pollinated. — About  550(1100?)  species,  especially  in  northern  temperate 
regions. — USES.  OFFICINAL  :  the  petals  of  Rosa  centifolia  and  gallica,  the  fruits 
of  the  Raspberry  (Rubus  id&us),  the  rhizome  of  Geum  urbanum,  the  flowers  of 
the  Koso-tree  (Hagenia  abyssinica  or  Brayera  anthelmintica). — The  bark  of 
Quillaja  saponaria  (Chili)  is  used  as  soap  and  contains  saponin.  "  Attar  of 
Eoses  "  from  Rosa  damascena,  centifolia  and  other  species,  especially  from  the 
southern  slopes  of  the  Balkans.  Many  species  and  varieties  of  Hoses  are 
ORNAMENTAL  plants :  from  S.  Europe,  Rosa  lutea  (the  Yellow  Eose),  R.  gallica 
(the  French  Eose)  and  R.  rubrifolia ;  from  W.  Asia,  R.  centifolia,  of  which  the 
Moss  Eoses  (R.  muscosa  and  cristata)  are  varieties,  and  R.  damascena ;  from  India 
and  N.  Africa,  R.  moschata  (the  Musk  Eose) ;  from  China,  R.  indica  (Tea  Eose) 
•etc.,  besides  the  native  species  and  the  varieties  which  have  been  derived 
from  them.  In  addition,  Kerria  japonica,  species  of  Potentilla,  Rubus  odoratus 
from  N.'  Am.,  and  many  species  of  Spiraea  from  South-eastern  Europe  and  N. 
Am.  ESCULENT  :  the  "  hips  "  of  R.  mollissima,  R.  pomifera,  etc. ;  the  fruits  of 

\ 


ROSIFLORJ:. 


461 


FIG.  602.— Diagram  of 
Prunus  virginiana. 


Rubtis-species  :  Raspberry  (R.  idteus),  Cloudberry  (R.  chamcemorus),  Blackberry 
(R.  fruticoms),  etc. ;  of  .Fra/jana-species  (F.  vesca,  collina,  grandijiora,  etc). 

Order  2.  Amygdalacese.  Trees  or  shrubs  with  rosaceous 
flowers ;  leaves  simple  with  caducous  stipules ;  a  regular,  perigy- 
nous  flower,  the  receptacle  being  partly 
thrown  off  by  a  circular  slit ;  sepals  5, 
petals  5  stamens  20-30;  gynwceum  simple, 
formed  of  1  carpel  (hence  oblique,  Fig.  502), 
with  terminal  style  and  2  pendulous  ovules, 
ripening  into  a  drupe  (Fig.  503).^- The 
leaves  are  penninerved  and  frequently  have 
glands  on  the  stalks  and  edges  ;  thorns 
(modified  branches)  often  occur,  i.e.  dwarf- 
branches,  which,  after  producing  a  few 
leaves,  terminate  their  growth  in  a  thorn 
{e.g.  Prunus  spinosa).  The  vernation  of  the  foliage-leaves  varies  in  the 
different  genera ;  in  the  Almond,  Peach,  Cherry,  and  Bird-Cherry  they  are 
folded;  in  the  Apricot,  Plum,  Sloe  and  Bullace,  rolled  together.  In  some  the 
flowers  unfold  before  the  leaves  (Amyydalus,  Armeniaca}.  That  the  gynoeceum 
is  formed  of  1  carpel  is  evident  in  this  as  in  other  instances  (e.g.  in  the 
Leguminosae,  which  are  closely  related  to  this  order),  from  the  fact  that  the 
carpel  is  oblique,  and  has  only  one  plane  of  symmetry,  and  similarly  in  the 
fruit  there  is  a  longitudinal  groove  on  one  side  which  indicates  the  ventral 
suture.  It  is  only  exceptionally  that  both  ovules  are  developed.  In  abnormal 
instances  more  than  1  carpel  is  developed. 

A.  FRUIT  HAIRY:  Amygdalus  {A.  communis,  Almond-tree)  has 
a  dry  pulp  which  is  detached  irregularly,  when  ripe,  from  the 
wrinkled,  grooved,  ovoid  and 
somewhat  compressed  stone. — 
Persica  (P.  vulgaris,  Peach-tree) 
differs  from  the  Almond  in  hav- 
ing a  juicy  pulp,  not  detachable 
from  the  stone,  which  is  deeply 
grooved  and  has  pits  in  the 
grooves  (Fig.  603).  (The  name 
of  the  genus  is  derived  from  Persia, 
though  it  is  a  native  of  China). — 
Armeniaca  (A.  vulgaris,  Apricot) 
has  a  hairy,  velvety  fruit,  but 
the  stone  is  smooth  and  has  two 
ribs  along  one  of  the  edges  ;  the 
pulp  is  juicy.  (The  generic  name  has  been  given  on  the  incorrect  assumption 
that  it  was  a  native  of  Armenia  ;  its  home  is  China.) 


FIG.  503.— Fruit  of  the  Peach.    The  pulp  is 
cut  through  so  that  the  stone  is  visible. 


462  DICOTYLEDONES. 

B.  FRUIT  GLABROUS  (i.e.  without  hairs)  :  Primus  (Plum)  has  a 
glabrous  fruit  with  bluish  bloom ;  the  stone  is  .compressed,  smooth 
or  wrinkled.  The  flowers  are  borne  solitarily  or  in  couples,  and 
open  before  or  at  the  same  time  as  the  leaves ;  they  are  borne  on 
shoots  without  foliage-leaves. — Cerasus  (Cherry)  has  a  glabrous, 
spherical  fruit,  without  bloom,  and  a  spherical  stone.  The  flowers 
are  situated  in  2-many-flowered  umbels  or  racemes,  and  open  at  the 
same  time  as  the  leaves  or  a  little  before  them.  Long-stalked  flowers 
in  umbels  are  found  in  C.  avium  (Wild  Cherry),  C.  vuluaris  (the  cultivated 
Cherry,  from  Western  Asia) ;  racemes  at  the  apex  of  leaf-bearing  branches  and 
small  spherical  fruits  are  found  in  C.  padus  (Bird  Cherry),  C.  virginiana,  C. 
laurocerasus  (Cherry-laurel),  C.  mahaleb. 

POLLINATION.  Primus  splnosa  (Sloe,  Blackthorn)  is  protogyuous,  but  the 
stamens  are  developed  before  the  stigma  withers.  Honey  is  secreted  by  the 
receptacle.  Cerasus  padus  (Bird-Cherry)  agrees  in  some  measure  with  P.  apinosu. 
In  the  flowers  of  the  Plum  and  Cherry  the  stamens  and  stigma  are  developed 
simultaneously  and  self-pollination  seems  general ;  the  stigma,  however,  overtops 
the  inner  stamens  and  thus  promotes  cross-pollination. — DISTRIBUTION.  114 
species  in  the  N.  Temp,  zone ;  few  in  the  warmer  regions  ;  the  majority  from 
W.  Asia.  C.  vulgaris,  from  the  regions  of  the  Caspian ;  Primus  spinosa, 
insititia  (Bullace),  domeslica  (Plum,  from  the  Caucasus,  Persia). — USES,  princi- 
pally as  fruit-trees  :  Cherry,  Plum,  Apricot,  etc. ;  "  Almonds  "  are  the  seeds  of 
Amyrjdalus  communis  (W.  Mediterranean),  "  bitter,"  "  sweet,"  and  "  shell  " 
almonds  are  from  different  varieties,  the  latter  being  remarkable  for  the  thin, 
brittle  stone.  In  the  majority  of  species  and  in  almost  all  parts  of  the  plant 
(especially  the  bark,  seed  and  leaves)  is  found  the  glycoside,  amygdalin,  which 
forms  prussic  acid.  Many  form  gum,  and  the  seeds  have  fatty  oils  ("Almond 
oil").  OFFICINAL:  the  seeds  and  oil  of  Ami/gdalus  commnnis,  and  the  fruit  of 
the  Cherry ;  in  other  countries  also  the  leaves  of  C.  laurocerasus. — The  stems 
of  Cerasus  mahaleb  are  used  for  pipes.  Ornamental  Shrubs  :  Amygdalus  nana, 
Cerasus  laurocerasus. 

Order  3.  Chrysobalanaceae.  Tropical  Amygdalaceae  with  zygomorphic 
flower  and  gynobasic  style.  200  species;  especially  Am.  and  Asia.  Chrysoba- 
lanus  icaco  (Cocoa-plum)  is  cultivated  on  account  of  its  fruit  (Am.) 

Order  4.  Pomaceae.  Trees  and  shrubs,  most  frequently  with 
simple  leaves  and  caducous  stipules.  The  flowers  (Fig.  505)  have 
5  sepals,  5  petals  and  generally  20  stamens  (10  +  5  +  5',  or  10  +  10  +  5). 
There  are  from  1-5  carpels,  which  unite  entirely  or  to  some  extent 
with  each  other,  and  with  the  hollow,  fleshy  receptacle  (the  flower 
becoming  epigynous),  (Figs.  505,  506,  507).  The  carpels  are  nearly 
always  free  on  the  ventral  sutures,  rarely  free  at  the  sides  also. 
The  wfrole  outer  portion  of  the  fruit  becomes  fleshy,  but  the  por- 
tions of  the  pericarp  surrounding  the  loculi  (endocarp)  are  most 
frequently  formed  of  sclerenchymatous  cells,  and  are  more  or  less 


JJOSIFLOEJ;. 


463 


firm  (the  "core").  The  nature  of  the  fruit  varies,  according  to 
the  thickness  and  hardness  of  the  endocarp,  being  either  a  "  berry" 
or  a  "drupe"  (see  A  and  J5).  When  the  endocarp  is  thin  and 
parchment-like,  the  fruit  has  the  characteristics  of  a  berry,  each 
of  the  5  loculi  generally  present  containing  several  seeds  ;  but  when 
this  is  hard  the  fruit  resembles  a  drupe,  only  one  seed  is  developed 
in  each  loculus,  and  the  number  of  the  loculi  is  reduced  to  one  or 
two.  There  are  nearly  always  2  ovules  in  the  loculi  of  the  ovary, 
but  in  Cydonia  there  are  a  large  number  in  2  rows.  In  the  genera 


FIG.  £05.— Floral  diagrai 
of  Mespilus  germ&nica . 


FIG.  504. — Longitudinal  and  transverse  section  through  the  flowers  of  A,  B  Cotoneaster ; 
C  Cydonia;  D  Mains  communis;  E  Raphiolepis ;  F  Cydonia;  G  Mespilus. 

which  have  stones,  only  one  seed  is  developed  in  each  stone.  The 
genera  are  distinguished  mainly  in  accordance  with  the  kind  of 
fruit  and  the  number  of  ovules  and  seeds. 

A.  SORBET.  THE  ENDOCARP  is  PARCHMENT-LIKE  OR  PAPERY  (drupe, 
with  thin  stone  or  berry) . 

?. .  Pyrus  and  Cydonia  •  carpels  completely  embedded  in  the 
cup-like  receptacle,  styles  always  free. — Pyrus :  the  fruit  is 
glabrous,  and  has  only  a  small  calyx,  withering  or  deciduous, 
and  a  5-locular  ovarv  with  at  most  2  ascending  ovules  in  each 

W.  B.  H  H 


464 


D ICOT  YLE  DORIES. 


loculus  (Fig.  504  D).  The  large  flowers  are  situated  in  few- 
flowered  umbels  or  corymbs.  P.  communis  (Pear ;  free  styles,  Fig.  507  ; 
it  has  the  well-known  pear-shaped  fruit ;  the  core  is  reduced  to  several  groups  of 


FIG.  503.—  Cydonia  valgans.    Longitudinal  section  of  fruit. 

sclerenchymatous  cells  embedded  in  the  pulp,  the  leaf-stalk  is  as  long  as  the 
blade). — Cydonia  (Quince)  has  a  hairy  fruit  with  many  seeds  in  2 
rows  in  each  loculus  of  the  endocarp  (Figs.  504  (7,  F ;  506);  the 
testa  of  these  seeds  is  mucilaginous.  C.  vulgaris,  large,  terminal 
flowers  on  lateral  branches, 
and  large  leaf -like,  per- 
sistent sepals. 

2.  Mains  and  Amelan- 
chier(Aronia}\  carpels  free 
on  the  ventral  edge ;  styles 
united.  Malus  communis 
(Apple)  the  fruit  is  "  um- 
bilicate  "  at  the  base  ;  no 
sclerenchymatous  cells  in 

the  pulp  ;   Styles  united  at  FIG.  507.— Longitudinal  section  of  Pear  flower. 


465 

the  base  (Fig.  504  D) ;  leaf-stalk  shorter  than  the  blade.  Sorbus 
(Mountain-ash)  differs  only  in  having  a  2-3-locnlar  fruit  with 
•extremely  thin  endocarp.  Cymose  inflorescences  in  umbellate 
•cymes.  S.  aucuparia  has  pinnate  leaves,  S.  aria  (White-beam)  and  other 
•species  have  simple  leaves. — Amelanchier  (the  Service-tree)  has  a  false  divi- 
sional wall  springing  from  the  dorsal  suture,  and  more  or  less  projecting  into 
•each  of  the  loculi  of  the  ovary  ;  Raphiolepis  (Fig.  504  E)  has  racemes  and  a 
juicy  berry;  Eriobotryajaponica  (Loquat). 

B.  CRATEGEJ:.  THE  ENDOCARP  is  HARD  AND  sour  ("drupes," 
generally  with  several,  sometimes,  however,  with  only  1-2  stones, 
rarely  one  multilocular  stone;  only  1  seed  in  each  of  the  loculi). — 
Cratce.gus  (Hawthorn,  May).  There  are  1-5  stones  in  the  spherical 
or  ovoid  fruit.  The  disc,  found  on  the  apex  of  the  fruit,  inside 
the  small,  withered  calyx,  is  small  (much  less  than  the  transverse 
-section  of  the  fruit).  Shrubs  with  thorns  (branches)  and  mode- 
rately largo  flowers  borne  in  corymbs.— M espilus  (Medlar)  differs 
from  the  last-named  only  in  having  a  large  disc  at  the  apex  of  the 
fruit,  inside  the  large,  leaf-like  sepals,  i.e.  almost  equal  to  the 
•greatest  diameter  of  the  fruit.  The  flowers  are  solitary  and  ter- 
minal.— Cotoneaster  is  chiefly  distinguished  from  the  others  by 
its  syncarps,  the  2-5  carpels  (and  stones)  being  free  from  one 
another,  and  only  united  to  the  receptacle  by  a  larger  or  smaller 
portion  of  their  dorsal  surface  (Figs.  504  A,  B).  Small  shrubs 
with  leathery  leaves,  generally  covered  with  white,  felted  hairs  on 
the  lower  surface,  and  with  small  flowers ;  the  fruit  is  red  or 
black. 

Pear,  Apple,  Mountain  Ash  and  Hawthorn  have  protogynous  flowers  which 
•secrete  honey,  and  are  conspicuous  to  ensure  insect  pollination. — 180  species  ; 
iiu  the  northern  temperate  regions. — Pear  and  Apple  are  especially  cultivated 
as  fruit  trees  in  a  number  o£  varieties;  the  Paradise  Apple  (Pyrus  baccata)  ; 
-especially  in  southern  countries  also  the  Quince  (from  N.  Persia  and  the 
Caucasian  districts),  Medlar  and  Amelanchier  vulgaris.  Malus  pumila  (Caucasus, 
Altai)  and  M.  dasyphylla  (Orient,  S.  Eur.)  are  regarded  as  primitive  forms  of 
the  Apple-tree  ;  M.  sylvestris,  which  grows  wild  in  European  forests,  appears  to 
have  been  less  used.  The  early  Lake-dwellers  in  Switzerland  had  the  apple- 
tree  both  wild  and  cultivated. — The  original  form  of  the  Pear  is  supposed  to  be 
Pyrus  achras  (Central  Asia) . — Many  of  the  species  of  Cratcegus,  some  with  double 
flowers,  and  Pyrus  (Ch&nomeles)  japonica,  with  brilliant  red  flowers,  are  culti- 
vated as  ornamental  shrubs.  OFFICINAL  :  Quince  pips,  on  account  of  the 
mucilaginous  testa. — The  fruits  contain  free  organic  acids  and  sugar;  prussic 
acid  -nay  be  obtained  from  the  seeds.  The  wood  of  the  Pear-tree  is  used  in 
manufactures. 


466  DICOTYLEDONES. 

Family  21.     Leguminosae* 

The  most  characteristic  feature  is,  that  the  gynceceum  is  1-1  ocular 
and  formed  of  1  carpel,  the  ventral  suture  of  which  is  turned  pos- 
teriorly. The  fruit,  in  most  instances,  is  apod  (legume),  which  opens 
generally  along  both  sutures,  the  two  valves  twisting  more  or  less 
in  opposite  directions.  In  other  instances  it  opens  along  one  suture  only, 
or  as  a  pyxidium  (Bed  Clover),  or  it  is  indehiscent,  in  which  case  it  is  more  or 
less  berry-like  (e.g.  the  Tamarind,  Carob-bean),  or  it  is  a  drupe  (e.g.  the  Ton- 
quin-bean),  or  a  1-few-seeded  nut  (e.g.  Melilntus),  or  a  lomentum,  which  divides 
transversely  into  as  many  joints  as  there  are  seeds  (Ornithopus,  see  Fig.  513). 

The  inflorescences  belong  to  the  centripetal  type  (i.e.  indefinite)  ; 
cymes  do  not  occur.  The  flowers  are  zygomorphic,  with  vertical 
plane  of  symmetry,  seldom  regular  ;  5-merous  with  but  a  few  ex- 
ceptions, 5  ,  and  slightly  perigynous.  The  following  diagram  if* 
the  most  general  (Fig.  511)  :  5  sepals,  with  the  unpaired  sepal 
median  and  anterior,  5  petals,  5  +  5  stamens,  all  in  alternating 
whorls,  1  carpel.  The  calyx  is  most  frequently  gamosepalous,  the 
gynoaceum  is  narrowed  down  at  the  base  to  a  short  stalk  and,  in 
the  majority,  is  more  or  less  bent.  The  seed  is  most  frequently 
kidney-shaped,  with  a  smooth,  hard  and  shining  testa,  the  hilum 
being  very  distinct.  Endosperm  is  wanting,  or  is  reduced  to  a  thin 
layer,  which  is  of  service  when  the  seed  swells  during  germina- 
tion. The  vegetative  parts  have  these  features  in  common,  namely, 
the  leaves  are  scattered,  stipulate,  and  almost  always  compound. 
Peculiar  sleep-movements  and  sensitiveness  are  found  in  some,  chiefly 
in  the  Mimosas.  Many,  probably  all,  LegumiriosaD  have  small 
tubercles  on  their  roots  which  are  produced  by  a  kind  of  bacterium, 
and  assist  in  the  assimilation  of  free  nitrogen.  Spontaneous  move- 
ments are  exhibited  by  Desmodium  gyrans  (Telegraph-plant). 

This  family  is  closely  allied  to  the  Eosiflorap,  with  which  it  agrees  in  the 
scattered  leaves,  the  presence  of  stipules,  the  generally  5-merous  and  most 
frequently  perigynous  flowers  with  eucyclic  stamens,  and  the  absence  of  endo- 
sperm. Amygdalacea  and  Chrysobalanacece,  with  solitary  carpels,  approach  on 
one  side  to  the  Leguminosse,  among  which  genera  with  drupes  are  also  found  ; 
Mimosaccce,  with  their  many  stamens,  form  a  connecting  link  on  the  other 
side.  In  this  respect  the  Mimosa-genus  Affonsea,  and  certain  Csesalpine*  and 
Swartzieas,  are  of  special  interest  in  having  more  than  one  carpel  (syn.rarp),  a 
condition  which  is  sometimes  met  with  abnormally  in  other  Leguniinosa',  as 
well  a^  in  Amygdalaceas.  About  7,000  species  of  the  Legumicosse  are  known. 

Order  1.  Csesalpiniacese.  These  are  leguminous  plants  with 
straight  embryo  and  a  flower  ivhich  is  not  papilionaceous  and  has  not 


LEGUMINOS^. 


467 


the  same  aestivation  (Figs.  508-510) ;  but  in  reality  there  is  not  a 
single  characteristic  which  absolutely  distinguishes  them  from  the 
Papilionacea?. — The  majority  are  aborescent ;  the  leaves  as  a  rule 
are  pinnate  or  bi-pinnate.  The  flower  is  5-merous,  most  frequently 
perigynous  and  slightly  zygomorphic ;  the  calyx  is  free  or 
gamosepalous,  the  corolla  polypetalous  with  ascending  imbricate 
(Estivation  (i.e.  the  two  lowest  petals  envelop  the  lateral  ones, 
and  these  again  the  posterior;  Fig.  508)  ;  10  free  stamens;  fruit 
various. 

Cassia  (Figs.  508-510)  is  the  largest  genus  (about  200  species)  ; 
it  has  an  almost  hypogynous,  zygomorphic  flower  with  5  free 
sepals  and  petals  ;  of  the  10  stamens  the  3  posterior  are  generally 
barren,  the  others  are  of  very  unequal  length  and  open  at  the 
apex  by  pores  (Fig.  509).  In  some  (the  Senna  group)  the  fruit  is 


?iGr.  508. — Floral  diagram. 


FIGS.  608-510.— Cassia  flo rib urc da. 
FIG.  509.— Flower.  FIG.  510. — The  same  in  long.  sect. 


a  flat,  short,  thin,  dehiscing  pod ;  in  others  (Cathartocarpus)  it  is 
round,  long,  woody  or  fleshy,  indehiscent,  and  divided  internally 
by  more  or  less  fleshy  transverse  walls  into  as  many  cells  as  there 
are  seeds. — The  following  also  have  DEHISCENT  FRUITS  :  Bauliinia 
(often  lianes,  tropical  climbers  with  tendrils  [stem-structures]  and 
anomalous  stems),  Copaifera,  Hce.matoxylon  (whose  pod  does  not 
dehisce  along  the  suture,  but  laterally),  Cercis  (simple  leaves;  the 
corolla  resembles  that  of  the  Papilionacea3,  but  the  posterior  petal 
is  the  smallest,  and  is  enveloped  by  the  2  lateral  ones,  which  are 
enveloped  in  their  turn  by  the  2  anterior). — FRUIT  INDEHISCENT: 
Tamarindu*  indica ;  -the  pod  is  almost  round,  often  a  little 
abstricted  between  the  seeds ;  the  wall  is  formed  by  a  thin, 
brittle  external  layer,  enclosing  an  acid  pulp ;  well-dey eloped 


468  DICOTYLEDONES. 

septa  are  present  between  the  seeds  ;  the  most  internal  layer  is 
parchment-like.  Calyx  4-merous  by  the  coalescence  of  2  sepals. 
Only  3  fertile  stamens.  —  Ceratonia  siliqua  (Carob-bean,  Locusts)  ; 
the  pod  is  long-,  compressed,  with  thick  sutures,  and  has  a  wall, 
the  central  part  of  which  is  more  or  less  leathery,  fleshy  and 
sweet;  there  are  transverse  septa  between  the  seeds,  as  in  the 
Tamarind.  Embryo  greenish  in  endosperm.  The  flower  is  without 
a  corolla,  5  stamens.  —  Pterogyne  (winged  fruit),  etc.  —  KiUMiiiui^ 
with  Krameria  is  an  anomalous  group. 

DISTRIBUTION.  80  genera,  with  740  species;  almost  exclusively  in  the  Tropics. 
The  Carob-tree  and  Cercis  grow  in  the  Mediterranean  basin.  The  largest  and 
most  widely  distributed  genus  is  Cassia,  which  is  found  as  trees,  shrubs,  and! 
weeds  iu  all  tropical  countries.  The  order  has  many  important  uses  to  man- 
kind. MEDICINAL  :  the  leaves  and  pods  of  Cassia  acutifoKa,  and  ang-uatifolicu 
(officinal,  Senna-leaves),  the  fruit-pulp  of  the  Cassm-sub-gemis,  Cathartocarpits. 
Rhatany  root  from  Krameria  triandr<i  (Peru,  officinal).  Balaam  is  extracted 
from  a  number  of  Co/?ai/mi-species  (Balsam  of  Copaiba)  from  S.  Am.  (offi- 
cinal), and  from  Hymencea  (Copal  balsam),  Tracliylobinm  and  others.  Edible. 
fruits  are  obtained  especially  from  the  Carob-tree  (from  the  East)  and  the- 
Tamarind  (officinal).  The  heart-wood  of  several  species  of  CcKsalpinia,  such 
as  C.  brasiliensis  (the  Pernambuco-tree),  echm&Ui  (Red-tree),  and  *api>((nt 
yield  dyes  ;  Hcematoxylon  (H.  campechianum,  Logwood),  Copaifera  bracteata 
(Amarant-tree).  —  Timber  is  obtained  from  many  (Melanoxylon  and  others). 
In  Europe  they  are  of  little  importance  as  ornamental  plants-,  these  being  con- 
fined principally  to  the  species  of  Gleditscliia  (G.  tviaeantlta,  from  N.  Am.) 
and  Cercis  (the  Judas-tree,  C.  siliqua  strum,  S.  Eur.),  which  are  cultivated  in 
gardens  ;  but  in  tropical  gardens  beautiful  flowering  species,  e*g.  of  Cassia, 
Poinciana,  Broivnea,  are  found,  and  the  most  beautiful  of  all  ornamental  plants, 
the  Indian  Ainherstia  nobilis. 

Order  2.      Papilionacese.       The    flower   (Figs.    511,    512)  is 
strongly  zygomorphic  and  somewhat  perigynous  (Fig.  512  B  ;  most 
frequently   more   on    one    side    than    the    other).       The   calyx   is 
gamosepalous  and  persistent.     The  polypetal- 
ous  corolla  has  descending  imbricate  asstiva- 
tion,  the  posterior,  large  leaf,  the  standard 

(Fi°s-  5n  -^  ;  512  B'>  e^  coveri»y  in  the  bud 

rhe  two  lateral  ones,  the  wing*  (Figs.  511  v  ; 
512  B',  a),  which  again  cover  the  two  an- 
terior ;  these  are  united  in  the  form  of  'a  boat, 
the  keel  (k  and  c)  ;  the  wings  and  the  two 
FIG.  5ii.  —  Diagram  of  petals  of  the  keel  are  very  unsym  metrical. 


Paba     vulgaris:      f     the        rpj^    the    keel     ig    forme(J    Qf     two     petals    is 
standard  ;    v  the  wings  ;  .  .,.,.,,  f  1N 

fc  the  keel.  seen   by  its  position  (in  front  or  one  sepal) 


LEGUMINOS^E. 


469 


and  by  the  two  often  more  or  less  free  claws.  The  10  (5  +  5) 
stamens  (monadelphous)  are  either  all  united  into  one  bundle,  or 
into  two  bundles  (diadelphous),  the  posterior  one  being  free  (Fig. 
512  C).  The  ovules  are  curved  and  also  the  embryo  (Fig.  512  0\ 
especially  the  hypocotyl,  so  that  the  radicle  assumes  a  position 
close  to  the  edge  of  the  thick,  fleshy  cotyledons.  Endosperm 
wanting ;  the  cotyledons  are  very  rich  in  proteid  reserve  material. 
The  forms  of  the  fruit  and  exceptions  are  described  under  the 
genera. 


c- 


FIG.  512.— Pisum  sativum  :  A  entire  flower ;  B  in  longitudinal  section ;  C  gynoaceum 
and  stamens  ;  D  gynceceum  ;  B'  corolla  dissected,  e  standard,  a,  a  -wings,  c  keel ;  D  seed 
opened  to  show  the  cotyledons  (c),  the  radicle  (r),  the  plumule  (g) ;  E  fruit  (legume) ;  F  seed. 

Geocarpic  fruits,  i.e.  those  which  penetrate  the  soil  during  their  development 
and  ripen  underground,  are  found  in  e.fj.  Arachis  hypogcea  (see  page  472), 
Trifolium  subterrancum,  Vicia  amphicarpcea.  Germination  takes  place  in 
various  ways.  In  the  majority  the  cotyledons  are  raised  above  the  ground  as 
green,  leaf -like  bodies ;  in  the  Vicieas  they  remain  thick  and  white,  and  are 
always  enclosed  in  the  testa,  and  are  therefore  never  able  to  take  part  in  the  work 
of  assimilation  ;  in  species  of  Phaseolus,  on  the  other  hand,  they  are  raised  well 
above  the  ground  and  become  green,  but  remain  however  thick  and  fleshy. 

i,  2.  The  two  groups  PODALYRIE^E  (the  majority  of  the  genera  are 
Australian)  and  SOPHORE;E  (Sophora,  Edivardslx,  etc.),  represent  the  oldest 


470  DTCOTYLEDONES. 

type,  as  they  have  10  free  stamens  and  so  form  the  transition  to  the  Cresal- 
piniaceas.     Nearly  all  are  trees  and  shrubs. 

3.  ASTKAGALEJ:.     Herbs   or  shrubs,   less  frequently  trees,  with 
imparipinnate  leaves  (without  tendrils).     The  flowers  are  gener- 
ally   borne     in    racemes    or    spikes.     Stamens    monadelphous    or 
diadelphous. — Astragalus    (Milk- Vetch)    has    the    legume   incom- 
pletely divided  longitudinally  into  2  loculi  by  a  septum  formed  by 
the   incurved    dorsal    suture.     Diadelphous. — Gli/cyrrhiza  (Liquo- 
rice) ;  Colutea  (Bladder-Senna)  from  S.  Europe ;  Robinia  (the  false 
Acacia)    with    thorny    stipules ;    Indigofera    (the    Indigo    plant) ; 
.  Amorpha  (which  has  only  one  petal,  namely  the  standard,  and  the 
fruit  a  nut),  Caragana,  Wistaria  (a  climbing  shrub),  Galega.     Car- 
michselia  australis,  when  old,  produces  flat  branches  with  scale- 
like  leaves. 

4.  VlCiE-E.     Climbing  herbs  with  paripinnate  leaves,  the  midrib 
ending  in  a  point  or  frequently  in  a  tendril,  Which  generally  is 
branched,  representing  lateral  veins  without  mesophyll ;  stamens 
diadelphous ;  the  cotyledons  remain  underground  on  germination. 
—  Vicia  (Vetch)  has  a  filamentous  style,  hairy  towards  the  tip,  and 
a  pod  with  many  seeds  ;  climbing  by  means  of  tendrils ;  the  leaves 
have  many  leaflets. — Faba  (F.  vulgaris,  Horse-bean)  is  erect,  with- 
out tendrils ;  its  pod  is  thick  with  spongy  septa  between  the  seeds. — 
Ervum  (Lentil)  has  a  pod  with  only  1-2  seeds,  and  sweeping  hairs 
(stylar-brush)  on  the  inner  side  of  the  style. — Pisum  (Pea ;   Fig. 
512)  has  very  large  stipules,  the  bent  style  has  a  hollow  groove  on 
the  anterior  side.    P.  sativwfo  (Common  Pea),  P.  arvense  (Grey  Pea). 
—Lathyrus  (Sweet  Pea)  generally  has  an  angular,  winged  stem  and 
most  frequently  only  a  few  pairs  of  leaflets.     The   style  is  flat- 
tened, with  sweeping  hairs  on  the  back.     In  L.  aphaca  the  stipules 
alone  are  developed  into  foliage-leaves,  while  the  remainder  of   the  leaf  is 
modified  into  a  tendril. — Cicer   has    a   nearly    straight  embryo  and 
imparipinnate  leaves  with  dentate  or  incised  leaflets.     C.  arietinus 
(Chick-pea). — Abrus  (precatorius,  etc.);  the  seeds  ("Crab's  eyes,"  "  Pater- 
noster peas,"  "  Jequirity  ")  are  scarlet  with  a  black  spot  round  the  hilum. 

5.  PHASEOLEJE.     Herbs,  twining  or  erect,  but  not  climbing  by 
tendrils  ;  the  leaves  are  imparipinnate,  generally  ternate,  and  bear 
small,  linear  bodies  resembling  stipules  at  the  base  of  the  stalks 
of  the  leaflets.     The  inflorescences  are  most  frequently  compound, 
groups  of  few  flowers  being  situated  on  short,  nodose,  lateral  axes 
borne   on  a   longer    stem.     On   germination    the    cotyledons    are 
raised    a   considerable   distance   above    the    ground,  and    become 


LEGUMINOS^.  471 

greenish,  but  do  not  become  leaf-like  ;  in  P.  multiflorus  they  remain 
underground.  Stamens  as  in  the  Vetches. — Phaseolus  (Kidney- 
bean)  :  the  keel  with  the  stamen  and  style  is  spirally  twisted, 
(to  the  right).  Herbs,  twining  to  the  left. — The  "  Calabar  -bean " 
(Physostiymd  venenosum},  Erythrina,  Clitoria,  Glycirie,  Soja,  Mucuna,  Apios, 
Canavalia,  Vigna,  Dolichos,  Cajanus,  Rht/nchosia,  etc. 

6.  TRIFOLIEJ:  (CLOVERS).    Herbs  with  ternate  leaves,  the  leaflets 
are  often  dentate  with  the  veins  prolonged  into  the  teeth ;  stamens 
diadelphous;  fruit  1-locular,  1-few-seeded,  pyxidium-like,  irregu- 
larly dehiscent,  or  more  frequently  a  nut.    The  flowers  are  generally 
borne  in  capitula,  racemes,  or  spikes. — Trifolium  (Clover).     The 
corolla  is  gamopetalous.     The   calyx  persists,  together  with  the 
corolla,  round  the  ripe  fruit.     The  inflorescence  is  a  spike,  capitu- 
lura  or  capitate  umbel  ;  the  leaves  are  ternate,  and  have  adnate 
stipules.—  Medicago  (Medick).     The  corolla  falls  off  after  flower- 
ing ;  fruit  curved  like  a  sickle  or  spirally  twisted ;  it  is  a  nut,  and 
opens  with  difficulty.     Leaves  ternate. — Melilotus  (Melilot)  has  a 
small,  spherical  or  lanceolate,  thick  and  wrinkled  fruit,  which  as 
a  rule  is  indehiscent.     The  inflorescence  is  a  raceme,  often  long, 
or    a   spike,    sometimes    a   capitulura.      Leaves    ternate. — Ononis 
(Rest-harrow)  differs   in    having  monadelphous    stamens  and  in 
being  more  shrub-like  and  bushy,  and  in  having  a  normal,  2-valved 
pod,   by  which  characteristic  it  approaches  the  Genisteas.     The 
flowers  are  generally  rose-coloured,   solitary,   or  in   few-flowered 
racemes  in  the  leaf-axils.     Thorns    (branches)  are  often  present; 
the  leaves  are  compound  with  only  one  small  leaflet  (the  terminal 
one),  or  ternate  with  adnate  stipules.. 

7.  LOTE$.     Herbs  with  ternate  or  imparipinnate  leaves,  with 
entire  leaflets.      In  the  latter  case,  when  the  lowest  pair  of  leaf- 
lets is  placed  quite  close  to  the  sheath,  the  stalk  is  wanting,  and 
apparently   a   trifoliate   leaf    with    large    stipules   is    developed. 
Flowers    in  an  umbel  or  capitulum.     Stamens  monadelphous  or 
diadelphous,  the  filaments  (either  all  of  them,  or  only  the  5  sepal- 
stamens)  are  widened  at  the  top. — Lotus  (Bird's-foot-trefoil)  has  a 
long,  round  pod. — Tetragonolobus. — Anthyllis  (Lady's-finger)  ;  the 
fruit  is  a  nut,  which  is  distributed  by  the  wind  by  means  of  the 
membranous,   bladder-like  calyx,   which  completely  encloses  and 
falls  off  with  it. 

8.  G-ENISTEJL    The  majority  are  shrubs  or  trees  with  apparently 
simple  leaves,  i.e.    compound  leaves    with  only    one  leaflet   (the 
terminal  leaflet),  or  ternate  leaves  ;  the  stipules  in  most  instances 


472 


DICOTYLEDON  ES. 


are  very  small  or  are  entirely  wanting;  stamens  monadelphous. — 
Genista  (Dyer's-weed)  has  apparently  simple  leaves ;  the  branches 
often  terminal e  in  a  thorn.  The  strongly-winged  stems  in  G.  tagittal'u 
are  its  most  important  organs  of  assimilation.  Bosnaxi  rnfa  has  flat  branches, 
its  leaves  being  reduced  to  small,  pointed  stipules. — Sarotkamnits  (Broom) 
has  switch-like,  angular  branches  and  often  both  the  apparently 
simple  and  ternatve  leaves  on  the  same  shoot ;  style  spirally  rolled. 
—  Cytisus  (Laburnum).  JJlex  (Furze;  in  U.  europecus,  the  seedlings  bear 
a  few  foliage  leaves,  but  the  leaves  succeeding  these  are  modified  into  thorns)  ; 
Spartium;  Crotalaria,etc. — Lupinus  (Lupin)  is  allied  to  this  group; 
it  has  a  thick,  often  somewhat  fleshy  pod,  and  digitate  leaves  with 
adnate  stipules. — Retama. 

g.  HEDTSAEEJE  are  especially  recognised  by  having  the  ovary 
divided  by  transverse  septa  into  as  many  cells  as  there  are  seeds, 
the  frnit  thus  becomes  a  lomentum,  dehiscing 
transversely  into  nut-like  joints  (Fig.  513). — 
Orm'£/topM.s(Bird's-foot);  Coronilla;  Hippocrepis; 
Onobrychis  (Sainfoin)  has  a  fruit  with  only  1 
joint  (i.e.  a  1-seeded  nut) ;  Desmodium;  Alhayi ; 
Hedysarum,  etc. — Arachis  hypogcea  (Earth-nut) 
has  a  pod  which  is  abstricted  between  the 
seeds,  and  is  indehiscent,  but  is  not  multi- 
locular  nor  a  true  lomentum ;  it  is  reticulately 
wrinkled  externally,  and  ripens  underground ; 
the  basal  part  of  the  ovary  is  prolonged  after 
flowering,  attaining  a  length  of  several  inches, 
and  buries  the  young  fruit  in  the  soil.  The 
embryo  is  straight. — Desmodium  gyrans  is  well- 
known  for  its  motile  leaflets. 
10.  DALBERGIEJE.  25  genera  ;  especially  in  Tropical  America  ;  the  majority 
are  trees,  a  few  shrubs  or  lianes  ;  the  leaves  are  simple  or  imparipinnate.  The 
fruit  is  indehiscent  in  all;  in  some  it  is  a  winged,  in  others  a  wingless  nut 
(BJachan-hnu,  Dalbergia,  Centrolobium,  etc.),  in  others,  again,  a  drupe,  e.g.  in 
Diptenjx  (Tonquin-bean)  and  Andira.  In  some  genera  the  embryo  is  straight. 

POLLINATION.  Especially  effected  by  Bees.  The  nectar  is  secreted  by  a 
ring  or  disc-like  portion  round  the  base  of  the  gynoeceum  or  the  inner  surface 
of  the  receptacle.  The  flower  is  constructed  with  a  peculiar  mechanism  to 
ensure  cross-pollination  by  insects.  The  pollen  is  shed  just  before  the  flower 
opens,  and  is  retained  in  a  pouch  formed  by  the  keel.  An  insect  visiting  the 
flower  uses  the  wings  and  keel  for  a  landing-stage,  and  in  attempting  to  reach 
the  honey  presses  down  the  wings  and  the  keel  which  are  locked  together  near 
the  standard;  the  st>lar-brush  by  this  means  is  forced  through  the  apical 
opening  of  the  keel  and  a  little  pollen  is  thus  swept  out  and  deposited  upon 
the  abdomen  of  the  visiting  insect  as  it  presses  against  the  apex  of  the  keel ; 


FIG.  513. — Hedysarum 
coronarium. 


LEGUMINOSJ:.  473 

the  insect  thus  carries  away  pollen  and  may  effect  cross-pollination.  In  the 
different  flowers  this  arrangement  is  modified  in  various  ways  to  promote 
pollination.  5000  species  (319  genera) ;  especially  in  the  Tropics,  where  many 
are  important  forest  trees. — The  following  plants  are  used  FOB  FOOD:  Pixunt 
sativum  (W.  Asia  ? )  and  arvense  (Italy)  ;  Phaaeolus  vulgaris  (Kiduey-beau, 
American;  Dolichos  sinensis  was  known  to  the  Greeks  and  Romans  under 
the  name  "  0ctcr?;Aos,"  ''  phaseolus  "),  P.  compressus  (French-bean),  etc.;  Faba 
Bulgaria  (Field-bean,  Horse-bean;  from  the  Old  World)  ;  Era  um  lens  (Lentil, 
Eastern  Mediterranean) ;  in  tropical  countries  the  oil-containing  seeds  of  Arachi« 
hypogaa. —  The  following  are  FODDEK  plants:  Vicia  sativa,  't>'dba  vulyaris, 
Onobrychis  sativa  (Sainfoin),  Medicago  sativa  (Lucerne),  and  lupulina  (Medick), 
species  of  Tri  folium,  Hedysarum  corunarium.  OFFICINAL  :  "  Liquorice  root," 
from  Glycyrrhiza  glabra  (S.  Europe);  "  Red  Sandalwood,"  from  Pterocarpiis 
santaliims  (Tropical  E.  Asia) ;  Gum  Tragacanth,  from  Astragalus-species  (E. 
Mediterranean) ;  Balsam  of  Peru,  from  Toluifera  pereine,  and  Balsam  of  Tolu, 
from  Toluifera  balsamum.  Calabar-beans,  from  Physostiguia  venenosum ;  Kino, 
from  Pterocarpus  marsvpium ;  the  pith  of  Andira  araroba  is  used  under  the 
name  of  "  Chrysarobin." — Of  use  TECHNICALLY  :  Genista  tinctoria  (yellow  dye) 
and  Indigof era-species  (Indigo),  the  bast  of  Crotalaria  juncea  (Sunn  Hemp); 
tbe  seeds  of  Dipteryx,  which  contain  Coumarin,  and  ate  highly  scented,  and 
Balsam  of  Mi/roxylon.  POISONOUS:  the  seeds  of  Laburnum  (Cytisus  laburnum), 
various  species  of  Lathyrus,  and  Abrus  precatoriu*  ;  the  latter  contain  two 
poisonous  proteids,  paraglobulin  and  albumose,  which  resemble  snake-poison 
in  their  effects.  The  following  are  ORNAMENTAL  plants;  Phaseolus  multijlorus 
(Scarlet  runner,  from  America),  Sobinia  pseudacacia,  Amorpha,  Colutea,  Coro- 
nilla,  Indignjera  dosua,  Wistaria  polystachya,  Cytisus  laburnum  (Laburnum,  S. 
Europe,  Orient.)  and  other  species. 

Order  3.  Mimosacese.  The  flowers  are  most  frequently 
hypogynous  and  regular,  the  a3stivation  of  the  corolla  is  valvate 
and,  in  the  majority  of  instances,  that  of  the  calyx  also.  The  flower 
is  4-merous,  less  frequently  5-  or  3-merous.  —  The  flowers  are 
generally  small,  but  are  always  borne  in  compact,  round  capitula 
or  spikes  (Fig.  514)  ;  they  are  hypogynous  or  perigynous.  The 
calyx  is  generally  gamosepalous  and  the  corolla  gamopetalous,  the 
latter  being  frequently  wanting.  The  stamens  are  equal  or 
double  the  number  of  the  petals  (Mimosa,  etc.,  in  M.  pudica,  e.g. 
S4,  P4,  A4,  Gl)  or  (in  Acacia,  Inga,  etc.)  in  a  large,  indefinite 
number,  free  or  monadelphous,  often  united  to  the  corolla  (Fig. 
514  &).  The  colour  of  the  flower  in  most  cases  is  due  to  the  long 
and  numerous  stamens.  The  fruit  is  various.  The  embryo  is 
straight  as  in  the  Csesalpiniaceae.  Entada  and  many  species  of 
Mimosa  have  a  flat,  straight,  or  somewhat  sickle-like  pod,  which 
resembles  the  siliqua  of  the  Cruciferse  in  that  the  sutures  (in 
this  instance,  however,  dorsal  and  ventral  suture)  persist  as  a 


474 


DICOTYLEDON  ES. 


frame,  but  the  intermediate  portion  divides,  as  in  the  transversely 
divided  siliqna,  into  as  many  nut-like  portions  as  there  are  seeds. 
Some  species  have  a  pod  of  enormous  dimensions.  The  seeds  of 
Entada  gigalobium  are  often  carried  from  the  West  Indies  to  the 
"N.  W.  coasts  of  Europe  by  the  Grulf  Stream. — The  fruit  of  Acacia 
in  some  species  is  an  ordinary  pod,  in  others  it  is  transversely 
divided,  or  remains  an  undivided  fruit,  a  nut. — This  order  includes 


FIG.  514. — Acacia  farnesianu. :  a  inflorescence  ;  b  flower. 

both  trees  and  herbaceous  plants,  which  are  often  thorny;  the 
leaves  are  usually  bipinnate  (Fig.  514)  and  are  sensitive,  and  also 
possess  sleep-movements. — Many  Australian  Acacias  have  com- 
pound leaves  only  when  young,  but  when  old  have  phyllodia,  i.e. 
leaf-like  petioles  without  blades,  placed  vertically.  A  large  number 
have  thorny  stipules,  which  in  some  (Acacia  sphcerocephala)  attain 
an  enormous  size,  and  serve  as  a  home  for  ants,  which  in  return 
protect  their  host- plant  against  the  attacks  of  other,  leaf -cutting 
ants. 


PASSIFLORIN.E.  475 

Other  genera  besides  those  mentioned  are :  Adenanthera,  Des- 
mantlius,  Parkia,  Inga  (with  rather  fleshy,  indehiscent  fruit), 
Calliandra,  etc. 

1350  species  (30  genera) ;  none  natives  of  Europe,  their  home  being  the 
Tropics  and  sub-tropical  regions,  especially  Australia  and  Africa. — Fossils  in 
Tertiary. — Gums  are  found  in  many  species  of  Acacia,  especially  the  African 
(Gum  arabic)  and  Australian,  of  which  some  are  officinal.  The  bark,  and  also 
the  fruits,  contain  a  large  amount  of  tannic  acid  and  are  used  as  astringents  and 
in  tanning  ("  Bablah  "  is  the  fruits  of  several  species  of  Acacia).  Catechu  is  a 
valuable  tanning  material  extracted  from  the  wood  of  Acacia  catechu  (E.  Ind.). 
The  flowers  of  Acacia  fames i ana  (Fig.  514)  are  used  in  the  manufacture  of  per- 
fumes. With  us  they  are  cultivated  as  ornamental  plants,  e.g.  A.  lopliantha  and 
many  others,  in  conservatories. 

Family  22.     Passiflorinae. 

The  flowers  are  most  frequently  regular,  5-merous  in  the  three 
most  external  whorls,  eucyclic  and  perigynous  or  epigynous,  less 
frequently  hypogynous.  A  characteristic  feature  is  that  the 


FIG.  515.—  Passiflora  ccerulea  (reduced). 

ovary  is  tricarpellary,  unilocular,  and  with  3  parietal  placenta  which 
sometimes  meet  in  the  central  line  (Cucurlitacece) .  The  styles  arc 
generally  free  and  bifid.  To  all  these  characteristics,  however, 
there  are  exceptions.  The  Cucurbitacese  are  sometimes  placed  among  the 
Syrupetalte,  close  to  the  Campanulas,  but  they  are  not  allied  to  the  S.ympetalas, 
from  which  they  differ  especially,  for  instance,  in  the  structure  of  the  ovule. 
The  position  of  the  Begoniaceae  in  this  family  is  also  open  to  doubt. 


476  DICOTYLEDONEE. 

Order  1.  Passifloraceae  (Passion-flowers).  The  majority 
are  herbs  which  climb  by  means  of  tendrils  (modified  branches) 
and  have  scattered,  stipulate  leaves,  often  palminerved  and 
lobed  (Fig.  515).  The  flowers,  which  are  often  large  and 
beautiful,  are  regular,  § ,  with  S5,  P5,  A5,  G3  ;  the  calyx  and 
corolla  are  perigynous,  and  immediately  inside  the  corolla  is 
the  "corona,"  consisting  of  numerous,  tapering,  filamentous 
bodies,  or  sometimes  united  in  rings,  most  frequently  petaloid 
and  coloured  ;  the  stamens  are  raised  on  a  long,  round  internode 
above  the  cup-like  receptacle ;  immediately  above  these  is  the 
gynceceum  with  its  3  free  styles  and  capitate  stigmas  ;  the  ovary 
is  unilocular  with  3  parietal  placentae.  Fruit  most  frequently  a 
berry.  The  seeds  have  an  aril. 

210  species  ;  especially  in  Tropical  America.  Several  Pass(/?om-species  are 
ornamental  plants,  and  the  fruits  of  some  species  are  edible. 

Order  2.  Papayacese.  The  best  known  representative  is  the 
Papaw  (Carica  papaya),  a  Tropical  American  tree  whose  stem  is 
usually  un branched,  and  bears  at  its  summit  several  large,  palmi- 
lobed  leaves  on  long  stalks.  The  stem  and  leaves  have  latex.  The 
large,  Melon-like  berries  are  edible,  and  for  this  reason  it  is  culti- 
vated in  the  Tropics.  Flowers  unisexual,  with  slightly  different 
structure  in  the  £  -  and  $  -flowers,  besides  intermediate  forms. 
The  ^-flower  has  a  gamopetalous,  the  $ -flower  a  polypetalous 
corolla.— The  milky  juice  contains  a  substance  with  similar  action 
to  pepsine.  10  stamens.  5  carpels. 

Order  3.     Turneraceae.     85  species  ;  especially  in  America. 
Order  4      Samydaceae.     160  species  ;  tropical. 

Order  5.  Loasaceae.  Herbaceous  plants  seldom  shrubs,  some- 
times climbing,  and  nearly  always  studded  with  stif  hairs,  in  some 
instances  stinging  or  hooked.  The  leaves  are  most  frequently 
palmilobed  and  without  stipules.  The  flowers  are  regular,  J  , 
polypetalous,  entirely  epigynous,  with  4-5  sepals,  petals  and  sta- 
mens, or  more  frequently  (by  splitting)  many  stamens,  those 
which  are  placed  before  the  sepals  being  generally  barren  and 
more  or  less  petaloid;  carpels  most  frequently  3,  united  into 
an  inferior,  unilocular  ovary  with  3  parietal  placentae,  above 
which  the  receptacle  is  generally  more  or  less  prolonged.  Fruit 
a  capsule ;  iu  Gronovia  an  ovary  with  1  ovule  and  fruit  a  nut. 

115  species  ;  principally  from  S.  Am.  A  number  of  annuals  are  often  grown  in 
our  gardens  :  Bartonia  aurea  (California)  ;  Mentzelia  ;  Cajopliora  ;  Gronovia. 


PASSIFLORIN^. 


477 


Order  6.  Datiscacese.  4  species,  especially  in  the  Tropics. — • 
Datisca  cannabini  (Asia  Minor)  resembles  the  Hemp  in  external 
appearance.  The  flowers  are  dioecious,  insignificant  ;  <£ -flowers: 
a  low,  gamosepalous  calyx,  no  corolla,  and  an  indefinite  number  of 
stamens  ;  ?  -flowers  :  epigynous ;  ovary  unilocular  with  free,  mostly 
bifid,  styles,  and  generally  3  parietal  placentae.  In  most  cases  the 
ovary  is  not  entirely  closed  at  the  top  (as  in  Reseda). 

Order  7.  Begoniacese.  This  order  principally  comprises 
herbs  or  under-shrubs  with  succulent  stems  (having  scattered  vas- 
cular bundles  in  the  pith);  the  leaves  are  arranged  in  two  rows 


FIG.  516. — Begonia  rex  (reduced). 

(a  divergence  of  I)  and  are  asymmetrical,  as  a  rule  more  or  less 
obliquely  cordate,  or  ovate  with  cordate  base  (Fig.  516)  ;  large, 
caducous  stipules  are  present.  Inflorescences  dichasial,  or  uni- 
pared  scorpioid  cymes  ;  the  flowers  are  unisexual ;  the  first  ones 
(the  oldest)  are  <$  -flowers,  while  ?  -flowers  are  found  especially 
on  the  younger  axes.  The  $  -flowers  have  most  frequently  2  +  2 
coloured  perianth-leaves,  and  many  stamens  collected  into  a  head 
in  the  centre  of  the  flower ;  the  ?  -flowers  are  epigynous  with  5 
coloured  perianth-leaves  (placed  spirally  with  a  divergence  of  f) 
and  a  trilocular  ovary,  bearing  3  bifid  styles  and  3  wings  (the 
wings  usually  of  unequal  size) ;  in  the  inner  angle  of  each  loculus 


478 


DICOTYLEDONES. 


there  is  one  large  projecting  placenta,  or  two  plate-like  placentae 
(the  bent  back  edges  of  the  carpels)  studded  with  ovules.  Fruit 
a  capsule,  with  many  extremely  small  seeds. — Begonia. 

420  species  ;  almost  all  from  the  Tropics  (Am.,  Asia).— Many  species,  with 
varieties  and  hybrids,  are  ornamental  plants  in  houses  and  conservatories, 
chiefly  on  account  of  the  form,  colour  and  markings  of  their  leaves  ;  but  also 
for  their  very  beautiful  flowers.  They  reproduce  easily  by  adventitious  buds 
from  leaves  and  portions  of  leaves  placed  on  damp  soil ;  some  have  bulbils. 
Like  the  OxalideEe  they  contain  an  acid  sap. 

Order  8.  Cucurbitacese.  The  flower  is  epigynous,  and,  as  a 
rule,  is  also  provided  with  a  leaf-like,  cup-  or  bell-shaped  receptacle 
above  the  ovary,  to  which  the  perianth  and  stamens  are  attached ; 
the  flowers  are  regular,  unisexual,  with  rudiments  of  the  other  sexr 
and  5-merous  :  sepals  5,  narrow  and  pointed,  with  the  median 
sepal  posterior  (Fig.  517),  petals  5,  stamens  5,  and  carpels  3 
(rarely  4-5)  ;  the  corolla  is  gamopetalous  in  the  majority,  poly- 
petalous  in  some ;  generally  plicate- valvate  in  the  bud.  The  anthers 


Fio.517. — Eclallium  agreste.    Diagram  of  a  $-  and  a  ? -flower. 

in  the  <$  -flowers  are  extrorse,  and  monoihecious.  i.e.  only  one  half  of 
each  of  the  anthers  of  the  5  stamens  is  developed,  the  pollen-sac  hav- 
ing frequently  a  peculiar  <^>-shaped  curve  (Fig.  518  A,  B)  ;  the 
stamens  are  either  all  united  into  a  column  (e.g.  in  Cucurbita),  or 
they  are  united  in  pairs,  so  that  only  one  remains  free  (Figs.  517  A  ; 
518  A}  ;  in  the  latter  case  there  appears  to  be  one  small  stamen 
with  a  cx^ -shaped,  curved  pollen-sac  and  two  larger  ones,  each  with 
two  curved  pollen-sacs  placed  as  in  Fig.  517  A.  The  original  form 
appears  to  be  Feuillea  with  free  petals  and  5  free  stamens.  Some- 
times the  rudiment  of  a  gynceceum  is  present.  The  carpels  are 
united  inco  an  ovary  with  3  (4-5)  placentae  formed  by  their  united 
edges.  These  are  thick,  fleshy,  and  bifid,  bearing  a  number  of  ovules 
on  each  s;(le  (Figs.  517  B:  518  C,  D)  ;  in  general  the  placentae  are 


PASSIFLORINJI:.  475> 

so  large  that  they  not  only  meet  in  the  centre,  but  also  fill  up  the 
ovary  as  far  as  the  wall  of  the  pericarp.  The  whole  interior  of  the 
fruit  thus  becomes  a  juicy  mass  in  which  three  lines  may  be  seen, 
meeting  in  the  centre  (the  boundaries  of  the  individual  placentae), 
and  near  the  circumference  6  groups  of  seeds  (Fig.  518  D).  When 
the  carpels  are  equal  in  number  to  the  petals  they  alternate  with 
them.  The  style  is  short  and  thick,  and  generally  divided  into  3 
(4-5)  branches,  with  a  horse-shoe  shaped  stigma  on  each  branch 


FIG-.  518.— Citrullus  colocyntMs :  A  (J -flower,  cut  open  and  spread  out;  B  stamen; 
C  ? -flower  in  long  section ;  h  receptacle  ;  ca  calyx;  D  transverse  section  of  ovary. 

(Fig.  518  0).  The  fruit  is  most  frequently  a  many-seeded  berry; 
in  some  it  attains  a  considerable  size  and  has  a  firm  external  layer 
(Cucurbita,  Lagenaria,  etc.).  The  embryo  is  straight,  has  no  endo- 
sperm, but  contains  a  large  quantity  of  oil.  The  exceptions  to  the 
above  characters  will  be  found  under  the  genera. 

Exclusively  herbs,  generally  with  stiff  hairs  and  yellow  flowers. 
Many  species  are  annuals,  others  are  perennial,  having  tuberous 
roots  or  hypocotyls.  The  leaves  are  scattered,  long-stalked,  in 
most  cases  more  or  less  heart-shaped,  palminerved,  palmilobed, 


w.  B. 


ii 


480  DICOTYLEDONES. 

and  exstipulate;  in  their  axils  are  found  both  flowers  (singly,  or  in 
an  inflorescence)  and  a  vegetative  bud,  and  outside  the  axil,  on  the 
anodic1  side  of  the  leaf,  a  simple  or  branched  tendril,  by  which  the 
plant  climbs  (exceptions  :  e.g.  Ecballium). 

The  position  of  the  flowers,  branches  and  tendrils  situated  in  and  near  the 
leaf-axils  is  as  follows.  In  the  leaf-axils,  a  flower  is  borne  (as  a  branch  of  the 
first  order),  £  or  $  ,  according  to  the  conditions  of  the  various  genera.  This 
branch  is  not  situated  in  the  centre  of  the  axil,  but  is  removed  slightly  towards 
the  anodic  side  of  the  leaf.  Of  its  two  bracteoles  as  a  rule  only  the  one  lying  on 
the  anodic  side  is  developed,  namely  as  a  tendril,  which  is  displaced  to  a 
position  outside  the  axil.  The  branch  of  the  first  order  bears  on  its  catodic  side 
an  inflorescence  (in  the  axil  of  the  suppressed  bracteole),  on  the  anodic  side  a 
vegetative  bud  which  grows  out  into  a  branch,  like  the  main  axis.  The  sub- 
tending leaf  of  this  branch  is  thus  the  tendril ;  but  when  it  has  several  arms 
the  condition  is  complicated  by  the  appearance  of  an  accessory  bud  which 
unites  with  its  subtending  leaf,  the  tendril,  its  leaves  also  becoming  tendrils 
^situated  on  an  undeveloped  internode) ;  the  many-branched  tendril  is  thus  a 
branch,  and  the  tendril-arms  are  its  leaves,  except  the  main  arm  which  is  its 
subtending  leaf.  Other  explanations  of  these  difficult  relations  have  been  given. 
— The  germination  is  somewhat  peculiar,. owing  to  the  fact  that  a  heel-like  pro- 
longation is  formed  at  the  base  of  the  hypocotyl  to  assist  in  separating  the  two 
halves  of  the  testa  from  each  other,  and  to  facilitate  the  unfolding  of  the 
cotyledons. 

Cucurbita  (Pumpkin,  Marrow)  has  branched  tendrils ;  the  flowers 
are  monoecious,  and  are  borne  singly  ;  the  corolla  is  bell- shaped,  and 
divided  almost  as  far  as  the  middle.  The  stamens  are  all  united 
into  a  tube ;  the  compressed  seeds  have  a  thick,  blunt  edge. 
— Cucumis  has  (generally)  unbranched  tendrils ;  the  $ -flowers  are 
borne  singly,  whilst  the  ^-flowers  are  borne  in  groups  :  the  corolla 
is  divided  nearly  as  far  as  the  base,  and  the  stamens  are  united 
2-2-1.  The  connective  is  elongated  above  the  anthers.  The  seeds 
have  a  sharp  edge. — Citrullus  (Fig.  518)  has  a  corolla  similar  to 
Cucumis,  but  ty  and  <$  -flowers  are  borne  singly  ;  the  stigma  is 
only  3-lobed,  the  fruit  most  frequently  spherical. — Ecballium 
(Squirting  Cucumber,  only  1  species,  E.  elaterium)  has  no  tendrils, 
and  is  therefore  not  a  climber.  The  oblong  fruit  is  pendulous 
from  the  apex  of  its  stalk,  and  when  ripe  is  distended  with  an 
acrid,  watery  fluid  ;  on  being  touched  the  fruit  is  detached,  and  the 
seeds,  together  with  the  watery  fluid,  are  violently  ejected  through 

1  11  we  suppose  a  spiral  line  drawn  through  the  leaves  upwards  on  a  stem 
•with  scattered  leaves  (in  the  shortest  way),  then  the  side  of  the  leaf  first 
touched  is  the  catodic,  or  descending,  and  the  other  the  anodic,  or  ascending 


PASSIFLORINJ1.  481 

the  aperture  formed  at  the  base  of  the  fruit.  The  ^-flowers  are 
borne  in  racemes  near  the  solitary  $  -flowers  (Fig.  51 7). — Bryonia 
(White  Bryoriy)  has  chiefly  unbranched  tendrils  and  small, 
greenish-yellow,  usually  dioecious  flowers  with  rotate  corolla, 
in  many-flowered  inflorescences  ;  the  small,  spherical  berry  has 
no  specially  firm  outer  layer,  and  generally  only  few  seeds. 
The  tap-root  and  a  few  of  the  other  roots  are  tuberous.  S. 
alba  (berry  black;  monoecious)  and  dioica  (berry  red ;  dioecious). 
Among  other  genera  may  be  mentioned :  Layenaria  (Gourd)  ;  the  fruit  has  a 
woody  external  layer  which,  after  the  removal  of  the  pulpy  integument,  may  be 
used  as  a  gourd.  Lujj'a  has  a  polypetalous  corolla  ;  the  fruit  is  dry,  and  consists 
internally  of  a  network  of  vascular  bundles  ;  it  opens  by  an  aperture  at  the 
summit.  Benincasa ;  the  fruit  has  a  close,  bluish  coating  of  wax.  Trichos- 
anthes  (Snake  Cucumber)  has  a  thin,  round,  long  and  curved  fruit.  Momordica ; 
the  fleshy  fruit  opens  and  ejects  the  seeds.  Cyclanthera  takes  its  name  from 
the  staminal  column  which  is  found  in  the  centre  of  the  $  -flower,  bearing  a  bi- 
locular,  ring-like  anther  which  opens  by  a  horizontal  cleft.  The  fruit  is  uni- 
locular  by  suppression,  has  1  placenta,  and  when  touched  opens  and  ejects  the 
seeds.  Sicyos  and  Sechium  have  only  unilocular  ovaries  with  one  pendulous 
ovule.  Sechium  has,  moreover,  5  free  stamens,  of  which  only  one  is  halved,  the 
other  4  having  both  halves  of  the  anther.  Fevillea  and  Thladiantha  also  have 
5  free  stamens.  Dimorphochlarnys  has  dimorphic  flowers. 

POLLINATION  is  effected  by  insects,  chiefly  bees  or  wasps,  the  nectar  being 
secreted  by  the  inner,  yellow  portion  of  the  receptacle  ;  in  the  $  -flower  access 
is  gained  to  the  nectar  through  the  slits  between  the  stamens,  which  arch  over 
the  nectary. — 85  genera  ;  about  637  species  ;  especially  in  the  Tropics.  Only 
two  are  found  in  the  whole  of  N.  Europe,  Bryonia  alba  and  dioica  ;  in  S. 
Europe,  Ecballium  also.  Most  of  the  cultivated  species  have  been  obtained 
from  Asia,  such  as  the  Cucumber,  Melon,  Colocynth,  several  Lw$a-species  (the 
"  Gourds "  mentioned  in  Scripture  are  Cucumis  chate) ;  from  Africa,  the 
Water-melon,  Cucurbita  maxima,  and  others  ;  from  S.  Am.,  no  doubt,  the 
Pumpkin  (C.  pepo  and  melopepo).  USES.  Many  species  are  used  in  medicine 
or  for  domestic  purposes.  Bitter,  poisonous  properties  are  found ;  the  fruits  of 
the  two  officinal  ones  are  purgative  :  Citrullu*  colocynthis  (Mediterranean,  E. 
India,  Ceylon)  and  Ecballium  elaterium,  as  well  as  various  tropical  species,  the 
roots  of  Bryonia,  etc. — The  following  are  cultivated  AS  ARTICLES  OF  FOOD  : 
Pumpkin  (Cucurbita  pepo,  etc.),  Cucumber  (Cucumis  sativus),  Melon  (Cucumis 
melo),  the  Water-melon  (Citrullus  vulgari*),  Sechium  edule  (Chocho),  certain 
species  of  Luff  a  (the  young  fruit).  The  Bottle  Gourd  is  cultivated  in  tropical 
countries  for  the  sake  of  its  hard  pericarp,  which  is  useful  for  bowls,  bottles, 
etc.  The  fruits  of  Luff  a  have  a  number  of  reticulately  felted,  tolerably  firm 
vascular  bundles,  which  render  them  serviceable  in  various  ways  (as  a  kind  of 
il  sponge ").  The  Cucurbits  are  of  no  use  in  the  manufactures.  Only  a 
few  are  cultivated  as  ornamental  plants,  chiefly  as  curiosities. 


482  D1COTYLEDONES. 

Family  23.     Myrtiflorse. 

The  leaves  are  most  frequently  opposite,  simple,  entire  (rarely 
dentate),  and  exstipulate.  The  flowers  are  regular  and  epigynous 
(perigynous  in  Lythracece  and  a  few  others),  ^  ,  polypetalous ;  the 
number  of  members  in  a  whorl  is  generally  4  or  5  (S,  P,  A,  or  most 
frequently  A  2,  G),  but  sometimes  it  becomes  (e.g.  Myrtles  and 
Lythracete)  very  large  in  the  androecium  by  splitting,  and  in  the 
gynceceum  also  is  often  different.  (When  suppression  takes  place 
it  is  principally  in  the  corolla  and  petal-stamens.)  In  nearly  all 
instances  the  calyx  is  valvate.  Gynceceum  multicarpellary, 
multilocular,  with  only  one  style  (except  Haloragidacefe}.  In  the 
majority  the  ovules  are  situated  on  an  axile  placenta  in  the  multi- 
locular ovary.  Endosperm  is  wanting  in  the  majority. — Less  import- 
ant exceptions :  Rhizophoracete  and  Gunnera  have  stipules.  Haloragidaceae 
have  several  styles  and  endosperm.  Rhizophora  also  has  endosperm. 

Order    1.       Lythracese.      Hermaphrodite,    perigynous    flowers 
which   are   most  frequently  6-merous,  viz.    S  6  (often  with   a  com- 
"  epicalyx,"  Fig.  519  c),  one  segment  posterior,  P6,  A6  +  6 
or  6  +  0  and  G  2-6,  forming  a  2-6-locular 
ovary  with  many  ovules    in  the    loculi, 
style  single,  and  capitate  stigma.     The 
gynceceum   is   free    at    the    base   of    the 
tubular,    or    bell-shaped,    thin,    strongly 
veined  receptacle,  which  bears  the  other 
leaf -whorls  on  its  edge  and  inner  side. 
Fruit   a   capsule.      No   endosperm.  —  To 
FIG.  519.— Lythrum  saiicaria.        this  order  belong  both  herbs,  shrubs  and 

c  the  "epicalyx." 

trees.        I  he     branches     are    frequently 

square,  the  leaves  always  undivided,  entire,  and  without  stipules, 
or  with  several  very  small  stipules,  and  often  opposite.  The  calyx 
is  valvate.  The  flower  is  regular  (except  Guphea)  and  frequently 
large  and  beautiful.  The  stamens  are  generally  incurved  in  the 
bud,  and  the  petals  irregularly  folded. 

Lythruin  (Loose-strife).  The  flower  is  diplostemonous  and 
6-merous,  with  a  long,  tubular  receptacle  with  epicalyx-teeth 
(Fig.  519  c).  The  12  stamens  are  arranged  in  two  tiers  on  the 
inner  side  of  the  receptacle.  The  gynceceum  is  bi-carpellary. 
The  flowers  are  borne  in  small  dichasia  in  the  leaf-axils,  and  their  number 
is  increased  by  accessory  inflorescences  beneath  the  main  inflorescence. — 
The  native  species,  L.  saiicaria,  is  trimorphic  (long-styled,  mid-  and  short- 
styled  forms,  Fig.  520).  Cross-pollination  is  chiefly  effected  by  humble- 


MYRTIFLOR^l. 


483 


bees  and  bees,  which   seek  the  nectar  formed  at  the  bottom  of  the  recep- 
tacle.    Other  species  are  only  dimorphic,  or  even  monomorphic Closely 

allied  are,  Nesasa,  Diplusodon,  Lagerstrosmia,  and  Cuphea,  whose 
flower  resembles  that  of  Lythrum, 
but  is  zygomorphic.  In  Cuphea  the 
receptacle  is  oblique  and  at  the 
back  prolonged  into  a  spur,  in  which 
the  nectar,  secreted  by  a  gland 
situated  behind  the  ovary,  is  col- 
lected ;  the  calyx  and  corolla  gradu- 
ally become  reduced  in  size  toward 
the  anterior  side  of  the  flower  ;  the 
reverse,  however,  is  the  case  with 
the  11  stamens  (the  posterior  one  is 
absent) ;  the  posterior  loculus  in  the 
bilocular  ovary  is  sometimes  barren ; 
the  fruit,  when  ripe,  dehisces  along 
the  posterior  side,  the  ovary  as  well 
as  the  wall  of  the  receptacle  being 
ruptured  by  the  placenta,  which 
expands  and  projects  freely.  The 
flowers  stand  singly  in  the  centre  of  the 
stem,  between  the  pairs  of  leaves.  This 
may  be  explained  as  follows  :  of  the  two 
foliage-leaves  in  each  pair,  one  supports  a  foliage-shoot,  the  other  a  flower ; 
the  foliage-shoot  remains  in  the  axil,  but  the  flower  is  displaced  through  the 
length  of  an  entire  internode  to  the  next  pair  of  leaves,  and  then  assumes  a 
position  between  these  two  leaves.  All  foliage-shoots  stand  in  two  rows,  the 
flowers  in  two  other  rows. 

Pep  Us  (Water-purslane),  a  small,  annual,  plant,  with  thin,  bell-shaped  re- 
ceptacle without  projecting  nerves.  The  small  flowers  have  no  petal-stamens, 
and  often  also  no  corolla ;  fruit  indehiscent. — Ammannia  is  closely  allied  to  it. 

365  species  ;  30  genera  ;  mostly  in  the  Tropics,  and  more  especially  S.  Am. 
—  Some  yield  dy?s,  e.g.  Lawsonia  inermis  (cultivated  in  Africa  and  Asia)  and 
Lagerstrcemeria  indica  •  some  contain  tannin  ;  others  are  orn  imental  plants, 
especially  in  gardens  in  warm  countries. 

Order  2.  Blattiaceae.  12  species.  Tropical  Asia  and  Africa.  Trees. 
Formerly  included  with  Punica,  but  best  placed  as  an  independent  order. 

Order  3.  Melastomaceae.  A  very  natural  and  very  large  order  (150 
genera;  2,500  species),  its  home  being  chiefly  in  tropical  S.  America,  especially 
the  Brazils  (termed  by  Schouw  "  The  kingdom  of  Palms  and  Melastomacese  "). 
There  are  both  herbaceous  and  arborescent  species,  which  are  easily  recog- 
nized by  the  opposite  or  verticillate,  simple  leaves  which  have  (with  the  excep- 
tion of  a  few  heather-like  species)  3-5-7-9  curved  veins  proceeding  from  the 
base  of  the  leaf,  and  connected  very  regularly  by  closely  parallel,  transverse 


FIG.  620. — Lythrum  salicaria.  One 
side  of  the  perianth  is  removed  from 
all  three  flowers.  A  is  long-styled, 
£  mid-styled,  and  C  short-styled.  The 
direction  of  the  arrows  and  dotted 
lines  indicates  the  best  (legitimate) 
methods  of  crossing. 


484  DICOTYLEDONES. 

veins.  The  flower  is  perigynous  or  epigynous  ;  its  type  is  that  of  the  Onagraceae 
(4-5-merous  ;  1  whorl  of  sepals,  petals  and  carpels,  2  of  stamens) ;  the  calyx 
is  valvate,  the  corolla  is  twisted  (to  the  left)  in  aestivation ;  the  stamens  are 
very  characteristic  ;  in  the  bud  they  are  geniculate  ;  the  anther  opens  in  the 
often  long,  beak-like,  prolonged  point,  with  1,  less  frequently  with  2  pores,  and 
has  generally  ear- like  appendages  at  its  base.  The  fruit  is  a  berry  or  cap- 
sule. These  large  and  beautiful  flowering-plants  play  a  very  important  part  in 
South  American  landscapes  ;  otherwise  they  are  of  slight  importance  (a  few  are 
cultivated  in  conservatories,  e.g.  Centradenia,  Medinilla,  Lasiandra,  TiboUchincir, 
Miconia,  etc.). 

Order  4.  CEnotheraceae  (or  Onagracese).  The  flowers  are 
arranged  in  racemes  or  spikes,  §  ,  epigynous,  regular,  polypetalous, 
4-merous  in  all  5  whorls  (1  whorl  of  sepals,  petals  and  carpels,  2  of 
stamens) ;  2-3— 5— 6-merous  flowers  are  less  frequent ;  the  calyx  is 
valvate,  the  corolla  twisted  in  aastivation  (the  left  edge  being  covered). 
Grynceceum  simple  with  multilocular  ovary;  the  style  is  undivided, 
filiform,  and  bears  a  capitate  or  4-partite  stigma ;  endosperm  want- 
ing ;  embryo  straight. — The  majority  are  herbs,  especially  water- 
and  marsh-plants;  several  are  shrubs.  No  essential  oils.  The  leaves 
are  alternate  or  opposite,  always  single,  and  without  (or  with  very 
small)  stipules.  The  odourless  flowers  sometimes  have  a  coloured  calyx. 
In  'some  instances  (e.g.  (Enothera,  Fuchsia)  the  receptacle  is  prolonged  more  or 
less  beyond  the  inferior  ovary,  and  finally  falls  off.  The  stamens  are  obdiplos- 
temonous  (carpels  epipetalous) ;  the  petal-stamens  are  sometimes  suppressed. 
The  anthers  in  some  genera  are  divided  into  storeys.  The  well-pronounced, 
triangular  pollen-grains  are  connected  together  by  viscous  threads.  Small 
stipules  are  sometimes  found,  e.g.  Fuchsia,  Lopezia. 

A.  Fruit  a  capsule.  (Enothera  (Evening  Primrose)  is  4- 
merous,  has  8  stamens,  a  tubular  receptacle,  and  an  oblong  capsule 


FIG.  521. — Flower  of  Lopezia. 

with  loculicidal  dehiscence  leaving  a  centrally  placed  column,  bear- 
ing the  seeds. — Epilobitvni  (Willow-herb)  deviates  from  CEnothera 
especially  in  the  seeds  being  hairy  (at  the  chalazal  end  of  the  seed). 
—  Cliamaenerium  is  a  Willow-herb  with  zygoraorphic  flowers. — The 


MYETIFLOR^.  485 

following  may  be  included  here  :  Clarkia,  Eucharidium  (an  (Enothera  with  4 
stamens  and  3-lobed  petals),  Godetia  and  JBoisduvalia,  Jussicea  (dehiscence 
septicidal),  Isnardia  (petal-stamens  absent,  sometimes  the  petals  also).— Lopezia 
has  a  peculiar,  zygomorphic  flower  (Fig.  521) ;  one  of  the  four  sepals  is  bent  for- 
wards and  the  other  3  backwards ;  the  posterior  petals  are  narrower  than  the 
2  anterior  ones  which  are  turned  obliquely  backwards  and  bent  like  a  knee,  with 
a  greenish  nectary  at  the  bend;  2  stamens,  one  only  fertile  (the  posterior), 
while  the  anterior  is  barren,  petaloid,  and  spoon-shaped ;  both  are  sensitive, 
which  is  essential  for  pollination.  In  Fig.  521,  a  represents  an  early  stage, 
in  which  the  stamen  and  style  lie  concealed  in  the  staminode  ;  6  is  the  $  stage, 
the  stamen  projects  from  the  centre  of  the  flower;  c,  the  $  stage,  the  style 
occupies  the  place  of  the  stamen. 

B.  Fruit  a  berry.     Fuchsia  generally  has  a  coloured  calyx 
and  tubular  receptacle  ;  the  corolla  may  be  wanting. 

C.  Fruit    a  nut.     Girccea  (Enchanter's   Nightshade)  has    a 
2-merous  flower  (S2,  P2,  A2  +  0  [petal- stamens  are  wanting],  G2). 
The  flowers  are  borne  in  racemes  without  bracts. — Gaura. 

D.  Fruit  a  drupe.     Trapa  (Horn-nut)  ;  a  peculiar  aquatic 
plant ;  the    submerged  stem  has  long   internodes  and  lanceolate 
leaves,  falling  off  at  an  early  period,  but  at  each  node  are  found  4 
long  roots  with  thin,  lateral  roots  (sometimes  erroneously  regarded 
as    leaves)   borne   pinnately;    the   stem   reaching   the  surf  ace -of 
the  water,  bears  a  rosette  of  rhombic  foliage-leaves,  with  large, 
inflated  stalks  containing  air,  and  forming  the  floating  apparatus 
of  the  plants.     In  the  axils  of  the  leaves  (as  in  Gunnera)  8  small, 
stipular  structures  are  present.     The  flowers  are  solitary  in  the  axils 
of  the  foliage-leaves  (S4,  P4,  A4  +  0,  G2),  seravepigynous.    There  is- 
an  8-lobed,  crenate  disc  on  the  free  portion  of  the  ovary ;  one  ovule 
in  each  loculus.     The  fruit  is  a  drupe  with  4  (or  2)  prominent  horns 
(the  persistent  sepals),  which  after  the  pulp  has  decayed  away  bear  a 
series  of  hooks  turned  downwards  on  each  side,  i.e.  sclerenchyma- 
tous  bundles  which  formerly  lay  concealed  in  the  pulp  of  the  sepals. 
The  germination   is  peculiar :    one  of  the  cotyledons  is  large,  and   its  thick 
extremity  remains  in  the  fruit,  the  other  however  is  small  and  is  pushed  out  at 
the  apex  of  the  fruit  together  with  the  radicle  and  plumule ;  the  development  of 
the  root  soon  ceases,  and  the  plumule  usually  grows  into  a  stem  entirely  without 
branches,  similar  to  the  one  described  above,  only  that  1-2  precisely  similar 
shoots  arise  in  the  axil  of  each  cotyledon,  so  that  each  embryo  produces  3-5- 
shoots.— Trapa,  by  its  mode  of  life,  its  1-seeded  fruit,  etc.,  forms  a  transition  to 
Haloragidacece. 

The  large- flowered  forms  are  adapted  for  insect-pollination  and  are  often- 
protandrous,  the  small-flowered  ones  are  homogainous  and  may  pollinate 
themselves.  (Enothera  is  adapted  for  hawk-moths  and  bees. — 330  species ; 
especially  in  temperate  climates,  chiefly  in  the  Northern  Hemisphere.  Epilo- 


486  D1COTYLEDONES. 

bium,  Circcea  are  natives  of  this  country ;  Trapa  is  extinct  in  this  country,  it 
has  been  found  in  a  semi-fossilized  condition  near  Cromer  and  in  bogs  in  Den- 
mark, and  existed  in  Sweden  until  a  few  years  ago  ;  (Enothera  has  been  intro- 
duced from  N.  Am. — A  number  of  N.  Am.  species  are  grown  as  ornamental 
plants  in  our  gardens.  The  seeds  of  Trapa  natans  are  edible,  and  used  as  food 
in  China. 

Order  5.  Haloragidacese.  This  is  a  reduced  form  of  the 
CEnotheracese,  and  principally  differs  from  these  in  the  presence  of 
endosperm  and/ree  styles.  Only  1  ovule  in  each  loculus. — 84  species 
distributed  over  the  entire  globe  ;  the  majority  are  aquatic  plants. 
The  most  advanced  type  is  Myriophyllum  (Water-Milfoil),  with  a 
regular,  epigynous  flower  (S4,  P4,  A4  +  4,  G4),  most  frequently 
diclinous  (monoecious)  ;  the  fruit  is  a  Z-^-partite  schizocarp. 
Aquatic  plants,  most  frequently  with  pectinate,  pinnate  leaves. — , 
Haloragis. — Gunnera  (a  dozen  species  from  the  Southern  Hemisphere)  forms 
the  next  step  in  the  reduction.  Large,  scattered,  rough-haired,  and  softly- 
spined  leaves,  with  small  flowers  in  crowded  inflorescences.  The  flower,  when 
most  complete,  has  S2,  P2,  A2  (petal-stamens)  and  G2,  forming  an  inferior, 
unilocular  ovary  with  1  ovule.  It  is  remarkable  for  the  great  number  of 
stipules  placed  in  transverse  rows  in  the  leaf-axils,  for  the  peculiar  glandular 
organs,  and  for  the  colonies  of  Nostoc,  which  are  found  embedded  in  the 
cortex  as  a  kind  of  parasite.— The  simplest  form  is  Hippuris  (Mare's- 
tail)  with  an  extremely  small,  crenate  or  entire  calyx,  without 
corolla,  and  with  only  one  stamen  and  one  carpel,  forming  an 
inferior,  unilocular  ovary  with  only  one  ovule.  Fruit  a  drupe 
with  thin  pulp.  It  is  an  aquatic  plant  with  creeping,  sympodial  rhizome, 
and  erect  unbrancned  shoots,  bearing  numerous  small,  verticillate  leaves.  The 
small  flowers  are  situated  singly  in  the  leaf-axils. 

Order  6.  Rhizophoraceae.  Tropical  trees  or  shrubs  (50  species,  the  best 
known  being  Hhizophora  mangle,  Mangrove)  which  grow  gregariously,  especi- 
ally along  the  banks  of  rivers  and  by  sea-coasts,  where  the  water  is  quiet  and 
brackish,  and  where  they  form  the  so-called  Mangrove-swamps.  Aerial  roots 
.are  formed  on  the  stems  and  branches  (Fig.  522  A).  The  seeds  germinate  in 
the  fruit,  which  by  arrest  contains  only  one  seed  (Fig.  522  B),  before  it  is 
detached  from  the  tree.  The  radicle  projects  considerably  from  the  seed,  and 
hangs  down  freely  in  the  air ;  when  the  embryo  is  finally  detached  from  the 
mother-plant,  the  separation  is  effected  by  the  hood-like  cotyledon,  which 
entirely  envelops  the  plumule,  becoming  detached  from  tbe  rest  of  the  embryo, 
which  falls  down,  while  the  hood-like  cotyledon  remains  enclosed  in  the  fruit. 
The  embryo,  after  it  has  fallen,  strikes  root,  and  continues  growing  in  the 
undisturbed  mud  under  the  trees,  or  perhaps  it  may  first  be  drifted  about  by 
the  water,  being  well  adapted  for  this  by  its  peculiar,  tough  nature,  and  large, 
intercellular  spaces. — It  may  also  further  be  remarked  that  the  anther  is  divided 
iuto  a  number  of  small  locnli.  The  leaves  are  stipu'ate.  The  endosperm  projects 
from  the  micropyle,  growing  out  from  the  base  of  the  seed,  and  thus  serves  as 
an  organ  of  suction  to  convey  nutriment  to  the  embryo  from  the  mother-plant. 


MYBTIFLOKJ1. 


487 


Order  7.  Combretaceae.  Trees  and  shrubs,  partly  . lianes.  An  inferior, 
unilocular  ovary  with  few  pendulous  ovules.  Conocarpua  and  Laguncularia 
form,  in  conjunction  with  the  species  of  Rhizophoracese,  the  tropical  Man- 
grove-swamps. Terminalia. — 280  species  ;  Tropics. 

Order  8.  Myrtaceae  (Myrtles) .  The  plants  belonging  to  this 
order  are  shrubs  or  trees,  the  majority  being  easily  recognised  by 
the  vegetative  characters.  The  leaves,  for  instance,  are  most  fre- 
quently opposite,  without  stipules,  undivided  and  entire,  parch- 


A  B 

FIG.  522. — RTiuopTiora  mangle  with  the  germinating  fruit  (much  reduced). 

Tnent-like  or  leathery,  evergreen,  aromatic,  finely  dotted  by  pellucid 
•glands  containing  essential  oils  ;  the  venation  is  penninerved  with  a 
nerve  just  inside  and  running  parallel  to  the  edge  of  the  leaf. 
The  flowers  are  regular,  epigynous  (Figs.  523,  524,  525)  and  $, 
most  frequently  4-  or  5-merous  in  the  calyx  and  corolla,  with  many 
stamens  (by  splitting,  so  that  they  are  often  in  several  distinct 
bundles)  and  an  ovary  with  one  style,  formed  of  2-5-many  car- 
pels; the  receptacle  is  most  frequently  united  for  its  entire  length 
with  the  ovary.  The  fruit  varies,  but  is  most  frequently  a  berry. 
The  embryo  is  thick,  often  curved,  with  united  cotyledons;  no 
endosperm. 


488 


DICOTYLEDONES. 


1.  MYRTE^I,  MYRTLE  GROUP.    Chiefly  American,  though  some  are 
found  also  in  Africa  and  Asia.     The  fruit  is  a  berry  with  generally 
2-5  loculi  in  the  ovary,  and  many  ovules  in  each. — Myrtus;  Eugenia 
(the  petals  fall  off  together  as  a  hood  in  the  Clove,  E.  caryophyllata, 
Figs.  523,  524)  ;  Myrcia;  Jambosa ;   Amomis ;  Psidium,  etc. 

2.  PUNICE^,  POMEGRANATE  GROUP.    Only  2  species  (Punica  grana- 
tum ;  from  Persia,  Afghanistan),  differing  in  several  respects  from 
the   typical  form  of   the   Myrtacea?.      The    leaves  are  generally 
opposite,  without  glands  and  marginal  veins.    The  receptacle,  calyx 
and  corolla  are  red  ;  the  latter  5-8-  (generally  6-)  merous.     Calyx 
valvate    and   corolla  folded    as  in  LythraceaB,  stamens    also  and 
epicalyx  as  in  this  order.     The  most  characteristic  feature  is  the 
inferior,   spherical    berry,  with    dry    pericarp,   formed   from   two 


523.  624. 

Ftes.  523,  521. — Eugenia  caryophyllata. 
FIG.  523.— Flowers  (nat.  size). 
FIG.  524. — A  bud  ("  clove  "),  long.  sec.  (mag.). 


FIG.  525.— Punica  granatum.    Flower, 
long.  sec.  (nat.  size). 


whorls  of  carpels  in  two  tiers  (Fig.  525)  ;  the  interior  whorl, 
which  is  also  the  lower,  has  3  carpels,  and  the  placenta?  are  situated 
in  the  inner  angles  of  the  3  loculi ;  the  external  whorl  is  5-merous, 
and  the  placenta?  have  originally  the  same  position  in  the  inner 
angles  of  the  loculi,  but  their  position  is  changed  to  the  outer  side 
of  the  loculi  owing  to  the  growth  of  the  wall  of  the  ovary,  which 
takes  place  early,  causing  the  carpels  to  become,  as  it  were,  turned 
inside  out,  so  that  the  part  which  was  turned  downwards  is  turned 
upwards,  and  the  part  which  was  turned  inwards  becomes  turned 
outwards  (as  in  Mesembrianthemum) .  The  edible  part  of  the 
fruit  is  the  fleshy  testa,  as  in  Ribes.  The  cotyledons  are  rolled 
together  spirally. 


MYRTIFLORE.  489 

3.  LECYTHIDE^E.     The    majority  are    South    American.      The  leaves  are 
scattered,  without  pellucid  glands,  and  frequently  dentate.     The  flowers  are 
zygomorphic.     The  woody  fruits  are  either  indehiscent,  or  open  by  a  lid.     To 
this  belong:  Bertholletia  (B.  excelsa),  the  seeds  well  known  as  "Brazil-nuts," 
LecytJds  (Sapucaia-nuts  from  L.  ollaria),  Barringtonia. 

4.  LEPTOSPERMETE.   Almost  entirely  from  Australia  and  the  East  Asian  and 
Pacific  Islands.     The  fruit  is  a  capsule.     The  leaves  are  scattered,  and  in  some 
placed  edgewise  by  the  twisting  of  the  leaf-stalks. — Eucalyptus,  the  Australian 
Gum-tree  ;  the  calyx  falls  off  like  a  lid  (Figs.  526,  527).     Some  of  the  species 
attain  gigantic  heights,  E.  amygdalina  140-150  m.  with  a  diameter  of  8  m. 
The  leaves  in  E.  globulus  are  opposite  and  dorsiventral  on  the  young  plant ;  on 
the   older   scattered,  placed   edgewise   by  the  twisting  of  the  leaf-stalk,  and 
isolateral ;     Metrosideros,    Calothamnus     (stamens     distinctly    polyadelphous), 
Melaleuca,  Leptotpermum,   Callistemon  (the  flowers  are  borne  in  spikes  whose 
axis  continues  to  grow  after  flowering,  thus  several  zones  of  fruits  may  be  seen 
on  the  same  branch). 


526. 

FIGS.  526,  527.  —  Eucalyptus  gl 


•  Fio.  526.—  Long.  sect,  of  flower.  FIG.  527.—  Flower  opening. 

5.  CHAM^LAUCiEa:.  Australian  shrubs  with  heath-like  appearance;  they 
differ  from  the  other  Myrtacese  in  having  a  unilocular  ovary  with  few,  basal 
ovules,  and  a  1-seeded  nut.  The  sepals  are  often  pappus-like,  and  divided  into 
many  bristles.—  Chamcelaucium,  Darwinia,  etc. 

This  large  order  (2,100  species)  is  confined  almost  entirely  to  the  Tropics, 
being  found  principally  in  America  and  Australia.  In  Europe,  only  Myrtus 
communis.—  Several  are  useful  on  account  of  the  large  quantity  of  volatile  oils 
(contained  in  internal  glands)  :  the  flower-buds  ("Cloves")  of  Eugenia  caryo- 
pliyllata  (the  Moluccas,  cultivated  in  the  Tropics,  Figs.  523,  524)  ;  the  unripe, 
dry  berries  ("  Pimento  ")  of  Myrtus  pimento,  (Pimenta  officinalis,  W.  Indies)  ; 
Cajeput  oil  is  extracted  from  Melaleuca  minor  and  leucadendron  (East  Asian 
Islands).  Eucalyptus  globulus  (Australia)  has  of  late  years  become  well  known 
on  account  of  its  rapid  growth,  its  hard  wood,  and  its  antipyretic  qualities^ 
it  is  cultivated  on  swampy  soils,  which  it  helps  to  drain.-  OFFICINAL  :  "  Cloves," 
and  the  cork  of  both  stem  and  root  of  Punica  granatum.  Several  have  EDIBLE 
FRUITS,  such  as  Psidium  guyava  (Guava,  var.  pomiferum  and  pyriferum,  Am.), 
Eugenia  cauliflora  and  others,  E.  jambosa,  Punica  granatum  (the  Pomegranate), 
etc.  EDIBLE  SEEDS  (with  abundance  of  fatty  oil)  :  "  Brazil  nuts  "  from 
Bertholletia  excelsa  (Trop.  S.  Am.).  "Bay-rum"  is  extracted  from  the  leaves 


490  DICOTYLEDONES. 

and  fruits  of  the  Bayberry-tree  (Pimento,  acris,  W.  Ind.) ;  Guava-rum  from 
the  berries  of  Euyenia  jloribitnda.  Tannin  is  found  in  large  quantities  e.g.  in 
Punica.  Gum  is  formed  by  many  Australian  Eucalypti  ("  Guni-trees "). 
ORNAMENTAL  PLANTS  cultivated  in  this  country  are  :  Myrlus  communis  (Mediter- 
ranean), several  in  conservatories,  especially  the  Australian  Leptospermeae, 
Eucalyptse  and  others. 

Family  24.     Umbelliflorse. 

The  flower  is  regular,  ^  ,  and  completely  epigynous,  5-  or4-merous, 
with  1  whorl  of  stamens  and  5—2  carpels.  Sepals  very  small,  tooth- 
like.  The  corolla  is  polypetalous,  most  frequently  valvate  in  cestivation 
(least  pronounced  in  the  Umbelliferous  plants).  Round  the  base  of 
the  styles,  which  are  generally  free,  there  is  an  epigynous  (undivided, 
or  divided)  nectar-disc  ("  stylar-foot "  :  Figs.  5281?,  C,  D ;  539); 
the  number  of  loculi  in  the  ovary  equals  that  of  the  carpels ;  only 
1  pendulous  (anatropous)  ovule  (Fig.  528  C)  in  each  loculus.  Endo- 
sperm copious  (Fig.  528  D).  To  this  must  be  added  that  the 
inflorescence  in  the  majority  of  cases  is  an  umbel  or  a  capitulum, 
especially  in  the  Umbelliferas  and  Araliacess.  Stipules  are  absent, 
but  most  frequently  the  base  of  the  petiole  forms  a  large  sheath. 

The  Umbelliflorffi  are  on  one  side  so  closely  allied  to  the  Frangulinae, 
especially  Rhamnacese,  that  they  may  perhaps  be  regarded  as  the  epigynous 
continuation  of  this  family.  On  the  other  hand,  the  similarities  to  the  Eubi- 
ales,  especially  those  between  Cornacese  and  Sambucese,  are  so  great  that  there 
is  scarcely  any  character  to  distinguish  them  except  the  polypetalous  corolla 
of  the  former  and  the  gamopetalous  corolla  of  the  latter.  Whether  this  is 
more  than  a  merely  analogous  resemblance,  and  if  not,  whether  the  Cornacese 
at  least  should  not  be  included  in  the  Rubiales,  must  be  left  in  abeyance. — The 
sepals  are  very  small,  as  is  generally  the  case  in  epigynous  flowers. 

Order  1.  Cornaceae.  The  majority  of  the  species  are  shrubs 
with  solid  internodes,  opposite  (rarely  scattered)  leaves,  which  are 
simple,  entire  (rarely  incised),  penninerved,  without  stipules  or 
large  sheaths;  flowers  4-merous  (most  frequently  S4,  P4,  A4,  G2), 
borne  in  dichasia  which  are  either  collected  into  corymbs  (e.g. 
Cornus  sanguinea),  or  in  closely  crowded  umbels  or  capitula 
(Cornus  mas,  C.  suecica),  in  which  latter  case  there  is  often  a  large, 
leafy,  or  coloured,  most  frequently  4-leaved  involucre  round  the 
base  of  the  inflorescence  ;  the  style  is  undivided,  with  lobed  stigma ; 
the  raphe  of  the  ovule  is  turned  outwards.  The  fruit  is  a  berry  or 
.a  drupe,  with  a  1-4-locular  stone  or  2  free  stones. 

Cornus  (Dog- wood,  Cornel)  has  S4,  P4,  A4,  G2.    Leaves  opposite. 


UMBELLIFLOKJ:.  491 

Drupe  with  a  bilocular,  2-seeded  stone.  —  Aucuba,   dioecious;   uni- 
locular  ovary  ;   1  ovule  ;   1-seeded  berrj.  —  Garrya.  —  Helwingia. 

80  species  ;  N.  Temp.  The  fruits  of  Cornm  mas  are  edible  ;  the  wood  is 
very  hard;  gum  is  found  in  some.  Several  species  of  Cornus  and  Aucubce 
japonica  (Japan)  are  cultivated  as  ornamental  shrubs. 

Order  2  Araliacese  (Ivies).  Principally  trees  or  shrubs  with 
solid  stems.  The  leaves  are  scattered,  simple  or  compound,  with  a 
sheath  more  or  less  developed.  The  flowers  are  most  frequently 
situated  in  umbels  or  capitula  which  are  either  borne  singly  or  in 
racemes,  or  in  paniculate  inflorescences.  The  small,  most  fre- 
quently yellowish-green  flowers  are  $-merous,  in  the  calyx,  corolla, 
and  andrcecium  ;  the  gynceceum  may  be  5-merous  or  may  have 
some  other  number  (2-oo).  "  The  styles  are  most  frequently  several, 
free  ;  the  raplie  cf  the  ovules  is  turned  inwards  as  in  the  Umbelli- 
ferous plants.  The  fruit  is  a  drupe  or  berry.  —  Stellate  hairs  often 
occur.  The  petals  generally  have  a  broad  base,  and  a  thick  apex  which  is- 
slightly  incurved,  and  a  distinctly  valvate  aestivation. 

Hedera  helix  (Ivy)  climbs  by  adventitious  roots.  The  leaves  are 
palminerved  and  lobed  on  the  sterile  branches,  but  often  ovate  and 
not  lobed  on  the  flowering  branches.  The  flowers  are  yellowish  -green 
and  open  in  the  autumn  ;  they  are  slightly  protandrous,  and  are  visited  by 
flies  and  wasps.  Berries  black.  Endosperm  ruminate.  —  Panax.  Aralia  (with 


375  species,  51  genera;  especially  in  the  Tropics  (E.  Asia).  —  The  Ivy, 
several  species  of  Aralia,  e.g.  A.  japonica  (Fatsia),  Gastonia  palmata,  are  culti- 
vated as  ornamental  plants.  Paper  is  manufactured  from  the  pith  of  Aralia 
papyri/era  (China). 

Order  3.     Umbelliferae.     The  stem  is  herbaceous  with   hollow 
internodes  ;  the  leaves  are  scattered,  and  have  a  broad,  amplexicaul 
base,  a  large,  most  frequently  inflated  sheath,  and  generally  a  pinnate 
(ofteii  very  much  dissected)  blade.    Entire  leaves  are  found  in  Hydro 
cotyle  vulgaris  ;  Bupleurum. 

The  flowers  are  $  ,  regular,  small,  but  collected  in  compound 
umbels,  that  is,  in  "  simple  umbels,"  which  again  are  borne  in 
umbels  (for  exceptions  see  Hydrocotylew)  ;  the  external  flowers  in 
the  simple  umbel  have  often  subtending  bracts,  which  surround 
the  base  as  an  involucre,  and  may  be  termed  the  small  involu- 
cre-, the  internal  ones  have  no  bracts;  when  involucral  leaves 
are  present  at  the  base  of  the  compound  umbel,  they  may  be 
termed  the  large  involucre. 

The  flower  has  5  sepals^(the  median,  as  usual,  posterior),  5  petals,. 
5  stamens  and  2  carpels  (in  the  median  line)  (Fig.  528).  The 
calyx  is  often  scarcely  indicated.  The  petals  have  a  short  claw. 


492 


DICOTYLEDONES. 


are  most  frequently  obcordate,  or  have  an  incurved  apex  (Fig.  528 
B,  (7),  being  incurved  in  the  bud  ;  they  are  white,  rarely  yellow 
(Fennel  and  Parsnips),  blue  or  red.  The  flowers  are  sometimes 
zygomorphic,  especially  those  on  the  circumference  of  the  umbel, 
and  in  that  case  it  is  the  petal  which  is  directed  outside 
(anterior)  which  is  the  largest,  and  the  two  posterior  are  the 
smallest  {e.g.  Heracleum).  The  stamens  are  incurved  in  the  bud. 
The  2  free  styles  unite  at  the  base  into  the  "  stylar-foot  "  (stylopod), 
a  swollen  nectary  (Fig.  528  -B,  0) ;  the  ovary  is  bilocular,  the  raphe 


FIG.  528. — Caucus  carota  with  flower  and  fruit. 

of  the  ovules  being  directed  inwards.  The  fruit  is  a  schizocarp, 
dividing  into  two  mericarps;  the  plane  in  which  these  separate 
coincides  with  that  of  the  union  of  the  carpels,  and  the  two 
nut-like  mericarps  are  in  most  genera  kept  together  for  awhile  at 
the  top  of  a  thin,  bifid,  or  undivided  stalk  (carpophore)  which  is  in 
direct  continuation  with  the  flower-stalk  (Fig.  537).  Each  mericarp 
has  most  frequently  5  more  or  less  strongly  projecting  ridges,  the 
primary  ridges  (Figs.  530,  532,  534,  535,  etc.),  of  which  3  lie  on 
the  back  of  the  mericarp,  the  dorsal  ridges,  and  2  on  its  edge  near 
the  plane  of  division,  the  marginal  ridges;  five  of  these  (10  ridges 


UMBELLIFLOKE.  493 

in  all  in  the  entire  fruit)  are  placed  opposite  the  calyx-  teeth 
and  the  others  between  them.  In  some  genera  there  are  in 
addition  4  secondary  ridges  to  each  mericarp  between  the  primary 
ones  (Fig.  528  E  :  the  secondary  ridges  bear  the  long  bristles). 
Inside  these  secondary  'ridges,  or  inside  the  grooves  between  the 
primary  ridges,  when  the  secondary  ridges  are  absent,  oil  ducts 
(vittse,  schizogenous  ducts)  are  found  in  the  pericarp,  most  fre- 
quently one  in  each  groove  ;  two  are  also  often  found  on  the 
ventral  side  of  each  mericarp  (Figs.  528  E,  530  oZ,  etc.).  The  seed 
is  most  frequently  united  with  the  pericarp.  The  embryo  is  small 
and  lies  high  up  in  the  large,  most  frequently  horny  endosperm 
(Fig.  528  _D).  —  The  endosperm  does  not  contain  starch,  but  oil, 
and  presents  three  different  forms,  of  important  systematic  value  : 
(a)  those  which  are  quite  flat  on  the  ventral  side  (i.e.  the  side 
turned  towards  the  plane  of  splitting)  (Figs.  528  #,  530,  531,  534, 
etc.)  :  the  majority  of  the  genera,  ORTHOSPERME^  (e.g.  Carum,  Pas- 
tinaca)  ;  (b)  those  in  which  the  endosperm  on  the  ventral  side  is 
provided  with  a  longitudinal  groove,  often  deep  :  CAMPYLOSPERME.E 
{e.g.  Anthriscus)  ;  the  transverse  section  is  nearly  a  crescent  (  Fig. 
532);  (c)  those  in  which  the  endosperm  is  concave  on  the  ventral 
side  (hollow  in  both  longitudinal  and  transverse  sections)  :  CCELO 
SPERMEJ:  (e.g.  Coriandrum)  (Fig.  538). 

The  genera  are  distinguished  first  of  all  by  the  endosperm  and  forms  of  fruit, 
the  ridges  and  oil-ducts  ;  then  by  the  form  of  the  umbel,  the  calyx  and  corolla, 
by  the  absence  or  presence  of  an  involucre,  etc. 

I.  HYDROCOTYLE^E,  PENNY-WORT  GKOUP.  Capitula  OY  simple  umbels 
(all  the  other  groups  have  compound  umbels).  No  oil-ducts. 
Orthospermous.  —  Hydrocotyle  (Penny-  wort). 
The  fruit  is  considerably  compressed  laterally 
(Fig.  529).  .The  calyx-teeth  are  small.  The 
leaves  are  peltate.  —  Didiscus.  —  Sanicula  (San- 
nicle).  The  umbels  are  small,  capitate,  gener- 
ally collected  in  a  raceme;  calyx-teeth  distinct. 
$  and  $  -flowers  in  the  same  umbel.  The 
fruits  are  round,  studded  with  hooked  bristles. 
No  carpophore.  —  Astrantia  has  an  umbel  sur- 
rounded by  a  large,  often  coloured  involucre, 
with  this  exception  it  is  the  same  as  the  pre-  FiQ.52Q.-Hydrocotyie 
ceding,  but  the  fruit  is  slightly  compressed, 
with  5  equal  ridges.  Hacquetia  (Dondia).  — 
Eryngium  (Sea  Holly)  :  leaves  often  thorny.  The  flowers  are  all 


8 


4.M 


DICTOYLEDONES. 


sessile,  the  inflorescence  is  thus  a  capitulum ;  each  flower  is 
often  subtended  by  a  bract,  which  is  thorny  like  the  involucre, 
resembling  the  burrs  of  the  Teasel.  The  sepals  are  large.— 
Lacjcecia  :  one  of  the  loculi  of  the  ovary  is  suppressed. 

2.  AMMIE^E,  CARAWAY  GROUP  (Figs.  530-532).  The  fruit  has 
only  the  10  primary  ridges ;  it  is  usually  short,  almost  spherical  or 
broadly  ovate  and  distinctly  compressed  laterally.  Oil-canals  are 


FIG.  530.— Fruit  of  Carum  petroselinuni  -.  fr  endo- 
sperm ;  oi  oil-ducts. 


FIG.  631.— Pimpineila.    Transverse 
section  of  fruit. 


most  frequently  present.  Orthospermous  (except  Conium). — Cicuta 
(Cow-bane).  Pointed  calyx-teeth.  Glabrous  herbs  with  pinnate  or 
bi-pinnate  leaves.  C,  virosa  has  a  thick,  vertical  rhizome,  divided  by  transverse 
septa  into  many  compartments  ;  the  leaflets  are  narrow,  lanceolate,  and  dentate  ; 
the  large  involucre  is  wanting. — Apium  (Celery).  No  calyx-teeth.  A. 
graveolens,  a  maritime  plant,  has  neither  large  nor  small  involucre  ; 
the  umbels  are  short-stalked  or  sessile. — Carum  (Caraway).  Calyx- 
teeth  small ;  the  large  involucre  is  wanting  or  is  only  few-leaved. 
C.  carvi  (Caraway).  C.  petroselinum,  Parsley  (Fig.  530).  Fal- 
caria-,  Ammi;  Helosciadium ;  Bupleurum  (Hare's-ear)  with,  simple 
leaves  and  yellow  corolla  ;  Pimpineila  (Fig.  531)  ;  Siwm ;  ^Ego- 
podium  (A .  podagraria,  Gout- 
weed)  has  bi-  or  tri-ternate 
leaves,  with  ovate,  dentate 
leaflets ;  the  large  involucre 
is  wanting. —  Conium  is  cam- 
pylospermous  (Fig.  532)  ;  the 
short,  broadly  ovate  fruit  has 
distinctly  projecting,  often 
wavy  crenulate  ridges.  C. 
maculatiim  (Hemlock)  has  a 
round,  smooth  stem  with 

FIG.  532.— Conium  maculotum.     Fruit  entire  . 

and  in  transverse  section.  purplish  Spots. 


UMBELLTFLOR2E. 


495 


3-  ScANDiCEJ].  This  group  has  a  distinctly  oblong  or  linear  fruit- 
which  is  slightly  compressed  laterally,  and  generally  prolonged  up- 
wards into  a  "beak";  wings  absent.  Campylospermous.  Otherwise 
as  in  the  Ammieae. — Anthriscus  (Beaked  Parsley)  has  a  lanceolate 
fruit,  round  on  the  dorsal  side,  without  ridges,  but  with  a  ten- 
ridged  beak. — Scandix  (Shepherd's-needle). — Chasrophyllum  (Cher- 
vil) :  fruit  lanceolate  or  linear  with  low,  blunt  ridges ;  beak  absent 
or  very  short.  C.  temulum  has  a  red-spotted,  hairy  stem. — Myrrhis 
(Cicely)  has  a  short  beak  and  sharp,  almost  winged  ridges,  if. 
odorata  (Sweet  Cicely)  has  very  long  fruits. 

4.  SESELINE^;,  FENNEL  GROUP  (Figs.  533,  534).  The  fruit  is- 
slightly  elliptical  or  oblong,  in  transverse  section  circular  or  nearly 
so,  without  grooves  in  the  dividing  plane ;  only  primary  ridges 
are  present.  Orthospermous. — Fceniculum  (Fennel)  has  yellow 


FIG .  533.— (Enanthe  phellandrium.  Fruit  entire 
and  in  transverse  section,  emb  The  embryo  ; 
ol  the  oil-ducts ;  fr  endosperm. 


Fia.  534. — Fceniculnn\  vulgar* 
Fruit  in  transverse  section. 


petals;  both  involucres  are  wanting;  the  fruit  is  oblong.  The 
ridges  are  thick,  all  equally  developed,  or  the  lateral  ridges  are 
slightly  larger  (Fig.  534). — JEthusa  (A.  cynapium,  Fool's  Parsley)  ; 
the  large  involucre  is  wanting  or  is  reduced  to  one  leaf",  the  small 
involucre  is  composed  of  three  linear  leaves  which  hang  down- 
wards on  the  outer  side  of  the  umbels.  The  fruit  is  spherical- 
ovate,  with  thick,  sharp,  keeled  ridges,  the  lateral  ones  of  which 
are  the  broadest. — (Enanthe  (Dropwort)  ;  the  fruit  (Fig.  533)  has 
usually  an  ovate,  lanceolate  form,  with  distinct,  pointed  sepals  and 
long,  erect  styles ;  the  ridges  are  very  blunt,  the  marginal  ones  a» 
trifle  broader  than  the  others. — Seseli,  Libanotis,  Cnidium,  Siler, 
Silaus,  Meum,  etc. 

W.  B.  K  K 


496 


DICOTYLEDONES. 


5.  PEUCEDANE2E,  PARSNIP  GROUP  (Figs.  535-537).  The  fruit  is 
most  frequently  very  strongly  compressed  dorsally,  with  broad, 
mostly  winged,  lateral  ridges.  Only  primary  ridges.  The  dorsal 
ridges  may  project  considerably,  but  are  not  winged.  Ortho- 
spermous. 


FIG.  535. — An'liangelica  officinalis. 
Transverse  section  of  fruit. 


FIG.  536.— Scorodosmafvetidum. 
Transverse  section  of  fruit. 


a.  The  winged  lateral  ridges  stand  out  from  each  other,  so  that 
the  fruit  appears  to    be  4-winged  (Fig.   535). — Angelica;    Arch- 
angelica  (Fig.  535)  ;  Levisticum  (Lovage). 

b.  The  winged  lateral  ridges  lie   close  together,  and  form  one 
wing  on  each  side  of  the  fruit  (Fig.  536). — Pastinaca  (Parsnip). 
Corolla  yellow.     The  dorsal  ridges  are  very  weak ;  the  oil-ducts 
do  not  reach  quite  as  far  as  the  base  of  the  fruit.     Both  large  and 
small  involucres  are  wanting ;  leaflets  ovate.     Anethum  (Dill)  is  a 
Parsnip  with  more  distinct  dorsal  ridges  and  filamentous  leaflets. 

Peucedanum  (Hog's-fennel)  ;  Ferula  (with 
Scorodosma,  Fig.  536,  and  Narthex)  ;  Dorema. 
— Heracleum  (Cow-parsnip) ;  the  flowers  in 
the  margin  of  the  umbels  are  often  very 
large,  zygornorphic,  and  project  like  rays, 
e.g.  in  H.  sibiricum.  The  fruit  is  very  flat, 
with  very  small  dorsal  ridges  ;  the  oil-ducts 
are  more  or  less  club-like  and  do  not  reach 
as  far  as  the  base  of  the  fruit  (Fig.  537). 
Itnperatoria  ;  Tordylium. 
6.  DAUCE^,  CARROT  GROUP  (Fig.  528).  The  fruit  has  18  ridges, 
i.e.  each  fruitlet  has  5  primary  and  4  secondary  ridges,  the  latter 
being  often  more  prominent  and  projecting  further  than  the 
primary  ones.  The  oil-ducts  are  situated  under  the  secondary 
ridges  (Fig.  528). 

a.    -ORTHOSPERMOTJS  :   Daucus  (Carrot).      The  secondary  ridges 
project  much  further  than  the  primary,  and  bear  on  their  crests  a 


FIG.  537. — Heracleum  sphon- 
dylium.    Fruit. 


UMBELLIFLOR^]. 


497 


series  of  hooked  spines  (Fig.  528  D,  E) ;  these  are  much  longer 
than  the  small  bristles  on  the  primary  ridges.  The  involucral 
leaves  of  D.  carota  (Carrot)  are  numerous  and  deeply  pinnate;  the  inflo- 
rescence contracts  during  the  ripening  of  the  fruit,  and  since  the  external 
umbels  have  longer  stalks  than  the  central  ones,  they  arch  over  them,  and 
the  inflorescence  becomes  hollow.  For  the  terminal  flower,  see  below. 
— Cuminum  ;  Laserpitium  ;  Melanoselinum. 

b.  CAMPYLOSPERMOUS  :  Torilis   (Hedge  Parsley).     The  primary 
ridges  are  covered  with  bristles  ;  the   secondary  ridges  are  not 
very  distinct  on  account  of  the  spines,  which  entirely  fill  up  the 
grooves.     Gaucalis  (Bur  Parsley). 

c.  CCELOSPERMOUS  :  Coriandrum  (Coriander)  has  a  smooth,  spheri- 
cal fruit   (Fig.   538)  with   a   distinct,  5-dentate   calyx,  the  two 
anterior  (i.e.  turned  outward)  teeth  being  generally  longer  than 
the  others  ;  the  two  fruitlets  scarcely  separate  from  each  other 


FIG.  538. — Coriandrum  salivum  :  b  secondary  ridges  j  d  primary  ridges  ;  /  endosperm  ; 

I  embryo. 

naturally;  all  the  ridges  project  only  very  slightly,  the  curved 
primary  ones  least,  the  secondary  ridges  most. 

POLLINATION.  The  flowers  are  adapted  for  insect-pollination ;  they  secrete 
nectar  at  the  base  of  the  styles  ;  individually  they  are  rather  small  and  in- 
significant, but.  yet  are  rendered  conspicuous  by  being  always  crowded  in 
many-flowered  inflorescences.  Protandry  is  common,  sometimes  to  such  an 
extent  that  the  stamens  have  already  fallen  off  before  the  styles  begin  to  develop 
(Fig.  539,  2).  Insect  visits  are  more  frequent  and  numerous  as  the  inflorescences 
are  more  conspicuous.  The  flowers  as  a  rule  are  £  ,  but  £  -flowers  are  often 
found  interspersed  among  the  others  (Fig.  539),  and  the  number  of  th^se 
becomes  greater  on  the  umbels  developed  at  the  latest  period.  A  terminal 
flower,  which  differs  from  the  others  in  form,  and  in  Daucus  carota  often  in 
colour  also  (purple),  is  sometimes  found  in  the  umbel.  The  nectar  lies  so 
exposed  and  flat  that  the  flowers  are  principally  visited  by  insects  with  short 
probosces,  especially  Diptera ;  bees  are  less  frequent  visitors,  and  butterflies 
rare.  — 1400  species  (175  genera) ;  especially  from  temperate  climates  in  Europe, 
Asia,  N.  Am.  About  68  species  in  this  country. 


498  DICOTYLEDONES. 

USES.  A  few  are  cultivated  as  ornamental  plants.  They  are,  however,  useful) 
in  medicine,1  and  for  culinary  purposes  on  account  of  the  essential  oils  and 
gum-resins  which  in  many  are  formed  in  root,  stem,  and  fruit.  The  FRUITS  of 
the  following  are  used:  Carum  carvi  f  (Caraway),  Carum petroselinum  (Pars- 
ley ;  also  the  leaves  and  root ;  its  home  is  the  Eastern  Mediterranean) ; 
Fcenictilum  capillaceum  f  (Fennel ;  S.  Europe) ;  Pimpinella  anisum  f  (Anise ;. 
E.  Mediterranean) ;  Coriandrum  sativum  f  (Coriander ;  S.  Eur.) ;  (Enanthe- 
phellandrium  (Water  Drop- wort );  Cuminum  cyminum  (Point  Caraway ;  Africa; 
cultivated  in  S.  Europe);  Anethum  graveolens  (Dill).  The  LEAVES  of  the  follow- 
ing are  used  as  pot-herbs :  Anthriscus  cerefolium  (Chervil) ;  Myrrhis  odoruta 
(Sweet  Cicely  ;  Orient.);  Conium  maculatum  f  (the  green  portions ;  Hemlock). 
Besides  Parsley,  the  ROOTS  of  the  following  are  used :  Carrot,  Parsnip,  Shim 
xisarum  (Sugar-root;  E.  Asia);  Chcerophyllum  bulbosum  (Chervil-root);  l.evi- 
stictnn  officinale  (foliage-shoots ;  S.  Europe) ;  Imperatoria  ostruthium  ;  Apiuni 
yrai  eolens  (Cflery,  the  root  in  conjunction  with  the  internodes) ;  Pimpinella, 


st 


FIG.  539.—  Anthriscus Silvester :  1  <J -flower;  2  $ -flower 


saxifraga  and  marjna  (Pimpinell) ;  Archangelica  (Angelica,  the  root  of  A, 
iiorvegica  was  formerly  an  article  of  food  in  Norway).  Poisonous  alkaloids  are 
found  in  a  few,  such  as  Fool's  Parsley  (JEthusa  cynapium),  Hemlock  (Conium 
maculatum),  Cow-bane  (Cicuta  virosa)  and  species  of  (Enanthe. — Gum-resin  is 
extracted  from  various  species:  "Galbanum"  from  F erula  galbaniflua  f  and 
rnbricalis  f  (Persia) ;  Asafoatida  from  Ferula  scorodosma  f  and  F.  narthex  f  ; 
Ammoniac-gum  from  Dorema  ammoniacum  f,  all  from  Central  and  S.  W.  Asia. 
"  Silpliium "  was  an  Umbelliferous  plant  which  grew  in  ancient  times  in 
Cyrene,  and  from  which  the  Romans  extracted  a  valued  condiment. 

Family  25.     Hysterophyta. 

This  family  (with  the  exception  of  Aristolochiaceae)  includes 
only  parasitic  plants.  Partly  on  this  ground,  and  partly  because 
they  all  have  epigynous  flowers,  they  are  considered  to  belong  to 
the  youngest  type  (which  is  expressed  in  the  name  vcrrepos,  the 
one  that  comes  after).  It  is  not  certain  to  which  of  the  preceding 
families  they  are  most  nearly  allied.  Again,  it  is  a  matter  of  doubt 

1  Those  marked  f  are  officinal,  and  when  no  home  is  stated,  the  plant  is  a* 
native. 


HYSTEROPHYTA. 


499 


•whether  the  Aristolochiaceffi   are  related  to   the  others ;  they  are  by  Engler 
united  with  Rafflesiacese  into  one  family,  Aristolochiales. 

Order  1.  Aristolochiaceae.  The  majority  are  perennial 
herbs  or  twining  shrubs,  whose  stalked,  simple,  and  generally  more 
or  less  cordate  or  reniform  leaves  are  borne  in  2  rows  and  are  ex- 
stipulate.  The  flowers  are  hermaphrodite,  epigynous,  regular  or 
zygomOrphic;  perianth-leaves  united,  simple  but  most  frequently 
petaloid  and  3-merous  ;  6  or  12  (in  Thottea  as  many  as  36)  stamens 
with  extrorse  anthers.  The  ovary 
is  more  or  less  completely  4-6- 
locular  with  ovules  attached  in 
the  inner  angles  of  the  loculi 
(Fig.  540  kf).  The  style  is 
short,  and  has  a  large,  radiating 
stigma  (Fig.  540  n).  Fruit  a 
capsule.  Seeds  rich  in  endo- 
sperm, 

Asarum  europseum.  Each 
shoot  has  2  reniform  foliage- 
leaves,  between  which  the  ter- 
minal flower  is  borne  (the 
rhizome  becomes  a  sympodium 
by  development  of  the  bud  in 
the  axil  of  the  upper  foliage- 
leaf).  The  flower  ia  regular 
and  has  a  bell -shaped  perianth 
with  3  outer  valvate,  and  3 
inner  small  segments  (which 
may  be  wanting).  12  (2  x  6) 
free,  extrorse  stamens,  6  carpels. 
— Aristolochia  clematitis  (Birth - 
wort)  has  an  erect,  unbranched 
stem,  bearing  many  flowers  in 
the  leaf-axils,  in  a  zig-zag  row 
(accessory  buds  in  a  unipared 
scorpioid  cyme).  The  flowers 

are       zygomOrphic      (Fig.      540),        (long.  sect.).    A  Before  pollination,  and  B 
formed  by  3  alternating,  6-mer-        after :  n  stigma ;  a  anthers  j  t  an  insect ;  Tcf 

ous  whorls.     The  perianth  has 

a  lower,  much-distended   part  (fc),  succeeded   by  a  narrow,  bent 

tube  (r),  which  passes  over  into  an  oblique,  almost  tongue-like  pro- 


500 


DICOTYLEDONES. 


jection  (6  vascular  bundles  indicate  that  the  number  6  is  prevalent 
here,  as  in  Asm-urn)  ;  6  stamens  (Fig.  540  a),  with  the  dorsal  por- 
tion turned  upwards,  are  united  with  the  short  style  to  form  a 
stylar  column  ;  they  are  placed  quite  beneath  the  6  commissural 
stigmatic  rays,  which  arch  over  them  as  short,  thick  lobes. 
Protogynous ;  POLLINATION  is  effected  in  Arist.  clematitis  by  small  flies ;  these 
enter  the  erect  unfertilised  flower  through  the  tube  (Fig.  540  A,  I)  without 
being  prevented  by  the  stiff,  downwardly-turned  hairs  which  line  the  tube  and 
prevent  their  escape ;  they  find  the  stigma  (n)  fully  developed,  and  may  pollinate 
it  with  the  pollen  they  have  brought  with  them.  The  stigmas  then  straighten 
and  wither  (B,  n),  the  anthers  open,  and  the  flies  may  again  be  covered  with 
pollen ;  but  the  hairs  which  blocked  up  the  tube  do  not  wither  until  the 
anthers  have  shed  their  pollen,  and  only  then  allow  the  imprisoned  flies  to 
escape  and  effect  cross-pollination.  Prior  to  pollination,  the  flowers  stand 
erect,  but  after  this  has  taken  place  they  become  pendulous,  and  the  perianth 
soon  withers. — A.  sipho  (Pipe-flower),  another  speciee,  is  a  climber,  and  often 
grown  in  gardens  ;  it  has  only  one  row  of  accessory  buds  in  the  leaf-axils.— 200 
species  ;  chiefly  in  S.  Am.  OFFICINAL  :  the  rhizome  of  Aristolochia  serpentaria 
(N.  Am.). 

Order  2.  Santalaceae.  Parasites  containing  chlorophyll,  which,  by  the  help 
of  peculiar  organs  of  suction  (haustoria)  on  their  roots,  live  principally  on  the 
roots  of  other  plants.  Some  are  herbs,  others  under-shrubs.  The  regular, 
most  frequently  £  -flowers  have  a  simple  perianth,  which  is  gamophyllous,  3-  or 
5-partite  with  the  segments  valvate  in  the  bud,  and  a  corresponding  number 
of  stamens  opposite  the  perianth-leaves.  In  the  inferior  ovary  there  is  a 
free,  centrally  placed,  often  long  and  curved  placenta  with  three  ovules  (one 
opposite  each  carpel) ;  these  are  naked,  or  in  any  case  have  an  extremely  insig- 
nificant integument.  Fruit 
a  nut  or  drupe.  Seed 
without  testa.  Endosperm 
fleshy.  225 species;  chiefly 
in  the  Tropics.— Thesium,  a 
native,  is  a  herb  with  scat- 
tered, linear  leaves  and 
small  5-merous  flowers  (P5, 
A5,  G3)  in  erect  racemes; 
the  subtending  bracts  are 
displaced  on  the  flower- 
stalks.  Fruit  a  nut. — Osyris 
(dioecious  shrub ;  3-rnerous 
flowers)  is  another  Europ- 
ean genus. — Santahim  al- 
bum, which  grows  in  E. 
Ind.,  yields  the  valuable, 
scented  Sandal-wood,  the 


FIG.  6<Jl. — A  fiuit  of  Myiodevdion  Iradiystacliyum 
(slightly  mag.)  germinating  on  a  branch. 


oil  of  which  is  used  medi- 
cinally.— Quinchamalium. 


HYSTEROPHYTA.  501 

Myzodendron  is  a  reduced  form  of  the  Santalacese ;  the  $  -flowers  are  without 
perianth ;  the  perianth  of  the  ?  -flower  is  3-merous.  About  7  species  ;  S.  Am. ; 
parasitic  on  a  Beech  (Nothofagus).  The  fruit  has  3  feathery  brushes,  alternat- 
ing with  the  lobes  of  the  stigma,  which  serve  as  flying  organs  and  to  attach 
the  fruits  to  a  branch  (Fig.  541),  the  brushes  twining  round  as  soon  as  they  come 
in  contact  with  it.  There  is  only  1  seed  in  the  fruit,  which  germinates  by  a  long, 
negatively  heliotropic  hypocotyl,  and  is  attached  by  a  radicle  modified  into  an 
haustoriuru. 

Order  3.  Loranthaceae  (Mistletoes).  Plants  containing 
chlorophyll  which  are  parasites  011  trees,  and  most  frequently 
have  opposite,  simple,  entire  leaves  and  regular,  epigynous, 
often  unisexual,  2-  or  3-merous  flowers,  with  single  or  double 
perianth.  Stamens  equal  in  number  and  opposite  to  the  perianth- 
leaves,  free,  or  in  varying  degrees  united  to  one  another.  The 
inferior  ovary  is  constructed  as  in  the  Santalaceae,  the  ovules  being 
situated  on  a  low,  free,  centrally-placed  placenta,  but  the  placenta 
and  ovules  unite  with  the  wall  of  the  ovary  into  one  connected,  par  en- 
chymatous  mass,  in  which  the  embryo-sacs  are  imbedded.  Only  1 
(less  frequently  2-3)  of  the  1-6  embryo-sacs  is  fertile.  The  num- 
ber of  the  carpels  however  varies.  The  fruit  is  a  l-seeded  berry, 
whose  inner  layer  is  changed  into  a  tough  slimy  mass  (bird-lime), 
which  serves  to  attach  the  fruits  to  other  plants. 

The  two  groups,  Loranthoidece  and  Viscoidece,  are  distinguished  by  the  fact 
that  the  former  has  a  distinct  "  calyculus,"  i.e.  an  entire  or  lobed,  or  dentate 
swelling  on  the  receptacle  below  the  perianth.  The  majority  of  the  Loran- 
thoidese  have  a  petaloid  perianth  ;  in  all  the  Viscoideae,  on  the  other  hand,  it  is 
sepaloid. 

The  Mistletoe  (Viscum  album,  Fig.  542)  is  a  native,  evergreen 
plant  which  may  be  found  growing  on  almost  any  of  our  trees 
'  (sometimes  on  the  Oak),  and,  like  other  Loranthaceae,  ifc  produces 
swellings  of  the  affected  branches.  Its  spherical  white  berries  (Fig. 
542  A)  enclose  (1-)  2-3  green  embryos;  they  are  eaten  by  birds  (especially 
Thrushes),  and  are  partly  sown  with  their  excrement,  partly  struck  or  brushed 
off  the  branches  of  the  trees,  the  seed  being  enclosed,  at  maturity,  by  viscin,  i.e. 
"  bird-lime."  The  seeds  may  also  germinate  on  the  branches,  without  having  first 
passed  through  the  alimentary  canal  of  the  birds.  On  germination,  the  hypo- 
cotyl-axis  first  appears,  as  in  Fig.  541,  and  bends  towards  the  branch ;  the  apex 
of  the  root  then  broadens,  and  forms  at  the  end  a  disc-like  haustorium,  from  the 
centre  of  which  a  root-like  body  grows  through  the  bark  into  the  wood,  and 
ramifies  between  the  bark  and  wood.  Suckers  are  developed  on  the  root  like 
strands  which  are  formed  in  this  manner,  without,  however,  having  a  root  cap; 
they  are  green,  and  penetrate  the  wood  by  the  medullary  rays  (Fig.  542  C). 
Adventitious  buds  may  also  be  developed  from  the  root-like  strands  which  break 


502 


DICOTYLEDONES. 


FIG.  543.— Viscum  album:  A  branch  with  leaves  and  berries:  a  scale-leaves;  b  foliage- 
leaves;  n  m  n  flowers  ;  B  seedling,  the  bark  .of  the  branch  being  removed ;  C  an  older 
embryo  which  still  retains  the  cotyledons. 


HYSTEROPHYTA. 


503 


through  the  bark  and  emerge  as  young  plants.  The  young  stem  quickly  ceases 
its  longitudinal  growth,  and  lateral  shoots  are  developed  from  the  axils  of  its 
foliage-leaves.  These  and  all  following  shoots  have  a  similar  structure  ;  each  of 
them  bears  a  pair  of  scale-leaves  (Fig.  542  A,  a)  and  a  pair  of  foliage-leaves  (Fig. 


FIG.  543.— To  the  left  the  Rafflesiaceona  Cytinus  hypocistus,  parasitic  on  the  roots  of  Cis- 
/      tits.    To  the  right  the  Balanophoi  aceous  Cynomorium  coccineum,  parasitic  on  the  roots  of 
Salicomia. 


504  DJCOTYLEDONES. 

542  A,  &),  and  then  terminates  its  growth,  if  it  does  not  produce  an  inflorescence ; 
new  lateral  shoots  proceed  from  the  axils  of  the  foliage-leaves,  and  the  branch- 
ing, in  consequence,  is  extremely  regular  and  falsely  dichotomous.  Only  one 
internode  (shoot-generation)  is  formed  each  year,  so  that  each  fork  indicates 
one  year.  The  foliage-leaves  fall  off  in  the  second  year.  The  inflorescence  is 
a  3  (-5)  -flowered  dichasium  (Fig.  542  A,  m  is  the  central  flower,  n  the  lateral). 
The  plants  are  dioecious;  the  $  -flower  as  a  rule  is  2-merous :  perianth 
2  +  2,  each  leaf  of  which  bears  on  its  inner  side  6-20  pollen-sacs,  each  of 
which  opens  by  a  pore  ;  this  relationship  may  be  considered  to  have  arisen 
from  the  union  of  the  perianth-leaves  with  the  multilocular  stamens  (2  +  2) 
placed  opposite  them.  The  $ -flowers  always  have  Pr  2  +  2,  G2.—  Loran- 
thus  is  also  found  in  Europe  (it  has  a  3-merous  flower),  especially  in  the 
central  and  south-eastern  districts,  on  Quercus  cerris  and  Q.  pu^escens;  but 
the  great  majority  of  the  520  species  grow  in  the  Tropics  on  trees  which  they 
ornament  with  their  often  brightly-coloured  flowers,  and  ultimately  kill  when 
present  in  too  great  numbers.  The  pollination  in  the  numerous  Loranthaceae 
with  unisexual  flowers,  is  effected  by  the  wind.  In  Viscum  album  this  takes 
place  in  autumn,  the  actual  fertilisation  in  the  following  spring,  and  the 
maturity  in  November  or  December ;  in  the  succeeding  month  of  May  the 
berry  is  ready  to  germinate,  and  falls  off. 
USES.  Birdlime  from  Viscum  album. 

Order  4.  Rafflesiaceae  and  Order  5.  Balanophoraceae.  These  orders  com- 
prise root-parasite*,  almost  entirely  devoid  of  chlorophyll ;  they  are  reddish  or 
yellow,  without  foliage-leaves  (Fig.  543).  As  far  as  our  knowledge  of  these  rare 
tropical  plants  extends,  they  have  thalloid  organs  of  vegetation  resembling  the 
root-like  strands  of  Viscum,  or  they  are  filamentous  and  branched  like  Fungus- 
hyphae ;  they  live  in  and  on  the  tissues  of  the  host-plant,  from  which  their 
flowering- shoots,  often  of  mushroom-like  form,  are  subsequently  developed 
(Fig.  543).  In  order  to  unfold  they  must  often  break  through  the  tissues  of 
the  host-plant. 

Of  the  BAFFLESIACE^E,  Cytinus  hypocistus  is  found  in  S.  Europe  living  on  roots 
of  Cistus -plants  and  to  some  extent  resembling  Honotropa  (Fig.  543).  Rafflesia 
is  the  best  known ;  it  lives  on  roots  of  Ciasus-species  (belonging  to  the  Ampe- 
lidacese)  in  Java  ;  its  yellowish-red,  stinking  flowers  attain  a  gigantic  size  (one 
metre  or  more  in  diameter),  and  are  borne  almost  directly  on  the  roots  of  the 
host-plant.  Besides  these  there  are  other  genera :  Bruf/mannia,  Pilostyles, 
Hydnora. — To  BALANOPHORACE^  (Fig.  543)  belong :  Balanophora,  Langsdorffia, 
Scybalium,  Sarcophyte,  Helosis,  etc.,  and  in  S.  Europe,  Cynomorium  coceineunu 

Sub-Class  2.    Sympetalae. 

The  characters  which  separate  this  from  the  first  Sub-class,  th& 
Choripetalce,  have  been  described  on  page  336.  They  consist  in 
the  following :  the  flower  is  always  verticillate,  generally  with  5. 
sepals,  5  petals,  5  stamens,  and  2  carpels  (in  the  median  plane), 
the  calyx  is  generally  persistent  and  gamosepalous,  the  corolla  is 
gamopetalous  and  united  to  the  stamens,  which  are  therefore 


SYMPETALJ:.  505 

adnate  to  it,  the  ovules  have  only  one  thick  integument  and  a  small 
nucellus.     (The  exceptions  are  noted  later.) 

This  Sub-class  is  no  doubt  more  recent  than  the  Choripetalse  ;   it  is  also 
peculiar  in  including  fewer  trees  and  shrubby  forms  than  the  latter. 

The  Sympetalae  may  be  separated  into  2  sections  : — 

A.  PENTACYCLIC^E  (FIVE-WHORLED).     The  flowers  in  this  section 
have  5  whorls  equal  in  number,  namely,  2  staminal  whorls  in  addi- 
tion to  the  calyx,  corolla,  and  carpels;  in  some  instances,  one  of 
the  staminal  whorls  is  rudimentary  or  entirely  suppressed,  but  in 
this  case  it  is  frequently  the  sepal-stamens  which  are  suppressed, 
and  the  whorl  which  is  present  stands  opposite  the  petals.     The 
flowers   are    regular.      The  number  of  carpels   equals    that  of  the 
sepals,  but  in  one  of  the  orders  (Bicornes)  they  are  opposite  the 
petals    (the    flower   being  obdiplostemonous)  ;    in  the    other  two 
orders   (Primulinse  and  Diospyrinse)  they  are  placed  opposite  the 
sepals  (the  flower  being   diplostemotfous) .      This    section  is   the 
most  closely  allied  to  the  Choripetalae,  since  the  petals  may  some- 
times be  found  entirely  free,  and  the  stamens  inserted  directly  on 
the  receptacle  (Ericacea?)  ;  ovules  with  two  integuments  are  also 
found.      It  is  very  doubtful,  whether  the  orders  included  under  this  head  have 
any  relationship  with  the  other  Sympetalae.     They  appear  in  any  case  to  repre- 
sent, older  types. 

B.  TETRACYCLIC^:  (FODR-WHORLED).     The   flowers   have  only  4 
whorls,  namely,  beside  sepals,  petals,  and  carpels,  only  one  whorl 
of  stamens,  which  alternates  with  the  petals ;  there  is  no  trace  of 
the  second  staminal  whorl,  and  when  the  number  of  carpels  is  the 
same  as  that  of  the  preceding  whorls  ("  isomerous  ")  they  alternate 
with  the  stamens  ;  but  in  most  cases  there  are  2  carpels  placed  in  the 
median  plane. (see  the  diagrams,  e.g.  Figs.  559,  567,  583,  590,  etc.). 
This  section  is  the  largest,  and  the  one  which  shows  the  character- 
istics of  the  Sympetalae  best.    Yery  irregular  flowers  are  met  with. 

The   following   families   belong    to   the     Pentacyclicse :     26, 
Bicornes  ;  27,  Diospy rin30;  28,  Primulinse. 

The  remaining  families  belonging  to  the  Tetracyclicse  are  : — 

a.  HYPOGYNOUS  flowers  (with  a  few  exceptions)  :  29,  Tubiflords; 
30,  Personate;  31,  Nuculiferds ;  32,  Contorts. 

b.  EPIGYNOUS  flowers:   33,  Rubiales ;    34,  Dipsacales ;  35,  Cam- 
panulinx;    36,  Aggregate.     The    ovaries  and    ovules    in    the  last 
family  are   always  reduced   to   one;    and    at  the  same    time  the 
fruits   become   nuts,   and   the  flowers    are    united   into    crowded 
inflorescences. 


506  DICOTYLEDONES. 

A.     Pentacyclicae. 
Family  26.     Bicornes. 

This  family  is  chiefly  composed  of  shrubs,  less  frequently  of 
small  trees,  or  perennial  herbs;  their  leaves  are  undivided,  most  fre- 
quently evergreen,  stiff  and  leathery,  and  always  without  stipules. 
The  flowers  are  $  and  regular,  rarely  slightly. zygomorphic,  most 
frequently  obdiplostemonous,  and  4-  or  5-merous  through  all  the 
5  whorls.  The  stamens  are  attached  to  the  receptacle,  and  as  a  rule 
are  quite  free  from  the  petals,  an  attachment  which  is  very  rare 
among  the  Gamopetalae.  They  have  a  simple  gynceceum  with  one 
undivided  style,  a  commissural  stigma,  and  a  multilocular  ovary, 
whose  axile  placentae  project  considerably  into  the  loculi,  and  bear 
a  large  number  of  ovules.  The  placentas  are  sometimes  not  united,  and 
in  consequence,  the  ovary  is  1-locular  with  incomplete  partition-walls,  e.g. 
Pyrola,  Monotropa.  Embryo  straight,  with  endosperm.  The  carpels 
are  placed  opposite  the  petals. 

The  diagram  is  generally  Sn,  Pn,  An  +  n,  Gn,  in  which  n  is  4 
or  5.  To  this  may  be  added,  that  the  corolla  is  in  'most  cases  ganio- 
petalous,  but  in  some  (especially  Pyrolacese)  perfectly  polypetalous  ; 
and  that  the  anthers  usually  open  by  pores,  and  often  have  two  horn- 
like appendages  (hence  the  name  "  Bicornes  ")  (Figs.  545,  546)  ; 
frequently  the  two  halves  of  the  anther  are  also  widely  separated 
from  each  other  at  the  upper  end,  so  that  the  pores  are  placed  each 
one  at  the  end  of  its  own  tube  (Fig.  546)  ;  the  pollen-grains  in 
the  majority  are  united  into  tetrads  (Fig.  542  D). — The  flowers,  as 
a  rule,  are  pendulous  and  borne  in  racemes,  coloured  (red  or 
white),  but  odourless.  When  the  fruit  is  a  capsule,  the  placenta 
with  the  seeds  attached  persists  as  a  central  column.  A  mycorhiza 
occurs  on  many. 

The  majority  of  plants  belonging  to  this  family  inhabit  cold 
and  temperate  countries,  or  high  mountains  in  tropical  regions  ; 
they  prefer  cold  and  dry  or  damp  places  (bogs,  heaths,  etc.). 
Plentiful  in  N.  America. 

Order  1.  Pyrolaceae.  Perennial  herbs ;  petals  most  frequently 
quite  free  from  each  other,  and  falling  off  singly  after  flowering ; 
the  anthers  are  without  appendages,  and  open  by  pores  (Fig.  544), 
or  by  a  transverse  slit.  The  placentae  are  thick.  The  seeds  in 
the  capsule-like  fruit  (loculicidal  dehiscence)  are  exceedingly  small 
and  light,  they  have  a  sac-like  testa  which  loosely  envelops  them, 
an  oily  endosperm,  and  an  extremely  simple  embryo,  which  consists 


BICORNES. 


507 


FIG.  514.—  Pyrola  minor:  A  portions  of  a  young  flower; 
B  the  stigma ;  C  portions  of  an  older  flower  (longitudinal 
section). 


only  of  an  ellipsoidal,  cellular  mass,  without  cotyledons  or  differen- 
tiation into  plumule  and  radicle. 

Pyrola   (Winter-green)   is  green,  and  has    also  large  evergreen 
foliage-leaves.     The  flowers,  5-merous,  are  most  frequently  borne 
in  racemes  without  a 
terminal    flower ;    the  J  € 

anthers    are   extrorse         jjjr  t    \^  c 

in  the  bud  with  the 
pores  in  the  lower  por- 
tion (Fig.  544  A),  but 
they  become  inverted 
at  a  later  period,  so 
that  the  pores  open 
at  the  top  (Fig.  544  C). 
P.  uni flora  has  a  single, 
terminal  flower ;  it  winters 
by  its  roots,  producing 
from  these  in  the  spring  aerial,  quite  unbranched  shoots.  Chimaphila  umbellata. 

Monotropa  (Yellow  Bird's-nest)  is  very  pale  yellow,  without 
chlorophyll,  succulent,  and  has  only  scale-like  leaves  closely  pressed 
upon  the  stem  ;  it  is  a  saprophyte.  The  raceme  has  a  terminal 
flower,  and  is  pendulous  before  flowering.  The  anthers  open  by  a 
semicircular,  transverse  cleft.  M.  hypopitys  reproduces  chiefly  by  root- 
shoots. 

About  30  species,  especially  N.  Europe,  N.  America,  and  N.  Asia. 

Order  2.  Ericaceae.  The  flower  (Fig.  545)  is  hypogynous,  the 
median  sepal  posterior;  corolla,  gamopetalous ;  the  stamens  are 
generally  2-horned,  and  the  fruit  is  a  capsule,  less  frequently  a 
berry  or  drupe.  At  the  base  of  the  ovary  is  a  nectar-secreting 
disc  (Fig.  545  I>).  This  order  comprises  shrubs  or  undershrubs 
(rarely  small  trees),  which  are  evergreen,  and  as  a  rule  have 
densely  crowded  leaves. 

I.  ERICE^;,  HEATH  GROUP.  Flowers  most  frequently  4-merous 
(S4,  P4,  A4  +  4,  G4,  united  in  a  4-locular  gynoeceum),  rarely  5- 
merous.  The  withered  corolla  persists  after  flowering.  The  leaves 
are  most  frequently  acicular,  opposite  or  verticillate;  the  buds  are 
without  scales.  The  fruit  is  a  capsule. — Calluna  (C.  vnlgaris, 
Ling)  has  a  deeply  4-cleft  corolla,  which  is  less  than  the  coloured 
calyx ;  capsule  with  septicidal  dehiscerice. — Erica  (about  420 
species ;  E.  tetralix,  Cross-leaved  Heath)  has  a  tubular  or  bell- 
shaped,  4-dentate  corolla,  which  is  much  longer  than  the  calyx. 
Capsule  with  loculiciJal  dehisccnce. — Pentapera. 


508 


DICOTYLEDONES. 


2.  ANDROMEDEJE.  The  flowers  are  5-merous  (S5,  P5,  A5  +  5,  G5), 
with  deciduous  corolla.  Capsule  with  loculicidal  dehiscence.  The 
leaves  are  scattered,  and  incline  more  to  the  ordinary  broad-leaved 
forms. — Andromeda;  Gaultheria;  Cassandra  (Lyonia) ;  Cassiope. 


FIG.  545. — Arctostaphylos  uva-ursi. 

3.  ARBUTE.E.  The  flowers  as  in  the  preceding  group  (Fig.  545), 
but  the  fruit  is  a  berry  or  drupe.  Arctostaphylos  (A.  uva-ursi,  Bear- 
berry)  has  a  drupe  with  5  stones  in  a  dry,  farinaceous  pulp  ;  in 
other  species  there  is  1  stone  with  several  loculi.  Arbutus  {A. 
unedo,  Strawberry-tree)  has  a  spherical  berry. 

Pollination  is  effected  by  means  of  insects,  especially  by  bees.  The  pollen 
is  light  and  dry,  and  is  shaken  out  through  the  pores  of  the  anthers  when  the 
insects  agitate  the  horn-like  appendages  during  their  visits.  Self-pollination 
takes  place,  no  doubt,  in  many  cases. —  800  species;  the  very  large  genus, 
Erica,  especially  in  S.  Africa  (the  Cape). — OFFICINAL  :  the  leaves  of  Arctosta- 
phylos uva  ursi.  Arbutus  unedo  (S.  Europe)  has  an  edible,  peculiarly  warted 
.(strawoerry-like)  fruit.  Many  .En'ca- species  are  cultivated  as  ornamental  plants. 

Order3.  Rhodoraceae  (Rhododendrons).  This  differs  from 
the  preceding  order  in  the  median  sepal  being  anterior,  and  hence 
the  position  of  the  other  floral  whorls  is  also  reversed.  The  flower 
is  hypogynous,  in  most  cases  5-merous ;  the  corolla  is  most  fre- 
quently deeply  cleft  or  polypetalous,  and  falls  off  after  flowering ; 
the  anthers  open  by  pores,  and  have  no  horn-like  appendages. 
Capsule  with  septicidal  dehiscence. — The  shrubs  or  small  trees 
belonging  to  this  order  have,  like  the  Vaccinese,  ordinary  foliage- 
leaves,  and  the  buds  are  generally  provided  with  large  bud-scales. 

Rhododendron  has  10  stamens,  and  a  slightly  zygomorphic  flower 
•with  deeply  5-cleft  corolla  (the  section  Azalea  has  frequently 


BlCORNESv  509 

only  5  stamens,  the  petal-stamens  being  absent).  They  are  Alpine 
plants  (200  species)  in  the  mountains  of  Asia,  especially  the 
Himalayas  ;  some  in  S.  Europe. — Menziesia. — Ledum  ;  small,  rusty- 
brown,  hairy  shrubs  with  polypetalous,  expanded,  star-like  corolla. 
— Kalmia  (N.  Am.)  has  a  cupular  corolla,  with  10  small,  pocket- 
like  depressions  in  which  the  anthers  are  concealed  until  the 
arched,  elastic  filaments  are  freed  from  this  position  by  means 
of  the  insects,  when  they  quickly  straighten  themselves  in  the 
centre  of  the  flower. — Phyllodoce ;  Loiseleuria  (5  stamens)  ; 
(Clethra  (?)  ;  also  placed  among  the  Ternstroamiacece). 

About  270  species.  Several  species  are  ornamental  plants.  Several  plants 
of  the  order  are  more  or  less  narcotic.  Ledum  palustre  has  been  used  as  a  sub- 
stitute for  hops. 

Order  4.  Diapensiaceae.  Hypogynous  flower.  3  floral-leaves  beneath 
the  flower  (So,  P5,  A5+0,  G3).  Stamens  on  the  throat  of  the  corolla. 
Pollen-grains  single.  Disc  absent.  Capsule  loculicidal. — 9  species  from  the 
Arctic  regions.  It  is  doubtful  whether  this  order  should  be  included  in  the 
Bicornes  ;  perhaps  it  would  be  more  correctly  assigned  to  the  Polemoniacccs. 

Order  5.  Epacridacese.  This  order  comprises  those  species  of  the  family 
which  are  confined  to  Australia  and  the  South  Sea  Islands.  They  are  shrub-like 
plants,  resembling  the  Ericacere  in  habit,  in  the  inflorescence,  and  in  the 
structure,  form,  and  colour  of  the  flower.  They  differ  especially  in  having 
only  1  whorl  of  stamens  (placed  opposite  the  sepals)  and  in  the  anthers  having 
only  2  loculi,  and  opening  by  a  longitudinal  slit.  Fruit  most  frequently  a 
drupe  (or  loculicidal  capsule).  Epacris-  and  Styphelia-  species  are  ornamental 
plants.  About  325  species. 

Order  6.  Vacciniaceae  (Bilberries).  The  flower  (Fig.  546) 
is  epigynous,  the  corolla  gamopetalous,  and  the  fruit  a  berry.  The 
latter  is  most  frequently  spherical,  and  bears  on  its  apex  the 
calyx,  which  is  generally  very  low,  almost  entire,  and  with  a  disc- 
like  expansion  inside.  The  flower  is  4-  or  5-merous  (Fig.  546  B,  D). 
The  anthers  have  2  pores,  and  are  most  frequently  2-horned 
(Fig.  546  F,  G).  Small  shrubs ;  the  leaves  are  scattered,  not 
needle-like. 

Vaccinium  (Bilberry,  Whortleberry)  has  an  urceolate,  gamo- 
petalous, only  slightly  dentate  corolla,  and  horn-like  appendages 
to  the  anthers  (Fig.  546).  V.  vitis  idcea  (Cowberry)  is  evergreen,  with 
flowers  in  racemes,  and  bright  red  berries;  V.  myrtillus  (Bilberry)  and  V. 
uliginosum  (Bog  Whortleberry)  both  have  black  berries  with  a  blue  bloom,  leaves 
deciduous. —  Oxy coccus  has  a-  polypetalous  corolla  with  the  petals 
projecting  backwards.  Anthers  without  appendages.  0.  palustris 
(Cranberry)  has  a  slender,  creeping  stem,  and  is  evergreen.  Dark  red  berry. 


510 


DICOTYLEDONES. 


Pollination  essentially  the  same  as  the  preceding  order. — 320  species  ;  es- 
pecially in  N.  Am.  Some  are  useful  on  account  of  their  edible  fruits,  especially 
Vaccinium  myrtillus  and  V.  vitis-idaa,  and  in  a  less  degree  Oxycoccus,  etc. 
The  fruits  of  V.  myrtillus  are  officinal. 


FIG.  546. — Vaccinium  uliginosum  (var.  •micropTiyllum).  The  parts  of  the  flower  A-E  are 
enlarged  5-6  times  ;  C  and  E  are  longitudinal  sections  ;  B  and  D  the  flower  seen 
from  above  ;  F  and  G  a  stamen  seen  irom  the  back  and  front  ;  H  the  style  and 
stigma. 

Family  27.     Diospyrinse. 

The  flowers  are  regular,  gamopetalous,  typically  diplostemonous, 
with  the  same  number  throughout  all  5  whorls,  thus  :  Sn,  Pn, 
An  +  n,  Gn,  where  n  most  frequently  =  5  (4-6),  rarely  3,  7  or  8. 
Of  the  two  whorls  of  stamens  the  one  opposite  the  sepals  is  often 
present  only  as  rudiments  or  is  entirely  suppressed,  and  the  com- 
pletely developed  stamens  are  thus  placed  opposite  the  petals.  The 
carpels  are  generally  placed  opposite  the  sepals.  The  ovary  is 
multilocular  with  the  ovules  attached  in.  the  inner  angles.  The 
fruit  is  most  frequently  a  berry.  The  seeds  are  large,  generally 
solitary,  or  a  few  in  each  loculus. — -AH  plants  belonging  to  this 
family  are  trees  or  shrubs  with  scattered,  single,  'most  frequently 
entire,  penninerved  and  leathery  leaves  without  stipules ;  the  ma- 
jority are  tropical  (America,  Asia),  some  are  found  in  N.  Am.  and 
the  Mediterranean. 

Order  1.  Sapotaceae.  Plants  with  latex;  anthers  extrorse,  1  erect  ovule 
in  each  loculus ;  fruit  a  berry  ;  the  seeds  with  bony,  shiny  brown  testa  have  a 
large,  lateral  hilum.  The  leaves  are  frequently  covered  with  silky  hairs. — A 


DIOSPYRIN.E. — PRIMULINJE.  511 

useful  order  in  several  respects  (400  tropical  species).  The  wood  of  some 
geuera,  such  as  Sideroxylon  (Iron-wood)  and  Bumelia,  is  as  hard  as  iron.  The 
latex  of  Palaquium  (P.  oblong i folium,  P.  gutta,  and  other  species),  Mimmops 
and  Payena  (Sumatra,  E.  Ind.),  is  the  raw  material  of  gutta  percha.  The 
following  have  very  delicious  fruits  :  Lucuma  mammosa,  Achras  sapota, 
Chrysophyllum  cainito  (Star-apple),  etc.  The  seeds  of  Bassia  (E.  Ind.)  contain 
a  large  quantity  of  a  fatty  oil.  Isonandra,  Mimmops  schimperi  are  often  found- 
in  the  Egyptian  royal  tombs. 

Order  2.  Ebenaceae.  Plants  without  latex,  often  dioecious ;  flowers  with  a 
more  or  less  leathery  perianth.  The  number  of  stamens  is  sometimes  increased 
(by  splitting  ?  ) ;  ovules  1-2,  pendulous  in  each  loculas.  Fruit  a  berry.  —250- 
species  ;  chiefly  tropical.  Some  are  well  known  on  account  of  their  hard  and 
black-coloured  heart-wood,  e.g.  Maba  ebenus  (the  Moluccas)  and  Diospyros 
ebe.num  (Ebony-wood,  from  Tropical  Asia)  and  others. — The  fruits  are  edible 
e.g.  of  Diospyros  lotus  (Date-plum,  Asia),  which  is  also  cultivated  as  an  orna- 
mental shrub,  together  with  several  other  species. 

Order  3.  Styracacese.  The  flower  is  more  or  less  epigynous,  and  the  corolla 
is  almost  polypetalous.  The  stamens  (by  splitting  ?)  are  more  than  double  the- 
number  of  the  petals,  and  often  united  at  the  base.  Stellate  hairs  are  frequent. 
— 235  species ;  Tropical  Asia  and  America,  a  few  for  example  in  the  East. — 
OFFICINAL:  Gum  -  benzoin  from  Sty  rax  benzoin  and  perhaps  other  species- 
(Sumatra  and  Siam).  Halesia  tetraptera  (N.  Am.)  is  an  ornamental  shrub- 
with  4-winged  fruits. 

Family  28.     Primulinse. 

The  flowers  are  regular,  § ,  hypogynous,  and  gamopetalous. 
The  stamens  are  equal  in  number  to  the  petals  (Fig.  547)  and  are 
placed  opposite  to  them.  The  ovary  is  unilocular,  with  a  free, 
central  placenta  with  1-many  ovules. — The  flower  is  a  further  de- 
velopment of  the  Diospyrinae ;  the  suppression  of  the  calyx- 
stamens,  which  commenced  in  this  family, 
is  carried  further  in  the  Primulinse,  so  that 
in  the  majority  of  cases  no  trace  of  them 
is  present,  but  in  certain  species  and  genera 
(Samolus,  Lysimachia  tliyrsiflora,  Soldanella, 
certain  Myrsinege)  some  small  bodies  (scales, 
teeth,  etc.)  are  found  in  the  position  of  the 
suppressed  stamens.  Again,  the  lateral 
portions  of  the  carpels  are  suppressed,  so 
that  the  ventral  placentce  with  the  ovules  PIG.  54,7.— Diagram  of 
are  separated  from  the  dorsal  portions,  and 

are  united  into  a  free  central  placenta ;  this  theory  is  supported  by 
the  branching  of  the  vascular  bundles,  the  development,  and  various 
comparative  considerations. — Sn,  Pn,  AO  +  n,  Gu;  where  n  =  4-8, 
generally  5.  The  carpels  are  placed  opposite- the  sepals  (Fig.  547), 

w.  B.  L  L 


512 


DICOTYLEDONES. 


li 


FIG.  518.—  Primula:  dimorphic  flowers. 
A  short-styled;  B  long-styled. 


Order  1.  Primulacese  (Primroses).  This  order  has  many 
ovules  attached  to  a  thick,  free,  central  placenta  (Fig.  547)  ;  style 

undivided  with  a  capitate  stig- 
ma ;  ovules  semi-anatropous ; 
fruit  a  capsule  with  many  seeds. 
All  the  plants  belonging  to 
this  order  are  herbs ;  stipules 
wanting  ;  the  flower  is  most 
frequently  5-merous  (S5,  P5, 
AO  +  5,  G5  ;  except  Centunculus 
and  Trientalis}.  The  corolla 
and  capsule  have  various  forms, 
but  the  capsule  generally  opens 
by  teeth  at  the  apex.  The 
ovules  are  semi-anatropous  (in 
Hottonia  they  are  anatropous), 
and  the  seeds  are  therefore 
peltate,  with  the  hilum  situated  in  the  centre  of  one  side.  The 
endosperm  is  fleshy  or  horny.  The  flowers  are  borne  either  in 
racemes  or  in  umbels;  as  bracteoles  are  typicallij  absent  (Fig.  547), 
cymose  branching  does  not  occur. 

Primula  (Primrose)  has  most  frequently  a  vertical  rhizome, 
bearing  a  rosette  of  leaves  at  its  summit,  arid  long-stalked  umbels; 
corolla  rotate  or  slightly  funnel-shaped ;  the  capsule  opens  at 

the  apex  by  5  teeth. 
The  flowers  in  some 
species  are  hetero- 
styled  (long- styled 
or  short-styled  ;  Fig. 
548).  Closely  allied 
are  Androsace  (with 
ovate.  cup -shaped 
corolla-tube  and 
ligular  scales,  alter- 
nating with  the 
corolla -lobes)  and 
Soldanella  (funnel- 
shaped  corolla  with 

FIG.  549.— Cyclamen  persicum.  ,       ... 

lacmiate    lobes    and 

most  frequently  ligular  scales). — Hottonia  (Water- Violet)   is   an 
aquatic  plant  with  pectinate    leaves  and    heterostyled  flowers.-- 


PRIMUUX2E. 


513 


Cortusa.  Dodecatheon.  Cyclamen  (Fig.  549)  has  solitary,  long- 
stalked  flowers,  and  a  rotate  corolla  with  the  lobes  reflexed  ;  the 
stalk  of  the  capsule  rolls  up  spirally;  the  tuberous  rhizome  is 
formed  by  the  hypocotyledonary  internode.  Only  1  cotyledon. — 
Lysimachia  (Money-wort) ;  stem-internodes  well  developed,  leaves 
opposite  or  verticillate,  calyx  almost  polysepalous,  corolla  deeply 
5-partite  (Fig.  550).  The  flowers  are  solitary  or  in  racemes. — 
Anagallis  (Pimpernel),  leaves  opposite,  flowers  solitary;  the  fruit  a 
pyxidium  (Fig.  551)  ;  similarly  in  Centunctdus,  which  is  4-merous. 
— Trientalis,  the  flowers  are  most  frequently  7-merous. — Glaux 
(Sea  Milk-wort)  is  a  creeping  maritime  plant  with  opposite  leaves ; 
flowers  solitary  in  the  leaf-axils,  corolla  absent,  but  with  coloured 
calyx.  The  petals  are  usually  developed  later  than  the  stamens  in  the  Prium- 
laceae  ;  but  in  this  instance  they  are  entirely  suppressed. — Samolus  (Brook- 


FIG.  5rO. — Lysimachin  t/tj/rst/tora. 


FIG.  551. — Anaqallis  arvensis. 
Fruit  dehiscing. 


weed)  differs  from  all  the  others  in  having  an  epigynous  flower; 
barren  sepal-stamens  are  also  present.  The  bracts  in  the  race- 
mose inflorescences  are  displaced  along  the  flower-stalks. 

POLLINATION.  Insect-pollination  in  the  majority;  cross-pollination  is  pro- 
moted in  some  by  heterostyly  (Fig.  548). — 300  species  ;  especially  in  northern 
temperate  zones ;  the  majority  on  mountains  (Soldanella,  Androsace,  etc.) ; 
almost  absent  in  the  Tropics.  A  large  number  are  ORNAMENTAL  PLANTS,  e.g. 
Primula  auricula  (from  the  Alps),  P.  sinensis  (China),  P.  elatior  (Oxslip,  a 
native)  and  grandijiora,  etc.  Cyclamen  europeeum  (Alpine  Violet) ;  the  tubers 
are  poisonous. 

Order  2.  Myrsinacese.  Trees  or  shrubs ;  evergreen,  tropical  Primulacese 
with  fleshy  fruits  and  few  seeds,  embedded  in  the  placenta.  The  leaves  are 
nearly  always  dotted  with  yellow  glands  (schizogenous  resin-receptacles). — 
550  species  ;  especially  Am. — ORNAMENTAL  PLANTS  :  Ardisia  crenulata  (W.  Ind.) ; 
other  genera:  Clauija,  Haesa,  Theophrasta  (barren  sepal- stamens),  Myrsine, 
Jacquinia  (barren  sepal-stamens),  etc. — ^giceras,  allied  to  this  order,  comprises 
arborescent  plants,  often  growing  with  Bhizophora  in  tropical  forests,  along 
the  shore.  The  embryo  germinates  while  still  in  the  fruit. 


514  DICOTYLEDONES. 

Order  3.  Plumbaginaceae.  This  order  has  a  position  of  the 
stamens  similar  to  that  in  Primulaceae  (S5,  P5,  AO  +  5,  G5),  but  it 
differs  from  these  in  the  flower,  which  has  generally  a  membranous, 
dry,  thin,  coloured,  folded,  almost  entire  calyx  and  an  almost 
entirely  polypetalous  corolla,  which,  as  a  rule,  has  twisted  aestivation 
and  is  only  united  with  the  stamens  at  its  base ;  but  more  especially 
it  differs  in  the  ovary,  which  bears  5  free  or  almost  free  styles  and 
only  1  basal  ovule  with  a  long,  twisted  funicle  (the  placenta  of  the 
Primulaceae  is  here  so  much  reduced  that  it  bears  only  1  ovule). 
The  fruit  is  a  nut  or  capsule.  The  radicle  is  turned  outwards. 
Endosperm  mealy. — To  this  order  belong  herbs  or  under-shrubs, 
which  are  especially  natives  of  the  sea-coast  and  of  salt-steppes  ; 
they  also  resemble  the  Primulaceae  in  the  scattered,  undivided, 
entire  leaves  (without  stipules),  often  in  rosettes,  and  the  inflor- 
escence borne  on  a  long  stalk.  In  opposition  to  the  Primulaceae, 
the  bracteoles  are  typically  present,  and  hence  the  branching  is  gene- 
rally cymose  (scorpioid). 

Armeria  (Thrift)  has  a  round  capitulum,  composed  of  closely- 
packed  dichasia,  surrounded  at  its  base  by  an  involucre  with 
peculiar  prolongations,  directed  downwards,  and  united  into  a 
sheath  protecting  the  intercalary  zone  of  growth.  The  pericarp 
is  finally  ruptured  at  the  base,  and  drops  off  like  a  hood. —  In 
Statice  (Sea-lavender),  the  unipared  scorpioid  cymes  are  pro- 
longed and  collected  into  panicle-like  inflorescences. — Plumbago  is 
the  genus  which  approaches  nearest  to  the  Brimulacese,  and  differs  most 
from  the  characters  given  above.  It  has  capitate  or  spike-like  inflorescences,  a 
salver- shaped  corolla,  and  the  stamens  are  not  attached  to  the  corolla.  The 
style  is  only  divided  at  the  extremity ;  the  calyx  is  not  membranous,  but  is 
covered  with  sticky,  glandular  hairs. 

250  species  ;  chiefly  in  the  Mediterranean  and  about  the  Caspian  Sea,  on  salt- 
steppes  and  beaches.  Some  are  Tropical ;  a  few  are  ornamental  plants. 

B.  Tetracyclicse. 

a.  Tetracyclicse  with  hypogynous  flowers. 
Family  29.  Tubiflorae. 

The  flower  is  regular,  §  ,  and  hypogynous.  The  gamopetalous 
type  is  present  in  this  family  with  great  uniformity,  without  sup- 
pression or  splitting ;  S5,  P5,  A5,  G2  (3-5).  The  stamens  are  all 
fertile, alternating  with  the  lobes  of  the  corolla.  Gynosceum  with  2, 
more  seldom  3-5  syncarpous  carpels.  Style  nearly  always  simple  ;. 
2  dorsal  stigmas.  In  each  carpel  2-oc  ovules.  At  the  base  of  the 


TUBIFLORJ;.  515 

ovary  is  found  a  yellowish  ring-like  nectary  (Fig.  552  C),  sometimes 
5-sinuate  or  5-partite. — The  leaves  are  nearly  always  scattered ; 
stipules  are  absent.— The  Solanacese,  which  formerly  were  classed  here,  are 
so  closely  allied  to  the  Personatse,  that  it  would  be  unnatural  not  to  place  them 
first  in  this  family  ;  and  the  Boraginaceee  (which  were  also  placed  in  the 
Tubifloras)  appear  to  be  best  united,  with  the  Labiatse  and  others,  into  one  family 
Nuculiferae. 

Order  1.  Polemoniaceae.  The  flowers  are  regular  ;  S5,  P5,  A5,  G3.  The 
calyx  and  corolla  have  united  leaves,  the  petals  tioisted  to  the  right  in  (estiva- 
tion (all  the  left  edges  being  covered).  The  ovary  is  3-locular  with  2-oo  ovules  in 
each  loculus ;  the  style  is  trifid  at  the  apex ;  the  fruit  is  a  3-valved  capsule. 
Embryo  straight;  endosperm  fleshy.  The  inflorescences  are  dichasia  passing 
over  into  unipared  helicoid  cymes  (the  shoot  of  the  loiver  bracteole  being  the 
more  strongly  developed).— Herbs  without  latex.  150  species  ;  especially 
Western  N.  Am. — Phlox  (salver-shaped  corolla  ;  entire,  opposite  leaves),  Pole- 
monium  (campanulate  or  almost  rotate  corolla;  scattered,  pinnate  leaves), 
Leptosiphon,  Gilia,  Collomia,  Cobcea  (climbing,  like  the  Vetches,  by  tendrils  at 
the  ends  of  the  leaves),  etc.  They  are  frequently  ornamental  plants. 

Order  2.  Hydrophyllacese.  This  order  approaches  very  closely  to  the 
Boraginaceae.  Herbs  with  pinnate  or  palmate  leaves;  S5,  P5,  A5,  G2 
The  lobes  of  the  corolla  are  imbricate  in  aestivation.  Generally  2  median 
carpels.  The  ovary  is  most  freqiiently  unilocular,  and  the  seeds  are  situated 
on  2  parietal  placenta;  capsule  2-valved  ;  embryo  straight ;  endosperm  fleshy. 
In  the  corolla-tube,  opposite  the  corolla-lobes,  there  are  frequently  append- 
ages of  various  forms,  which  resemble  those  of  Cuscuta.  The  inflorescences 
correspond  exactly  with  those  of  the  Boraginaceas,  being  unipared  scorpioid 
cymes,  which,  prior  to  opening,  are  tightly  rolled  up. — 130  species,  especially 
in  N.  Am.  (California,  etc.).  Many  annual  species  of  Phacelia,  Nemophila, 
Whitlavia,  Entoca,  Cosmanthus,  etc.,  are  cultivated  in  gardens  as  ornamental 
plants.  Hydrolea  (has  a  bilocular  ovary,  and  two  free  styles). 

Order  3.  Convolvulacese  (Bindweeds).  The  flower  is  regu- 
lar, hypogynous,  with  5  almost  free  sepals  (quincuncial  aestivation), 
P5,  A5,  G2  (rarely  3-5).  The  corolla  is  very  characteristic;  it 
is  (with  various  forms)  almost  entire,  or  slightly  5-lobed,  and 
folded  longitudinally  in  the  bud  in  such  a  way  that  5  projecting, 
flat  portions,  tapering  towards  the  top  and  frequently  differing  in 
colour  and  hairiness  from  the  rest,  are  visible  externally  and 
applied  close  together,  while  the  remainder  of  the  corolla  is  folded 
inwards  (Fig.  552  A)  ;  and  hence  the  whole  corolla  is  strongly 
twisted  to  the  right  in  the  bud.  The  gynoeceum  most  frequently  has 
a  bilocular  ovary;  in  each  loculus  there  are  only  2  (erect)  anatropous 
ovules  on  the  placenta,  which  is  not  especially  thickened  (Fig.  552 
D,  E) ;  each  loculus  is  sometimes  divided  into  two  by  a  false  septum 
(a  relationship  with  the  Boraginacece,  etc.);  style  simple  with 


516 


DICOTYLEDONES. 


most  frequently  a  bilobed  stigma,  or  a  bipartite  style.  The  fruit 
is  nearly  spherical,  most  frequently  a  capsule.  The  seeds  are  erect, 
and  have  a  large  hilum  at  the  base.  The  embryo  is  curved,  with 
leaf-like,  thin,  bilobed,  most  frequently  folded  cotyledons;  endo- 
sperm absent  or  mucilaginous. 

I.  CONVOLVULEJ;,  BINDWEED  GROUP.  The  majority  are  twining 
(to  the  left)  herbs,  with  latex.  The  leaves  are  scattered,  without 
stipules,  often  long- stalked,  and  nearly  always  with  cordate  base  ; 
some  are  palmately  lobed.  The  flowers  are  most  frequently  solitary 
in  the  leaf-axils,  large,  quickly  withering. — Convolvulus  (Fig.  552), 
Calystegia  (unilocular  ovary,  2  large  bracteoles),  Ipomcea,  Batatas, 
Evoknlus  (with  a  doubly  bifid  style),  Colony ction,  Pharbitis,  etc. 


FIG.  552. — Convolvulus  tcammonia. 

2.  DICHONDRE^E.     This  group  is  a  more  primitive  form,  not  twining,  and 
without  latex.     It  has  2  j'ree  carpels  with  basal  style  (as  in  Boraginacese)  and 
valvate  corolla. 

3.  CUSCCTE^;,  DODDER  GROUP  (Fig.  553).    Parasites, with  round, 
filamentous  stems,  bearing  only  scale-like  leaves  and  almost  desti- 
tute  of   chlorophyll    (they    are    reddish    or  yellowish)  ;  they  are 
parasitic  upon  other  plants,  around  which  they  twine,  first  with 
narrow,  compact  coils  from  which  haustoria  (Fig.  553  A)  are  de- 
veloped which  enter  the  host-plant,  and  then  with  wider  coils  by 
which  they  raise  themselves  to  other  portions  of  their  host  or  try 
to  reach  other  plants.     On  germination  a  very  temporary  primary 
root  is  developed,  which  bears  root-hairs  as  far  as  the  tip  (root- 
cap  is  wanting)  ;  it  only  serves  as  a  kind  of  reservoir  for  water,  and 
perishes  very  soon  after  the  seedling  has  fastened  on  to  a  host. 
The  embryo  is  filamentous  and  rolled  up  spirally  (Fig.  553  C),  and 


PERSONA'J  '&. 


517 


is  sometimes  destitute  of  cotyledons.  The  flowers  are  crowded 
into  capitulate  inflorescences,  complicated  by  accessory  shoots 
(Fig.  553  A)  ;  they  have  S5,  P5  (imbricate  aestivation),  A5  (and 
beneath  the  stamens  5  scales  on  the  corolla-tube),  G2.  Fruit  a 
capsule  opening  by  *lid.—Cuscuta  europcea,  G.  epilinum  (Flax-Dod- 
der), C.  epithymum  (Lesser-Dodder),  G.  trifolii  (Clover-Dodder), 
etc.,  are  parasitic  on  different  hosts,  or  parasitic  each  on  its  own 
particular  host. 


FIG.  553.— Cuscuta  trifolii,  parasitic  on  Red  Clover.  A  A  portion  of  the  stem  with  an 
inflorescence  and  haustoria  (mag.);  B  seed  (nat.  size);  0  seed  (mag.)  ;  D  embryo  (nat 
size). 

840  species ;  the  majority  in  the  Tropics,  especially  Am.  Many  are  ornamen- 
tal plants.  OFFICINAL  :  some  on  account  of  their  purgative  properties  :  the 
tuberous  roots  of  Ipomcca  purga  (Jalap,  from  Mexico)  and  the  dried  latex 
("  Scammony  ")  of  Convolvulus  scammonia  (from  the  East).  The  tuberous  roots 
of  Batatas  edulis  (Trop.  S.  Am.)  are  used  as  a  common  vegetable  (Sweet  Potato) 
in  the  Tropics. 

Family  30.     Personatae. 

The  type  of  the  flower  is:  S5,  P5,  A5  (of  which  one,  or  in 
gome  cases  several,  are  suppressed),  and  G2.  The  flowers  are 


518  DICOTYLEDONES. 

hypogynoiis,  ^  ,  perfect  with  gamopetalous  corolla,  but  most  fre- 
quently irregular  (medianly  zygomorphic,  except  Solanacece),  the 
corolla  being  bilabiate  (divided  into  a  posterior  part  of  two  lobes 
and  an  anterior  part  of  three  lobes),  and  the  stamens  4,  didynamous 
(the  posterior  being  suppressed).  The  ovary  has  2  loculi  (only  1 
in  Utriculariacece,  Gesneriacece,  Orobanche)  •  the  placenta  in  the 
first-named  orders  (1—7)  is  most  frequently  very  thick,  and  bears  a 
great  many  ovules  (Figs.  554,  555,  557,  562)  ;  the  number  of  ovules 
in  the  last  orders  (8-9)  is  considerably  reduced  (Fig.  570). 

Special  mention  may  be  made  of  the  apparently  4-merous 
flower  which  is  found,  e.g.  in  Veronica  and  Plantago  (Figs.  567, 
562(7,  570,  571),  and  which  arises  from  the  typical  5-merous 
flower  by  the  suppression  of  the  posterior  sepal  and  the  posterior 
stamen,  and  by  the  union  of  the  two  posterior  petals  into  one. — 
Terminal  flowers  very  seldom  occur  on  the  main  axis,  and  would 
not  harmonise  well  with  the  very  irregular  form  of  the  flower. 
When  they  do  occur,  they  are,  as  a  rule,  "  peloric,"  i.e.  regular  (in 
Linaria  vulgaris  two  kinds  of  peloric  flowers  occur, — one  with  5 
spurs,  and  one  without  spurs).  The  halves  of  the  anthers  are  often 
divided  as  far  as  the  base,  and  laterally  so  widely  separated  from 
each  other  as  to  assume  an  almost  straight  line  (Figs.  563,  564). 
There  is  generally  a  nectary  ("  disc  ")  round  the  base  of  the  ovary, 
often  5-lobed  (or  divided  into  free  glands). — A  common  vegetative 
characteristic  is  the  absence  of  stipules. 

The  9  orders  of  the  Personatae  are  :  1,  Solanacese  ;  2,  Nolanaceae ;  3,  Scrophu- 
lariaceae  ;  4,  Utriculariaceas ;  5,  Gesneriaceae  ;  6,  Bignoniaceae  ;  7,  Pedaliaceas ; 
S,  Acanthaceas ;  9,  Plantaginaceae. 

Order  1.  Solanacese.  The  flower  (Figs.  554,  555,  559)  is 
hypogynous,  regular  (zygomorphic  in  Hyoscyamiis},  $,  and  gamo- 
petalous,  with  S5,  P5  (most  frequently 
imbricate  or  valvate),  A5,  G2,  the  2  car- 
pels being  placed  obliquely  (Fig.  554)  ; 
the  bilocular  ovary  has  a  very  thick  axile 
placenta  (Figs.  554,  555  H,  557),  which 
extends  almost  as  far  as  the  wall  of  the 
ovary.  The  fruit  is  a  capsule  or  berry; 
the  seeds  are  more  or  less  reniform,  and 
the  embryo  is  curved  (rarely  straight),  in 

Fi0.654_Dia~gramofP*tuma.      'a   ^^   endosperm    (Figs.    555  F,   G-, 

561). — Both  arborescent  and  herbaceous 

forms  are  found  in  the  order;  leaves  scattered  without  stipules, 


PERSONATE. 


619 


but  with  variously  formed  laminae  (always  penuinerved).  A 
peculiar  leaf-arrangement  is  found  in  many  species,  viz.  the  leaves 
are  borne  in  pairs,  a  large  and  a  smaller  one  together ;  these  pairs 
stand  in  2  rows,  and  the  flowers  are  then  situated  between  the 
individual  leaves  in  each  pair,  apparently  not  in  a  leaf  axil.  The 
inflorescences  are  frequently  unipared  scorpioid  cymes  without 
floral-leaves. 


FIG.  655. — Atropa  belladnnna :  A  is  reduced. 

Zygomorphic  flowers  occur,  and  thus  form  a  transition  to  the  closely  allied 
Scrophulariacese ;  the  zygomorphy  sometimes  shows  itself  only  in  the  relative 
length  of  the  stamens,  sometimes  also  in  the  corolla  (Hyoscyamus). — Nicandra 
is  5-merous  throughout  all  the  whorls. — The  peculiar  relative  leaf -arrangement 
in  this  order  occurs  from  sympodial  branching  and  displacement.  The  most 
simple  is,  e.g.  Datura  (Fig.  556  A) ;  each  shoot-generation  in  the  floral  parts  of 
the  plant  has  only  2  foliage-leaves  if1  and  /2),  and  then  terminates  in  a  flower  ; 
the  axillary  buds  of  both  the  foliage-leaves  are  developed  and  form  a  dichasium, 
but  since  the  leaves  are  displaced  on  their  axillary-shoots  as  far,  or  almost  as 
far,  as  the  first  leaf  of  these  axillary-shoots,  the  flowers  are  borne  singly  on  the 
dichasial  branches,  and  all  the  branches  appear  to  be  without  subtending  leaves 
(Shoot  I  is  white,  II  shaded,  III  white,  etc.,  diagram  A).  Scopolia  and  others 
(Fig.  556.  B)  differ  in  that  the  lowest  and  smallest  (/*)  of  the  two  leaves  on 
each  shoot  is  barren,  and  is  therefore  not  displaced ;  but  the  upper  one  (the 
second  bracteole,  /2)  is  displaced  as  in  the  first  instance,  and  consequently  it 
assumes  a  position  near  the  first  leaf  (the  shaded  leaf/2  of  shoot  I  being  placed 
near  the  white  leaf  f1  of  shoot  II,  etc.,)  of  the  next  youngest  shoot-generation, 


520 


DICOTYLEDONES. 


and  hence  the  leaves  are  borne  in  pairs ;  the  flower  placed  between  the  two 
leaves  of  a  pair  is  therefore  the  terminal  flower  of  the  shoot  to  which  the 
smaller  of  the  two  leaves  belongs,  and  the  larger  leaf  is  the  subtending  leaf  for 
the  floral  shoot  itself. 


FIG.  656.— Diagrammatic  representation  of  the  branching  in  Solanacea?.    The  various 
shoot-generations  are  white  or  shaded. 

A.  FRUIT  A  CAPSULE.  Nicotiana  (Tobacco)  has  a  2-valved 
capsule  with,  septifragal  dehiscence ;  the  valves  separate  at  the 
apex ;  the  corolla  is  funnel-shaped,  tubular,  salver-shaped  or 
campanulate.  The  flowers  in  panicles. — Datura  (D.  stramonium, 
Thorn-apple)  has  a  (frequently  spiny)  capsule  (Fig.  558),  which  is 


FIG.  657. — Fruit  of  Hyoscyamus  niger  after 
removal  of  calyx. 


FIG.  558.— Fruit  of  Datura  stramonium. 


falsely  A-locular  (at  the  top,  bilocular)  and  opens  septif ragally  with 
4  valves.     The  lower  part  of  the  calyx  persists  as  a  thick  collar 


PERSONATE. 


521 


(see  Fig.  558).  The  corolla  is  funnel-shaped.  The  flowers  are 
solitary,  large. — Hyoscyamus  (II.  niger,  Henbane)  has  a  pyxidium 
(Fig.  557)  enclosed  in  the  campanulate,  completely  persistent, 
thick-walled  calyx.  The  flowers  are  slightly  zygomorphic,  and 
borne  in  unipared  scorpioid  cymes.  Scopolia  (pyxidium) ;  Falriana 
(Heather-like  shrub) ;  Petunia  (slightly  zygomorphic  flower ;  funnel-shaped 
corolla) ;  Nierembergia ;  Brunfelsia  (almost  a  drupe) ;  Franciscea ;  Browallia. — 
Among  those  with  capsular  fruits  are  found  the  most  anomalous 
forms,  which  by  their  zygomorphic  flowers  and  often  didynamous 
stamens  present  the  transition  to  the  Scrophulariaceae :  Salpig- 
lossis ;  Schizantlius  (lobed  petals ;  2  perfect,  and  3  rudimentary 
stamens). 

B.  FRUIT  A  BERRY.  Solanum  (Nightshade)  ;  rotate  corolla 
(Fig.  559).  The  stamens  have  short  filaments,  the  anthers 
stand  erect,  close  together  round  the  style,  like  a  cone  in  the 


559.  560.  CC1. 

Fies.  559-561.— Solarium  tuberosum. 

FIG.  559.— Flower  (i).  FIG.  560.— Stamen,  ejecting  pollen. 

FIG.  561. — Longitudinal  section  of  seed. 

centre  of  the  flower,  and  open  by  pores  at  the  apex  (Fig.  560). 
S.  tuberosum  (the  Potato-plant);  the  Potato-tuber  is  a  swollen,  underground 
stem;  the  "eyes"  are  buds,  situated  in  the  axils  of  its  scale-like,  quickly- 
perishing  le&ves.—Lycopersicum  resembles  Solanum  in  the  flower,  but 
the  united  anthers  open  by  longitudinal  clefts  and  have  an  apical 
appendage.  The  cultivated  species,  L.  esculentum  (Tomato),  has 
often  a  higher  number  than  5  in  the  flower,  and  in  the  fruit  several 
loculi  of  unequal  size. — Physalis  (Winter  Cherry)  ;  the  calyx  ulti- 
mately swells  out  in  the  form  of  a  bladder,  becomes  coloured,  and 
loosely  envelopes  the  spherical  berry. — Capsicum  (Guinea  Pepper- 
plant)  ;  some  species  have  very  large,  irregular,  rather  dry  (red, 
yellow,  black)  berries,  which  are  unilocular  in  the  upper  part.— 
Ltjcium  (false  Tea-plant) ;  the  corolla  is  salver-  or  funnel-shaped ; 
shrubs;  often  thorny. — Atmpa  (A.  belladonna,  Deadly  Nightshade, 


522  DICOTYLEDONES. 

Fig.  555)  ;  corolla  campanulate  ;  the  calyx  projects  beneath  the 
spherical,  black  berry.  The  flowers  are  'borne  singly. — Mandra- 
gora;  (Mandrake)  ;  Nicandra  (ovary  often  5-locular). — A  small  tropical 
group:  CESTRE^E  (Cestrum,  Habrothamnus,  etc.)  has  an  almost  straight  embryo, 
which  may  also  be  found  e.g.  in  species  of  Nicotiana.  Belated  to  the  Scro- 
phulariaceae. 

About  1.500  species  ;  the  majority  within  the  Tropics,  outside  these  limits 
especially  in  America.  Solanum  nigrum  is  a  common  weed. — The  Potato-plant 
(Solanum  tuberosum),  from  Peru  and  Chili,  was  introduced  into  Europe  in  1584 
by  Sir  Walter  Ealeigh.  (Potatoes  =  Batatos).  The  fruits  of  several  serve  as  con- 
diments :  Chilies  or  Pod-pepper  (Capsicum  annuuru  and  longum),  and  the  Cayenne- 
pepper  (C.  baccatum  and  others),  whose  fruits  also  are  officinal,  were  brought  to 
Europe  from  S.  America  by  Columbus,  and  are  commonly  cultivated  in  Tropical 
America;  Lycopersicum  esculentum  (Tomato)  and  others  from  Peru;  Solanum 
ovigerum  (Egg-plant) ;  Solanum  melongena,  etc.  Poisonous,  acrid,  narcotic  pro- 
perties (alkaloids,  etc.,  solanine,  nicotine,  atropine,  hyoscyamine)  are  found  in 
many:  Atropa  belladonna  (from  S.Europe;  the  roots  and  leaves  are  officinal); 
Solanum  dulcamara  (Bitter-sweet ;  formerly  officinal),  S.  toxicariiim  (Guiana) ; 
Datura  stramonium  from  Asia  (leaves  and  seeds  officinal),  D.  sanguinea,  metel, 
tatula,  and  others  ;  Hyoscyamus  (officinal :  the  leaves  and  seeds  of  H.  niger) ; 
Nicotiana  tabacinn  (Virginian  tobacco,  officinal :  the  leaves),  N.  rustica  and 
others  from  Trop.  America  (Tobacco  was  introduced  into  Europe  in  1560) ; 
Osfrww-species.  Duboisia  myoporoides  (Australia);  the  leaves  contain  hyoscya- 
mine and  are  used  in  medicine.  A  number  of  species  of  these  genera  are  orna- 
mental plants. 

Order  2.  Nolanacese.  These  most  resemble  the  Convolvulaceae  in  t  >e 
corolla,  but  the  Solanacese  in  their  branching,  and  leaf-arrangement  (in  pairs, 
etc.).  The  diagram  is  the  same  as  in  Nicandra  with  5  carpels,  but  the  fruits 
of  this  order  most  frequently  form,  by  invaginations  in  various  directions,  an 
ovary  (with  1  style)  consisting  of  numerous  and  irregularly  grouped,  1-ovuled 
cells;  the  fruit  is  a  schizocarp  with  many  1-seeded  fruitlets. — Nolana  (Wtstern 
S.  America)  :  a  few  are  ornamental  plants. 

Order  3.  Scrophulariacese.  The  flower  is  hypogynous,  $ , 
zyyom  orphic,  with  the  usual  type :  S5,  P5,  A5,  and  G2,  the  latter 
placed  in  the  median  plane  ;  some  genera  have  all  5  stamens  de- 
veloped (Fig.  562  A),  but  most  frequently  the  posterior  one  is  sup- 
pressed and  the  flower  becomes  didynamous  (Fig.  562  S}.  The  fruit, 
as  in  the  capsular-fruited  Solanacese,  is  abilocular,  2-valved  capsule, 
with  a  thick,  axile  placenta,  and  most  often  septicidal  dehiscence 
(Fig.  563  C}.  The  numerous  seeds  are  not  reniform  as  in  many 
Solanacece,  and  have  a  straight,  or  only  slightly  curved  embryo,  with 
abundant  endosperm  (Fig.  563  D). — The  majority  are  herbs;  some 
are  arborescent ;  the  leaves  are  opposite  or  scattered,  but  stipules 
are  wanting  as  in  the  whole  family. 

The  Scrophulariaceas  are  closely  allied  to  the  Solanaceae,  and  there  is,  properly 


PERSONATE. 


523 


Bpeaking,  no  characteristic  feature  which  absolutely    separates   them.      The 

somewhat  irregular  corolla,  with  five  stamens  of  unequal  length  in  Verbascum,  is 

*    also  found  in  Hyoscyamus  ;  curved  and    straight  embryos  are  found  in  both 

orders.    The  aestivation  of  the  corolla  in  the  Scrophulariaceae  is  simple  imbricate, 


FIG.  582.— Diagrams.    A  Verbascum ;  B  Linaria;  C  Veronica. 

in  the  Solanaceae  most  frequently  folded  imbricate  (in  Atropa  and  those  allied 
to  it,  imbricate  without  folding).  The  genera  (about  164)  are  distinguished 
according  to  the  form  of  the  corolla,  number  of  stamens,  inflorescence,  arrange- 
ment of  the  leaves,  etc.  Verbascum  belongs  to  the  most  primitive  5-stamened 
forms,  and  from  it  proceed  a  long  series  down  to  Veronica,  with  only  two  stamens 
and  most  frequently  the  posterior  sepal  suppressed. 

I.  ANTIRRHINEJ:,  SNAPDRAGON  GROUP.  This  has  most  frequently 
a  descending  aestivation  of 
the  petals  (the  posterior 
petals  are  outside  the  lateral 
ones,  which  again  enclose 
the  anterior;  Fig.  562  J., 
B}.  The  plants  belonging  to 
this  group  are  not  parasites. 

a.  5-stamened.  —  Ver- 
bascum (Mullein,  Fig.  563  J) 
has  a  slightly  irregular, 
rotate  corolla ;  five  stamens 
(frequently  covered  with 
woolly  hairs),  of  which  the 
two  anterior  ones  are  the 
longer  and  differ  often  also 
in  other  respects.  The  inflor- 
escences are  racemose,  often  with 
several  series  of  accessory  di- 
chasia  in  the  axil  of  each  primary 
floral-leaf.  The  leaves  are  scat- 
tered and,  together  with  the 
stems,  are  often  covered  with  a 
grey  felt  of  branched  hairs.  Pi&.  S63'.—  Verbascu 


524 


DICOTYLEDONES. 


b.  4-stamened,  didynamous  (Fig.  564).  —  Scrophularia 
(Fig-wort,  Fig.  565)  has  cjmose  inflorescences  in  a  panicle;  the 
corolla  (Fig.  565)  is  urceolate,  short  two-lipped ;  the  posterior 


FIG..  581. — Antirrhinum  majus.     A  flower,  and 
the  upper  lip  of  a  flower  with  the  stamens. 


FIG.  666.— Digitalis 
purpurea. 


stamens  are  present  as  a  scale  below  the  upper  lip  of  the 
corolla  (Fig  565  s).  S.  nodosa  has  a  tuberous  rhizome. — Pentstemon;  the 
posterior  stamen  is  barren  and  very  long. — Antirrhinum  (Snapdragon). 
The  corolla  (Fig.  564)  is  personate,  i.e.  bilabiate,  but  with  the 


FIG.  565.— Scrophularia  nodosa.  Protogynous  flower  in  various  stages :  A  ?  stage ;  g 
the  stigma  projecting  from  the  throat  of  the  corolla  ;  B  the  same  in  longitudinal  section; 
C  S  stage,  the  stigma  is  bent  down  and  its  former  position  occupied  by  the  stamens ;  s 
etaminode ;  g  stigma ;  d  nectary. 

under  lip  arched  to  such  an  extent  that  it  meets  the  upper  lip, 
closes  the   corolla  throat,  and  entirely  conceals  the  stamens  and 


PERSONATE.  525 

style  ;  the  corolla- tube  is  produced  into  a  short  pouch  at  the  base  on 
the  anterior  side.  The  capsule  is  oblique  and  opens  by  2-3  pores, 
formed  by  small,  dentate  valves.  In  Linaria  (Toad-flax)  the 
pouch  is  produced  into  a  spur.  Sometimes  there  are  traces  of 
the  posterior  stamens.  The  capsule  opens  by  large  pores  (one 
for  each  loculus),  produced  by  large,  many-partite  valves.  L. 
vulgaris  reproduces  by  suckers. — Liigitalis  (Foxglove,  Fig.  566) 
has  long  racemes  with  drooping  flowers  ;  the  posterior  sepal  is 
small  (a  step  towards  complete  suppression,  as  in  Veronica)  ;  the 
corolla  is  obliquely  campanulate,  and  generally  nearly  4-lobed,  the 
two  posterior  petals  coalescing. — Alonsoa;  Nemesia ;  Chelone ; 
Herpestis  ;  Mimulus ;  Torenia;  Vandellia ;  Limosella  (L.  aquatica, 
Mud- wort,  native);  Scoparia ;  Capraria;  Erinus  (found  on  the 
Roman  Camp  at  Chesters,  Northumberland,  and  supposed  to  have 
been  introduced  from  Spain  by  the  Roman  soldiers);  Celsia  (near 
Verbascum);  Maurandia  ;  LopJiospermum ;  Rhodochiton ;  Collinsia ; 
Nycterinia,  etc. 

c.  2-stamened. — Gratiola  (Water-hyssop).  5-partite  calyx. 
The  upper  lip  of  the  corolla  is  undivided  or  slightly  bifid;  the 
two  anterior  stamens  are  either  en- 
tirely absent  or  are  reduced  to  stami- 
nodes  (a  transition  to  Veronica). — 
Veronica  (Speedwell),  most  frequently 
4-partite  calyx  ;  4-lobed,  rotate,  zygo- 
morphic  corolla  with  2  perfect  stamens 
and  no  trace  of  the  others  (Figs.  567, 
562  c)  ;  capsule  with  loculicidal  de- 
hisfence.  Calceolaria ;  the  corolla  has 
two  slipper-like  lips. 

2.    RHINANTHBJE,  YELLOW  -  RATTLE 

GROUP.      Herbs,  all  of  which  (with  the  Fl&-  667.-riower  of  Veronica. 

exception  of  Lathrcea)  are  annual  parasites  with  green  foliage- 
leaves.  They  attach  themselves  by  haustoria  to  the  roots  of  other 
plants  and  draw  nourishment  from  them.  The  majority  turn 
black  when  dried.  Racemose  inflorescences.  In  many  the  calyx 
is  4-partite,  the  posterior  sepal  being  absent,  or  very  small.  The 
corolla  is  distinctly  bilabiate  (Fig.  568),  with  most  frequently  as- 
cending (Estivation ;  in  the  majority  it  does  not  become  detached  at 
the  base,  but  by  means  of  a  ring-like  cut  some  distance  up  the  tube  ; 
4  didynamous  stamens ;  pollen-grains  dry,  easily  falling  out ;  the 
anthers  are  often  furnished  at  the  base  with  bristles  or  hairs  (Fig. 


526  DICOTYLEDONES. 

568)  which  play  a  part  in  the  pollination,  the  probosces  of  the 
insects,  being  forcibly  pushed  against  them,  agitate  the  anthers  and 
shake  out  the  pollen-grains.  Capsule  with  loculicidal  dehiscence. 
— Euphrasia  (Eye-bright),  Melampyrura  (Cow-wheat),  Rhinanthus 
(Yellow-rattle),  Odontites  (Bartsia),  Pedicularis  (Louse- wort),  and 
Lathrcea  (Tooth- wort)  all  have  native  species.  The  last  named 
is  pale  yellow,  or  reddish  (without  chlorophyll) ;  it  is  a  parasite  on  the 
roots  of  the  Hazel,  Beech  and  other  shrubs,  having  an  serial  stem,  and  an 
underground,  perennial  rhizome,  covered  with  opposite,  scale-like,  more  or  less 
fleshy  leaves  with  a  number  of  internal  glandular,  labyrinthine  cavities.  The 
inflorescence  is  a  unilateral  raceme.  It  approaches  Gesneriacece  in  having  a 
unilocular  ovary  with  two  parietal  placentae. 


FIG.  568. — Euphrasia  officinalis.  Flower  of  the  large  and  the  small-flowered  forms; 
showing  the  anthers  and  stigmas. 

The  mechanical  contrivances  for  POLLINATION  are  so  numerous  that  no 
general  principle  can  be  laid  down.  Personate  flowers,  like  those  of  Antir- 
rhinum are  only  accessible  to  strong  insects,  such  as  humble-bees,  which  can 
force  themselves  between  the  two  lips,  and  so  become  dusted  with  pollen  on 
the  back.  In  Euphrasia  and  other  Ehinanthece  the  insects  become  covered  with 
smooth,  powdery  pollen  when  they  shake  the  anther-apparatus  in  touching  the 
hairs  and  bristles  mentioned  above.  Scrophularia  nodosa  is  protogynous  (Fig. 
565).  Digitalis  purpurea,  however,  is  protandrous.  Mimulus  luteus  and  some 
others  have  sensitive  stigmatic  lobes,  which  shut  up  on  being  touched.  The 
Fmmica-species  constitute  a  series,  from  large-flowered  down  to  small-flowered 
forms,  and  parallel  with  them  are  found  various  gradations  from  insect-  to  self- 
pollination.  In  some  (as  Euphrasia  officinalis,  Rhinanthus  crista  galli)  there 
are  two  kinds  of  flowers  :  large,  which  are  pollinated  by  insects,  and  small, 
which  arc  self -pollinated  (Fig.  5G8).  Lathrcea  squamaria  (Tooth- wort)  is  a  pro- 
togynous spring-flowering  plant,  largely  visited  by  humble-bees.  Others  have 
cleistogamic  flowers.  Nycterinia  capensis  opens  its  flowers  at  night. 


PERSONATE. 


527 


2,000  species;  chiefly  from  the  Temp.  OFFICINAL:  Digitalis  purpurea  (the 
leaves;  Europe),  a  poisonous  plant.  Verbascum  thapms  and  thapsiforme 
Veronica  officinalis  ('<  Herba  V."),  Gratiola  officinalis  ("  Herba  »)  have  medicinal 
uses.  The  whole  of  the  Scrophulariaceas  are  more  or  less  suspicious,  if  not 
actually  poisonous,  and  none  serve  as  food.  Many  are  OIINAMENTAL  PLANTS  : 
Mimulm  luteus  (N.  America),  Paulownia  imperial™  (the  only  species ;  in  Japan  ; 
a  tree),  Antirrhinum  vulgare  (S.  Eur.),  Linaria,  Pentttemon,  Veronica,  Calceolaria 
(Peru,  Chili,  etc.). 


FIG.  669. — Leaf  of  ETtricularia  vulgaris,  with  bladder.  Median  lonsifcudinal  section 
through  a  bladder  containing  a  Cyclops.  At  a  a  hair  of  the  upper-lip,  at  i  2  bristles  of  the 
under-lip  of  the  entrance  (a,  b) ;  in  the  latter  are  placed  4  bristles  h ;  k  stalk  of  the  bladder 
in  which  is  seen  a  vascular  bundle.  (After  CohD.) 

Order  4.  Utriculariaceae.  To  this  order  belong  only  per- 
ennial, insectivorous, -aquatic,  and  marsh-plants  (200  species)  with  a 
more  or  less  characteristic  appearance.  They  differ  from  the 
Scrophulariaceae,  especially  in  having  2  stamens  (the  anterior) 
and  a  unilocular  ovary,  with  free,  central  placenta  (like  that  of  the 

W.  B.  MM 


528  DICOTYLEDONES. 

PrimulacesD).     For  the  rest  the  flower  is  distinctly  bilabiate,  both 
in  the  calyx  and  corolla.     Two-valved  capsule  ;  no  endosperm. 

Pinguicula  (Butter-wort)  has  a  rosette  of  leaves  close  to  the 
ground ;  these  are  sticky,  covered  with  glandular  hairs,  and  roll 
round  any  small  insects  which  may  be  caught  upon  them  ;  flowers 
solitary,  terminal  on  a  long  scape ;  calyx,  5-partite ;  corolla  with 
spur.  The  embryo  germinates  with  1  cotyledon. —  Utricularia 
(Bladder- wort).  Our  native  species  are  floating,  without  roots, 
with  hair-like,  divided  leaves,  studded  with  peculiar  bladders  (in 
the  Tropics  there  are  terrestrial  species,  with  ordinary  foliage). 
The  bladders  (Fig.  569)  have  an  aperture,  closed  by  a  valve  open- 
ing inwards,  so  that  small  aquatic  animals  are  allowed  to  enter,  but 
are  not  able  to  escape ;  they  are  thus  entrapped  in  the  bladders, 
and  are  probably  used  as  food.  Calyx  bipartite  ;  corolla  personate 
with  spur. 

The  embryo  of  Utricularia  is  very  imperfect,  scarcely  more  than  a  spherical, 
cellular  mass,  with  a  few  slight  leaf-rudiments.  On  the  germination  of  U. 
vulgaris,  several  bristle-like  leaves  develop  into  a  compact  rosette ;  the  stem 
then  develops,  and  also  the  finely-divided,  bladder-bearing  leaves.  A  primary 
root  is  not  developed.  The  stems  branch  copiously  and  in  a  very  peculiar 
manner.  The  growing-point  of  the  stem  is  rolled  spirally. — The  stigmatic  lobes 
are  sensitive  and  close  on  being  touched ;  self-pollination  often  takes  place, 
however,  in  Pinguicula. 

Order  5.  Gesneriaceae.  The  flower  in  this  order  may  be  both  epigynous 
(Gesnerieai)  and  hypoyynous  (Cyrtandrece) ,  but  otherwise  is  nearly  the  same  as  in 
Scrophulariaceae,  only  that  the  ovary  is  unilocular,  with  2  parietal,  often  bifid,  pla- 
centa. Of  the  5  stamens  the  posterior  is  rudimentary,  or  (more  rarely)  entirely 
wanting,  and  the  others  are  didynamous  (Cyrtandreee  have  often  only  2  stamens) ; 
their  anthers  are  generally  glued  into  a  quadrangular  mass.  The  majority  are 
herbs  with  juicy  stems,  opposite,  verticillate  or  scattered  leaves  without  stipules, 
often,  like  the  stems,  thick  and  juicy,  soft-haired  or  glabrous.  The  corollas 
are  often  highly-coloured  (scarlet,  red-yellow,  etc.,  and  spotted  internally),  large 
and  magnificent,  so  that  many  species  are  ornamental  plants.  GESNEBIE.E 
(often  epigynous)  have  endosperm  ;  S.  Am. — CYETANDREJE,  hypogynous,  with- 
out endosperm  ;  Asia,  S.  Africa. — Streptocarpus,  neither  the  primary  root  nor 
primary  shoot  attains  development ;  one  of  the  cotyledons  dies,  while  the  other 
grows  and  becomes  a  very  large  foliage-leaf,  from  which  spring  adventitious 
roots  and  adventitious  inflorescences. 

500  species.  Gloxinia,  Achimenes,  Gesneria,  Alloplectus,  Tydcea,  Columnea, 
Ncegelia,  sEschynanthus,  and  others,  especially  in  the  forests  of  tropical 
America.  Some  are  epiphjtes  on  trees,  others  prefer  the  leaf -mould  of  the 
forest  and  crevices  of  cliffs.  Several  genera  have  peculiar,  catkin-like,  under- 
ground shoots,  with  scale-like  compact  leaves ;  others  have  tubers. 

Orobanche  (Broom-rape)  is  allied  to  this  order  as  a  parasitic  form.  It  is  a 
parasite  on  the  roots  of  other  plants,  not  like  Lathrcea  by  means  of  thin  root- 


PERSONATJ3.  529 

branches  with  haustoria,  but  growing  with  the  base  of  its  stem  in  close  contact 
with  its  host,  and  probably  even  often  protruding  a  kind  of  thallus  into  it,  in  a 
manner  similar  to  the  Loranthaceae.  Its  aerial  shoots  are  not  entirely  destitute 
of  chlorophyll,  but  are  not  green  ;  they  only  bear  scale-leaves  and  terminate  in 
a  raceme  or  spike-like  inflorescence. — Some  Oro&anc/te-species  are  detrimental 
to  various  cultivated  plants  (Hemp,  Lucerne,  Tobacco,  etc.).  The  flowers  are 
strongly  zygomorphic  ;  the  posterior  sepal  is  often  wanting,  and  the  anterior  are 
united  to  the  two  lateral  ones.  Ovary  unilocular,  as  in  Gesneraceae,  with  2  or 
4  parietal  placentae. — The  exceedingly  small  seeds  have  a  very  rudimentary 
embryo,  formed  of  an  ellipsoidal,  cellular  mass,  without  indication  of  cotyledons 
or  other  organs. — About  100  species  ;  especially  in  the  Mediterranean  region. 

Order  6.  Bignoniaceae.  500  species;  nearly  all  trees  and  shrubs,  and  to  a 
great  extent  lianes,  climbing  by  tendrils  (modified  leaves),  which  are  sometimes 
terminated  by  a  special  clasping  apparatus.  These  lianes  have,  as  a  rule,  an 
anomalous  stem  structure,  the  wood  being  either  divided  into  four  wedges  at  right 
angles  to  each  other,  separated  by  four  grooves  filled  with  secondary  wood- 
parenchyma,  or  a  greater  number  of  wedges  occur,  by  the  cambium  ceasing  to 
form  wood  in  several  places.  The  leaves  are  most  frequently  opposite  and  com- 
pound ;  the  flowers  in  the  main  are  similar  to  the  didynamous  Scrophulariacese, 
and  especially  resemble  those  of  Digitalis  purpurea  ;  they  are  bilabiate,  large, 
and  beautiful,  campanulate  or  trumpet-shaped,  many  of  the  prettiest  ornamental 
plants  in  the  Tropics  belonging  to  this  order.  The  fruit  is  most  frequently  a 
large,  woody,  2-valved,  siliqua-like,  septifragal  capsule,  whose  valves  separate 
from  the  flat  and  broad  partition-wall,  which  bears  the  large,  generally  winged 
:seeds  :  Tecoma ;  Bignonia. — In  gardens :  Catalpa  syringcefolia  (Trumpet-wood) ; 
Tecomd  radicana  (from  S.  Am.). — "  Palisander  "-wood  is  from  Jacaranda  (S 
Am.). — Eccremocarpus  (N.  Am.)  forms,  by  its  unilocular  capsule,  a  transition 
to  the  Gesneriaceae  (E.  scaber;  herbaceous). 

Crescentia  is  allied  to  this  order ;  C.  cujete  (Calabash)  is  its  best  known 
.•species.  The  fruit  (unilocular  with  2  parietal  placentae)  is  a  very  large, 
spherical  or  ellipsoidal  berry,  with  a  firm,  finally  woody  outer  layer.  After 
the  removal  of  the  juicy  interior,  these  are  commonly  used  as  drinking  vessels 
in  Tropical  America. 

Order  7.  Pedaliaceae.  Sesamum  (orientale  and  indicum) ;  very  important 
oil-plants,  which  from  olden  times  have  been  cultivated  in  tropical  Asia  and 
Africa  for  food  and  as  medicinal  plants,  and  are  now  cultivated  in  America 
also.  The  seeds  are  used  as  a  raw  material  in  the  manufacture  of  soap  in 

Europe. To  this  order  also  belong  Martynia  and  Craniolaria,  which  have  a 

long  horned  capsule  and  sensitive  stigmas. — 46  species. 

Order  8.  Acanthaceae.  1,500  species;  mostly  erect,  slender,  branched 
herbs  or  shrubs,  rarely  arborescent,  especially  in  S.  Am.  and  Ind.  The 
.branches  frequently  have  swollen  nodes ;  the  leaves  are  opposite,  penninerved, 
undivided,  more  or  less  lanceolate  or  elliptical,  and  generally  leave  a  distinct 
scar  when  they  fall  off.  Stipules  are  wanting.  The  flowers  are  solitary  or  in 
dichasia,  which  are  arranged  in  4-rowed  spikes  or  racemes,  each  flower  with 
its  subtending  bract,  which  may  be  brightly  coloured,  and  most  frequently  also 
with  two  bracteoles.  With  regard  to  the  corolla  (which  is  often  labiate,  in  any 
.case  irregular,  and  frequently  prettily  coloured),  the  2  or  4  didynamous  stamens 


530  DICOTYLEDONES. 

(of  whose  anthers  one  half  is  inserted  lower  than  the  other,  or  suppressed)  and  the 
gynoeceum,  the  Acanthaceas  are  true  Personatae,  approaching  most  nearly  to  the 
Scrophulariaceae :  they  differ  from  the  other  orders  especially  in  the  fruit, 
which  is  a  bilocular,  2-valved,  often  elastically  dehiscing  capsule,  which  never 
has  more  than  2  rows,  and  in  some  only  2  seeds  in  each  loculus,  the  seeds 
heing  often  compressed  and  borne  on  strony,  curved  or  hook-like  fnnicles  (reti- 
nacula)  which  persist  after  dehiscence.  Embryo  curved  without  endosperm; 
radicle  pointed  downwards. — Cleistogamic  flowers  are  found  in  several  species. 
Cystoliths  are  common. 

The  following  grow  wild  in  Europe  :  Acanthus  (spinosus  and  mollis,  whose 
pinnatitid  leaves  served  as  models  for  the  capitals  of  the  Corinthian  columns). 
The  posterior  sepal  is  the  largest  of  all  the  leaves  of  the  flower,  and  covers  the- 
other  parts  like  a  helmet ;  the  2  anterior  sepals  are  united,  and  the  two  lateral' 
ones  are  small  and  greenish  ;  the  corolla  has  no  upper-lip,  but  only  a  3-lobed 
under-lip.  The  anthers  are  bilocular ;  the  filaments  ultimately  become  very 
firm. — Justicia,  Eranthemum,  Goldfussia,  Thunbergia  (a  twiner), ^Ruellia,  Die- 
liptera,  etc. — Ornamental  plants  in  conservatories. 

Order  9.  Plantaginaceae  (Plantains).  The  flowers  (Figs. 
570,  571)  are  regular,  ^,  hypogynous,  with  a  4-part,ite,  persistent 
calyx,  a  gamopetalous,  scarious  corolla  with  4  projecting  lobes,. 
4  stamens,  incurved  in  the  bud,  later  on  projecting  considerably, 
about  equal  in  length,  and  a  bilocular  ovary  with  one  long,  fila- 
mentous, undivided,  feathery,  papillose  style  (see  Fig.  571).  The 
ovary  is  most  frequently  bilocular  with  1-few  ovules  in  each 
loculus.  An  hypogynous  disc  is  wanting.  The  fruit  is  a  pyxidium 
with  1-few  peltate  seeds  attached  in  each  loculus  (Littorella  is  in 
several  respects  an  exception).  All  species  are  herbs,  the  majority 
with  leaf-rosettes  near  the  ground,  and  the  flowers  in  spikes  or 
capitula. 

The  labiate-like  flowers  are  in  this  case  entirely  concealed  under  a  regular,, 
apparently  4-merous  exterior.  The  structure  of  the  flower,  however,  is  the  same- 
as  in  the  Scrophulariacece,  only  the  reduction,  which  is  found  in  Veronica  (com- 
pare Figs.  562  C,  567  with  570,  571),  is  also  present  in  this  instance  and  the- 
lobes  are  also  more  equally  developed  ;  the  posterior  petal  corresponds  to  the  bi- 
lobed  upper-lip  ;  the  posterior  stamen  and  the  posterior  sepal  also  are  entirely 
wanting.  In  the  development  of  the  flower  there  is  no  trace  of  posterior  sepal 
or  stamen,  and  the  posterior  petal  arises  from  one  primordium,  but  the  two 
anterior  sepals  arise  before  the  lateral  ones.  The  position  of  sepals  and  petals 
does  not  agree  with  that  of  a  true  4-merous  flower,  which  is  represented  in  Fig. 
361  E.  The  bracteoles  are  always  suppressed  in  Plantago. 

Plantago  (Plantain,  Rib-grass).  The  foliage-leaves  are  most 
frequently  scattered,  entire,  with  curved  veins,  arranged  in  a 
rosette  close  to  the  ground  on  an  unlimited  rhizome ;  the  spike- 
like  inflorescence  is  borne  on  a  long  scape ;.  in  some  (P.  psyllium)> 


NDCULIFER.E.  531 

the  leaves  are  opposite  on  a  stem  with  well-developed  internodes, 
and  the  inflorescences  are  borne  in  their  axils.  The  order  also 
presents  a  transition  from  insect-pollinated  to  wind-pollinated 
flowers.  The  flowers  are  protogynous,  wind-pollinated  in  /'.  major  and  P. 
lanceolata,  partly  also  in  the  other  species,  but  insect-pollination  also  occurs, 
and  P.  media  has  three  kinds  of  flowers,  some  of  which  are  adapted  for  wind- 
pollination  (Fig.  571),  others,  with  short  filaments,  for  insects.  Littorella 
lacustris  (Shore- weed)  is  the  most  reduced  of  the  Plantaginacese  :  an 
aquatic  plant  with  rosettes  of  round,  awl-like  leaves  and  diclinous 
(monoecious)  flowers.  In  the  axils  of  the  foliage-leaves  is  a  very  short 
3-flowered  spike,  formed  by  2  sessile  9  -flowers,. and  above  them  a  long-stalked 
cj  -flower ;  all  the  flowers  are  lateral,  the  terminal  one  being  absent,  as  in 
Flantago.  The  $  -flower  is  essentially  the  same  as  in  Plantago,  but  the  $  -flower 
has  a  scarious  corolla,  with  a  narrow,  3-4-dentate  mouth,  which  closes  tightly 
round  the  nut-like  fruit. 


570.  571. 

FIGS.  670,  571.— Plantago  media. 

FIG,  570. — Diagram  of  Plantago  media. 

FIG.  571. Two  different  forms  of  the  flower  (magnified):  1,  chiefly  adapted  for  pollina- 
tion by  wind;  2,  for  insect-pollination,  a  The  stigma;  b  the  calyx;  fc  the  corolla. 

The  genus  Plantago  constitutes  nearly  the  entire  order  (200  species).  Some 
are  widely  distributed  weeds  (e.g.  P.  major,  "  The  white  man's  footstep  ").  In 
P.  psyllium  (S.  Eur.)  the  integument  of  the  seeds  is  mucilaginous,  and  swells 
•considerably  in  water. 

Family  31.     Nuculiferse. 

The  flowers  are  Jiypogynous  and  zygomorphic  (in  Boraginacece 
•and  Cordiacece,  however,  they  are  regular,  except  Echium  and  An- 
chusa  arvensis}.  The  calyx  is  gamosepalous,  the  corolla  bilabiate 
(except  in  the  two  orders  mentioned),  mostly  after  f ,  i.e.  divided 
into  a  2-leaved  posterior  portion,  and  a  3-leaved  anterior  portion. 


532  DICOTYLEDONES. 

The  aestivation  of  the  corolla  is  nearly  always  descending. — In 
Boraginacece  and  Cordiacece  there  are  5  stamens  of  equal  length  ; 
in  the  other  orders  4  didynamons  ones,  or  only  2  fertile ;  the 
posterior  stamen  is  sometimes  developed  as  a  staminode,  sometimes 
fertile  (in  Stilbacece).  The  ovary  is  formed  of  2  median  carpels 
(except  some  Verlenacece) ,  with  (1-)  2  ovules  on  each  carpel ;  in  the 
majority  of  the  orders  it  is,  however,  divided  by  a  false  partition- 
wall  between  the  dorsal  and  ventral  sutures,  into  4  loculi,  each  of 
which  is  often  raised  independently,  causing  the  style  to  be 
situated  in  the  depression  between  the  four  lobes  ("  gynobasic  " 
style,  Figs.  572,  573,  575,  579).  The  fruit  in  these  orders  most 
frequently  becomes  a  4t-partite  scliizocarp  with  nut-like  fruitlets. 
The  other  orders  have  a  1  (-2)-locular  ovary. — The  leaves  are 
simple,  without  stipules. 

The  family  is  related  to  (and  proceeds  from)  the  TuUjJorce,  especially  Convol- 
vulacece,  which  has  an  almost  similar  construction  of  the  ovary.  It  is  doubtful 
whether  the  Cordiacece  and  Boraginacece  should  be  classed  with  the  others. 

The  orders  are  :  1,  Cordiaceae  ;  2,  Boraginaceaa ;  3,  Verbenaceae  ;  4,  Labiatae ; 
5,  Selagmaceae ;  6,  Globulariaceae ;  7,  Stilbaceaa. 

Order  1.  Cordiaceae  unites  Convolvulacese  and  Boraginaceae.  Tree-like 
plants  with  5-  (4-10)  merous  flowers,  doubly  bifid  style,  and  drupe  with  4  or 
less  loculi.  No  endosperm  ;  cotyledons  folded. — 185  species  ;  tropical. 

Order  2.  Boraginacese.  The  vegetative  parts  are  very 
characteristic :  herbs  with  cylindrical  stems  and  scattered,  undivided, 
nearly  always  sessile,  entire  leaves,  without  stipules,  and  generally, 
together  with  the  other  green  portions  of  the  plant,  covered  with 
stiff  hairs,  consequently  rough  and  often  even  stinging  (hence  the 
other  name  for  the  order  Asperifolice) .  The  inflorescences  are 
unipared  scorpioid  cymes  with  the  branches  coiled  spirally  ("  heli- 
coid,"  Fig.  573)  before  the  flowers  open.  The  flower  is  perfect, 
regular  (obliquely  zygomorphic  in  JEchium  and  Anchusa  arvensis), 
hypogynous,  gamopetalous :  S5,  P5  (often  with  ligular  out- 
growths), A5,  G2,  but  each  of  the  two  loculi  of  the  ovary  becomes 
divided  by  a  false  partition- wall  into  two,  each  of  which  contains 
one  pendulous  anatropous  ovule  with  the  micropyle  turned  up- 
wards ;  the  four  loculi  arch  upwards,  so  that  the  ovary  becomes 
4-lobed,  and  the  style  is  then,  as  in  the  Boragece,  placed  at  the 
base  ("gynobasic")  between  the  four  projections  (Figs.  572,  573). 
The  fnrt  is  a  4s-partite  scliizocarp  with  four  nut-like  fruitlets  (Fig. 
572y.-^Endosperm  is  wanting  (except  in  Heliotropium). ;  the  radicle 
is  turned  upwards. 


NUCULIFERJE.  533 

The  INFLORESCENCES  are  often  double  unipared  scorpioid  cymes ;  the  bud  of  the 
second  bracteole  is  developed,  that  of  the  first  suppressed ;  in  some  cases  both  the 
bracteoles  are  suppressed  (Myosotis,  Omphalodes,  etc.),  but  in  other  instances  all 
the  first  bracteoles  (a)  only  are  suppressed,  and  the  others  are  then  situated  in  two 
rows  towards  the  under  side  of  the  coiled  axis,  while  the  flowers  are  situated  on 
the  upper  side.  Displacement  of  the  branches  or  of  the  floral-leaves  sometimes 
takes  place.  The  flowers  are  often  red  at  first,  and  later  on  become  blue  or 
violet ;  they  hardly  ever  have  any  smell.  The  fruit  entirely  resembles  that  of 
the  Labiates,  but  the  radicle  of  the  latter  is  turned  downwards.  The  fruitlets 
present  small  differences  which  have  systematic  importance  ;  they  are  hollow 
or  flat  at  the  base,  attached  to  a  flat  or  columnar  receptacle,  etc. 

1.  HELIOTROPIE^.    This  group  deviates  from  the  characteristics 
mentioned  above  in  the  undivided  ovary  and  terminal  ("  apical ") 
style.    In  this,  as  well  as  in  the  fact  that  in  some  genera   (Tourne- 
fortia,  Ehretia,  etc.)  the  fruit  is   a  drupe,  it  connects  this  order 
with  the   Cordiaceas.     Heliotropium,  Tiaridium,  and  others  have 
schizocarps. 

2.  BORAGES,  BORAGE  GROUP.  Style  gynobasic ;  fruit  a  schizocarp. 
A.     The  throat  of  the  corolla  is  without  ligules,  or  with  very 

small  ones. — Pulmonaria  (Lung- wort)  ;  funnel-shaped  corolla;  a 
whorl  of  hairs  in  the  corolla -throat.  —  Echium  (  Viper 's-bugloss) 
has  zygomorphic  flowers,  the  plane  of  symmetry  almost  coinciding 
with  that  of  the  very  well-developed  inflorescence  (through  the 
fourth  sepal)  ;  the  corolla  is  obliquely  funnel-shaped,  the  style 
is  more  deeply  cleft  at  the  apex  than  in  the  others;  stamens 
2  longer,  2  shorter,  and  1  still  shorter. — Cerinthe  has  a  tubular 
corolla  with  five  small  teeth  and  two  bilocular  fruitlets.  The 
bracts  are  large  and  leafy,  and,  like  all  the  rest  of  the  plant, 
are  almost  glabrous. — A  few  Lithospermum-species  have  a  naked 
corolla-throat ;  others  have  small  hairy  ligules,  which  do  not  close 
the  corolla-throat.  The  fruitlets  are  as  hard  as  stone,  owing  to 
the  presence  of  carbonate  of  lime  and  silica. — Mertensia  (Steen- 
hammera')  ;  Amelia;  Nonnea  (small  ligules). 

B.  The  corolla-throat  is  closed  by,  or  in  any  case  provided 
with  ligules,  i.e.  scale-like  bodies  or  small  protuberances,  situated 
in  the  throat  of  the  corolla  opposite  the  petals,  and  which  are 
invaginations  or  internal  spurs  of  the  petals  (Fig.  572  D). — The 
nuts  in  Cynoglossum  (Hound's-tongue)  bear  hooked  bristles  over  the 
entire  surface,  or,  in  Echinospermum,  only  on  the  edge.  The 
following  have  smooth  nuts  -.—Symphytum  (Comfrey)  has  a  cylin- 
drical, campanulate  corolla,  and  prolonged-triangular,  pointed 
ligules.—  Borago  (Borage)  has  a  rotate  corolla  with  projecting, 


534 


D1COTYLEDONES. 


emarginate  ligules ;  the  stamens  have  a  horn-like  appendage,  pro- 
jecting upwards  from  the  back  of  the  filament.  The  fruitlets  are 
hollow  below.  —  Anchusa  (Alkanet,  Fig.  572).  The  corolla  is 
salver-shaped;  the  ligules  small,  hairy  protuberances.  A.  (Ly- 
copsis)  arvensis  has  an  S-curved  corolla-tube. — Myosotis  (Forget- 
me-not,  Fig.  573)  ;  rotate  corolla,  with  small  (yellow)  protuber- 
ances in  the  throat ;  scorpioid  cyme  without  floral-leaves  ;  fruitlets 

O 


FIG.  572.— Anchusa,  officinalis :  A  diagram;  the  bracteole  a  is  suppressed  (dotted);  0 
supports  a  flower.  B,  C  Myosotis,  the  fruit,  entire  and  with  the  calyx  in  longitudinal 
section.  I>,  F  Alkanva  tinctoria:  T>  the  corolla  opened  (y)  ;  e  the  ligule  ;  /,  g  the  anthers; 
E  gynosceum  (f);  F  fruit,  with  three  fruitlets;  i  an  aborted  locnlus  ;  li  disc. 

flat. — OmpJialudes  ;  fruitlets  hollow  at  the  back,  with  a  scarious, 
turned-in,  toothed  edge. — Asperugo  (Mad- wort);  the  calyx  grows 
after  flowering,  becoming  large,  compressed,  and  deeply  bifid. 

CROSS-POLLINATION  is  most  commonly  effected  by  insects  (especially  bees). 
There  are  a  great  many  contrivances  for  pollination;  some  flowers  are  protan- 
drous  (Echium  vulyare,  Borago  officin.),  others  are  heterostylous  (long-  and  short- 
styled:  Pulmonaria  officin.) ;  the  corona  (ligules)  is  a  protection  against  rain, 
and  excludes  certain  insects.  Some  are  barren  when  self-pollinated  (Pulmonaria 


NUCULIFERJE. 


535 


PIG.  573.— Myosotii.    Inflor- 
escence and  gynoeceum.    - 


officmalis,  Echium  vnJgare) ;  others  which  have  but  little  honey,  may,  failing 
insect-pollination,  fertilise  themselves,  and  in  Myosotis  versicolor  this  regularly 
occurs  by  the  growth  of  the  corolla  during  flower- 
ing, so  that  the  anthers  are  brought  into  contact 
with  the  stigma.  Honey  is  secreted  on  the 
hypogynous  disc.— About  1,150  species,  growing 
especially  in  the  northern  temperate  zone.  Mu- 
cilage is  found  (e.g.  in  the  officinal  root  of  Cyno- 
glossum  qfficinale,  in  the  root  of  Symphytum] ;  red 
dyes  are  found  in  some  roots  (e.g.  Alkanet-root,  the 
root  of  Alkanna  tinctoria,  which  is  also  medicinal ; 
S.  E.  Europe,  Asia  Minor);  some  are  poisonous: 
Cynoglossum,  Echium, Anchusa,  etc.  Several  species 
are  ornamental  plants.  Heliotropium  (Peru)  is 
cultivated  chiefly  on  account  of  its  pleasant  scent ; 
essential  oils  are  otherwise  very  rare. 

Order  3.  Verbenaceae.  The  majority  are 
shrubs  ;  a  few  are  herbs  or  trees  (Teak-tree) ;  some 
are  lianes.  The  branches  are  often  square.  The 
leaves  are  opposite  or  verticillate,  without  stipules; 
in  some  compound.  The  inflorescences  are  ra- 
cemes, spikes,  capitula,  or  dichasia.  Five  sepals;  five  petals  in  a  gamo- 
petalous,  zygomorphic  corolla,  which  is  often  bilabiate,  but  rarely  to  such  an 
extent  as  in  the  Labiatae,  and  the  upper  lip  in  some  is  larger  than  the  under, 
in  others  smaller;  stamens  four  didynamous,  or  two;  the  ovary  is  entire  (not 

.grooved  or  divided),  1-  or  2-locular,  or,  as  in  the  Labiatse,  divided  into  four 
loculi  with  an  erect  ovule  in  each,  but  in  some  the  antbrior  carpel  is  suppressed. 
One  terminal  style.  The  fruit  is,  e.g.  in  Verbena,  a  4-partite  schizocarp  with 
nut-like  fruitlets;  in  Vitex  (digitate  leaves)  a  drupe  with  a  4-locular  stone;  in 
Clerodendron  a  similar  fruit,  with  four  free  stones ;  in  Lantana  a  bilocular  stone, 
or  two  unilocular  stones.  The  radicle  is  turned  doivnwards.  Endosperm  small 
or  absent. — Lippia,  Stachytarpheta,  Bouchea,  Priva,  Githarexylon,  Callicarpa, 

•  etc. — The  Verbenaceae  are  closely  allied  to  the  Labiatae ;  they  differ  especially 
in  the  ovary  not  being  4-lobed  with  gynobasic  style,  but  undivided,  almost 
spherical  or  ovoid  with  a  terminal  style.  Again,  the  leaves  are  not  so  con- 
stantly opposite,  and  the  inflorescences  are  various. 

730  species;  especially  in  the  Tropics;  there  are  several  in  America,  especially 
Lantana-species  ;  shrubby  weeds. — Many  of  those  mentioned  are  ORNAMENTAL 
PLANTS  especially  Verbena;  Vitex  agnus  castus  is  a  S.  European  shrub.  Lippia 
citriodora  (S.  Am.)  etc.,  have  strongly-scented  leaves ;  the  Teak  tree  (Tectona 
grandis)  is  one  of  the  largest  trees  in  East  India,  and  has  a  very  hard  wood. 

Avicennia  is  allied  to  this  order;  it  inhabits  the  Mangrove  swamps  on  tropical 
coasts.  The  endosperm  emerges  from  the  ovule,  carrying  the  embryo  with  it ; 
the  embryo  ultimately  bursts  the  endosperm  and  lies  free  in  the  loculus  of  the 
fruit ;  this  is  then  filled  by  the  embryo  with  its  large,  green  cotyledons,  which 

,  are  borne  on  an  already  hairy  or  rooted  stem.  The  seedling  thus  developed 
falls  from  the  tree,  together  with  the  fruit,  and  strikes  root  in  the  mud.  One 
special  cell  of  the  endosperm  at  an  earlier  period  becomes  a  highly-developed 
organ  of  suction,  growing  into  a  much-branched  sac,  very  lich  in  protoplasm. 


536  DICOTYLEDONES. 

Order  4.  Labiatae.  The  special  characteristics  are:  the  square 
stem,  the  opposite  leaves  (without  stipules),  the  inflorescences  which 
are  formed  by  two  double  unipared  scorpioid  cymes,  the  labiate 
corolla,  the  4  didynamous  stamens  (the  posterior  being  entirely 
suppressed)  (Fig.  574),  and  the  4-partite  schizocarp  with  nut-like 
fruitlets.  The  floral  formula  is  S5,  P5,  A5  (the  posterior  stamen 
is  generally  absent),  G2. 

They  are  chiefly  aromatic  plants  (herbs,  shrubs,  e.g.  Lavender, 
or  trees),  volatile  oil  being  formed  in  internal  cells  or  in  the 
glandular  hairs,  which  cover  all  green  parts.  The  stem  is 
always  more  or  less  markedly  square  ;  the  leaves  are  borne 
upon  the  flat  sides,  and  are  simple  and  penninerved,  but  vary  in 
the  other  characters.  The  inflorescences  are  double  unipared 
scorpioid  cymes,  which  may  be  situated  at  some  distance  from 
one  another  in  the  axils  of  the  foliage-leaves  (Fig.  575  A),  but 
frequently  when  the  subtending  leaves 
are  bract-like,  they  are  crowded  into 
spike-like  inflorescences  (Lavandula, 
iMimv'i  \  Mentha,  Salvia,  etc.),  each  of  the  so- 
li \(  SB  J'JJf  called  "whorls"  (verticillaster,  glome- 
«-*^\vffc  U-l I  **-*•  rulus)  being  a  double  unipared  scorpioid 
oyme  (Fig.  574).  (Solitary  flowers  are 
found  in  e.g.  Scutellaria,  and  Origanum). 
FIO.  574.— Diagram  of  Lamium  The  calyx  is  strongly  gamosepalous,  5- 
toothed,  often  bilabiate  (Fig.  575  B). 

The  corolla  is  strongly  bilabiate  (Figs.  575,  576,  etc.),  with  2 
lobes  in  the  upper  lip  and  3  lobes  in  the  under  lip  (an  approach 
to  regularity  occurs  only  when  the  upper  lip  is  small,  and  thus 
resembles  one  lobe,  as  in  Mentha  (Fig.  578)  and  Lycopus,  so  that 
the  corolla  approaches  the  4-merous  corolla  of  Veronica  and  Plan- 
tago).  The  posterior  stamen  in  the  diagram  (Fig.  574*)  is  entirely 
suppressed ;  in  most  of  the  genera  the  posterior  lateral  stamens 
are  the  smaller  (Fig.  575  D),  and  are  entirely  suppressed  in  some 
(see  below)  ;  in  others,  e.g.  Nepeta,  they  are  the  longer.  2  stamens 
are  found  in  Salvia,  Rosmarinus,  Lycopus,  etc.  The  two  halves  of 
the  anthers  are  often  separated  from  one  another,  and  are  placed 
at  an  angle  with  each  other.  The  gynoeceum  has  1  style  with  a 
bifid  extremity  (Fig.  575  C)  bearing  the  stigma;  the  true  bilocular 
ovary  is  divided  by  a  false  partition-wall  into  4  loculi,  each  with 
1  erect  ovule  (Fig.  575  JET).  These  4  loculi  project  so  strongly 
that  the  ovary  becomes  deeply  4-lobed  with  the  style  situated  in 


NUCUL1FER.E. 


537 


the  centre  of  the  lobes  and  at  their  base,  "  gynobasic  "  (Figs.  575, 
579).  A  ring-like,  often  crenate,  nectary  surrounds  the  base  of 
the  ovary  (Fig.  575  G,  H).  The  embryo  in  this  order,  as  in  the 
Verbenacece,  is  directed  downwards  (Fig.  575  «7)  (it  is  directed  up- 
wards in  the  Boraginacece,  which  have  an  entirely  similar  fruit). 
Endosperm  absent. 


FIG.  575. — Thymus  vulgaris. 

The  142  genera  are  mainly  distinguished  according  to  the  form  of  the  calyx 
and  corolla,  the  number,  direction,  and  length  of  the  stamens,  the  forms  of 
the  nuts,  etc. 

I.  AJUGEJ:,  BUGLE  GROUP.  Calyx  10-nerved ;  the  upper  lip 
is  small ;  4  stamens.  The  ovary  is  not  so  strongly  lobed  as  in  the 
following  group,  so  that  it  is  most  nearly  allied  to  the  Verbenacece. 
The  nuts  are  reticulately  wrinkled.  Ajuga  (Bugle)  has  a  very 


538 


DICOTYLEDONES. 


PIG.  576. — Lamium  album:  A  lateral  view  of  flower;  B 
longitudinal  section ;  C  ovary  with  nectaries  (a) ;  D  the 
apex  of  the  style;  e,  upper  lip  of  corolla;  c,  b,  c  the  three 
petals  of  the  lower  lip ;  /  anthers ;  g  stigma. 


small  Tipper  lip.  The  upper  lip  of  Teucrium  (Germander)  is  deeply 
cleft,  and  the  two  lobes  are  bent  on  their  respective  sides  towards 
the  under  lip,  which  in  consequence  appears  to  be  5-lobed,  and 
the  upper  lip  to  be  wanting. 

2.     STACHYDEJ;,  BETONY  GROUP.     The  calyx  is  5-  or  10-nerved. 
The  upper  lip  of  the  corolla  is  most  frequently  strongly  arched  or 

helmet  -  shaped  ;  4 
stamens,  the  anterior 
pair  the  longer  (Fig. 
576). 

a.  A  somewhat 
regular  and  5-10- 
dentate  calyx  with 
projecting  stamens. 
— Stachys  (Betony, 
Woundwort)  ;  the 
C  '  lobes  of  the  under 

lip  are  rounded  off. 
The  anterior  fila- 
ments, after  pollina- 
tion, bend  outwards. 
Betonica. — Ballota  (Horehound)  ;  the  calyx  is  funnel-shaped,  and 
has  triangular,  long,  pointed,  awn-like  teetlci.^-Galeopsis  (Hemp- 
nettle)  has  two  conical  protuberances  on  the  under  lip  between 
the  lateral  and  the  central  lobes.  The  anthers  open  by  2  unequal 
valves.  Lamium  (Dead-nettle,  Fig.  576)  has  dentate,  lateral  lobes 

on  the  under  lip.  L.  album 
(White  Dead-nettle),  L.  ru- 
brum,  etc.  Galeobdolon. — Leo- 
nurus ;  Phlomis. 

b.  Tubular,  regular,  often 
10-toothed  calyx  and  con- 
cealed stamens. — Marrubium 
vulgare  (Fig.  577)  ;  10  calyx- 
teeth,  hooked  at  the  apex ; 
many  almost  spherical  whorls 
of  flowers  in  the  axils  of  the 
foliage-leaves,  at  some  dis- 
tance from  one  another. — 
Sid&ritis. 
c.  Strongly  bilabiate  calyx,  the  lips  doting  together  after 


FIG.  577 .—Marrubium  vulgare. 


NUCULIFER^]. 


539 


flowering.— Scutettaria  (Skull-cap);  the  two  lips  of  the  calyx  are 
entire,  the  upper  lip  has  a  large  spur,  and  drops  off  on  the  ripen- 
ing of  the  fruit.  The  flowers  are  generally  solitary  and  turned 
to  one  side.— Prunella,  (Heal-all)  ;  the  calyx  is  compressed,  its  two- 
lips  are  strongly  dentate,  the  upper  lips  closing  slightly  round  the- 
under.  The  stamens  have  a  tooth-like  projection  beneath  the- 
anthers. 

3.  NEPETE.E,  'CATMINT  GROUP.  13-15  nerves  in  the  calyx;, 
this  deviates  from  the  other  groups  in  the  posterior  stamens  being 
the  longer.  The  upper  lip  is  slightly  arched.  Nepeta  (Catmint), 
also  Glechoma  (Ground  Ivy),  with  regular,  and  Dracocephalum 
with  irregular  calyx. 


FIG.  578.—  Mentha  aquatica,  var.  crispa. 

4.  SATUREIEJ],  MINT  GROUP.  The  upper  lip  is  flat,  most  fre- 
quently ovate,  or  almost  spherical,  and  emarginate  (Fig.  578). 
The  calyx  is  most  frequently  5-10-nerved.  4  stamens,  the  anterior 
being  the  longer;  rarely,  2  stamens  only. —  Mentha  (Mint,  Fig. 
578)  has  a  regular,  5-dentate  calyx,  a  small,  almost  regular,  4- 
partite  corolla,  and  4  erect  stamens  of  nearly  equal  size.  The 
verticillasters  are  many-flowered,  and  are  often  collected  into 
cylindrical  inflorescences.  Herbs. — Lycopus  (Gipsy- wort);  corolla 
almost  regular.  2  stamens,  the  posterior  lateral  ones  are  wanting. 
Preslia  :  4-dentate  calyx,  4-partite,  regular  corolla;  4  stamens  of 
equal  size. —  Thymus  (Thyme,  Fig.  575)  has  a  strongly  bilabiate 
calyx,  the  throat  being  closed  by  a  whorl  of  hairs  (Fig.  575  B). 
The  corolla  is  distinctly  labiate.  Under-shrubs,  with  small  entire 
leaves;  verticillasters  few-flowered  and  separate. —  Origanum  (Mar- 
joram) ;  spike  or  capitate  inflorescences  with  the  flowers  solitary- 


540 


DICOTYLEDONES. 


in  the  axils  of  the  rather  large  and  distinctly  4-rowed  (often 
slightly  coloured)  floral-leaves.  Melissa.  Calamintha.  Clinopo- 
dium  (Wild  Basil).  Satureia.  Hyssopus  (Hyssop)  ;  small,  entire 
leaves;  the  verticillasters  are  situated  unilaterally  in  a  slender, 
spike-like  inflorescence.  Lavandula  (Lavender) ;  shrubs  with 
verticillasters  collected  in  cylindrical,  long-stalked  inflorescences ; 
the  calyx  is  tubular,  has  13-15  nerves,  the  posterior  tooth  is  much 
larger  than  the  others.  Stamens  and  style  do  not  project.  Coleus 
differs,  among  other  characters,  in  having  united  filaments  ;  the  stamens  and 
style  are  bent  down  and  concealed  in  the  boat-shaped  under  lip. 


FIG.  579.— Salvia  officinalis. 

5.  MONAKDE.E,  SALVIA  GROUP.  Only  the  2  anterior  stamens  are 
developed. — Salvia  (Fig.  579)  ;  calyx  deeply  bilabiate ;  the  upper 
lip  of  the  corolla  is  generally  strongly  compressed.  Rudiments  of 
the  two  lateral  stamens  are  present.  The  connective  in  the  two 
fertile  stamens  is  long  and  filamentous,  and  bears  at  the  upper 
end  a  normal  half-anther,  but  at  the  lower  one  a  barren,  often 
broader  portion,  against  which  the  insect  is  obliged  to  push  its 
proboscis  during  its  visits  to  the  flowers,  causing  the  pollen- 
bearing  half -anther  to  be  pressed  down  against  its  back.  Floral- 
leaves  often  coloured. — Rosmarinus  (Rosemary)  ;  a  shrub  with 
leathery  linear  leaves,  with  rolled  back  edge.  A  small  tooth  on 
the  filament  represents  the  barren  half  of  the  anther.  Monarda. 

The  POLLINATION  is  generally  effected  by  insects,  especially  bees  ;  the  under- 
lip  is  the  landing-stage  and  the  pollen  is  deposited  on  their  backs.  Cross-fertilisa- 
tion is  promoted  by  dichogamy  ;  honey  is  secreted  by  an  hypogynous  disc  and 
collected  in  the  corolla-tube.  Some  genera  are  homogamous  (Lamium,  Gale- 
opsis,  etc.) ;  others  are  dichogamous  (protandrous) ;  a  few  are  gynodixcious  : 


CONTORTS.  541 

$  -  and  £  -flowers  in  various  relative  sizes  (Glechoma  hederaceum,  Thymiu, 
Salvia  prafensis,  and  others).  The  entrance  of  uninvited  guests  to  the  honey 
is  often  rendered  difficult  by  whorls  of  hairs,  etc.  In  numerous  instances  the 
upper  lip  protects  the  pollen  from  rain.  Cleistogamy  is  found  e.g.  in  Lamium 
amplexicaule. 

2,700  species;  distributed  over  the  entire  globe,  but  the  greater  number  in 
Mediterranean  countries  (especially  in  the  Eastern  regions),  where  many  are 
shrub-like. — Poisonous  and  acrid  properties  are  absent.  On  account  of  their 
volatile  oils  they  are  principally  used  as  condiments,  for  perfumery  and  in 
medicine  (the  officinal  parts  are  therefore  nearly  always  "  folia  "  and  "  herba," 
in  Lavandula  the  flowers,  and  the  volatile  oils  extracted  from  them).  Such 
are  :  l  Mentha  piperita  f  (Peppermint) — menthol  is  obtained  from  this  species 
and  from  M.  arvensis — M.  viridis  f  (Spearmint),  M.  crispa  (Curly-mint),  Thymus 
vulgaris  (Garden  Thyme),  Melissa  officinalis  (S.  Eur.),  Hyssopus  officinalis 
(Hyssop,  S.  Eur.),  Origanum  majorana  (Marjoram,  from  the  Mediterranean), 
O.  vulgare  (Wild  Marjoram),  creticum,  smyrnceum,  etc.,  Salvia  officinalis 
(S.  Eur.),  Rosmarinus  offlcinalis  (oil  of  Bosemary,  S.  Eur.),  Lavandula  vera\ 
(oil  of  Lavender,  S.  Eur.).  Also:  Satureia  hortensis  (S.  Eur.),  Ocimum 
lasilicum  (E.  India),  Pogostemon  patchouli  (E.  India),  etc. — As  ornamental 
plants,  e.g.  Monarda,  Plectranthus,  and  Coleus  (foliage-plants,  often  with 
red  stems  and  leaves),  Stachys  lanata  (white,  woolly),  Phlomis,  Salvia-species, 
Perilla,  etc. 

Order  5.  Selaginaceae.  130  species ;  small,  most  frequently  heath-like 
shrubs  or  herbs,  mainly  from  S.  Africa.  They  differ  from  the  other  Nuculiferae 
especially  in  the  bilocular,  transversely-placed  anthers  of  the  4  stamens  (2 
stamens  divided  as  far  as  the  base  (?)).  The  ovary  has  2,  or  by  suppression 
only  1  loculus,  each  with  1  ovule,  and  the  fruit  is  a  schizocarp  dividing  into 
two,  or  is  a  1-seeded  nut.  Eadicle  turned  upwards. — A  few  are  ornamental 
plants  (Selago,  Hebenstreitia). 

Order  6.  Globulariaceae.  12  species ;  especially  in  the  Mediterranean.  They 
form  an  analogy  to  the  Composite,  and  in  the  main  resemble  Jasione  montana 
in  appearance,  the  flowers  being  crowded  into  a  spherical  head  (hence  their 
name)  and  supported  by  bracts,  but  without  involucre ;  the  ovary  is  unilocular 
with  1  pendulous  ovule.  The  l-seeded  nut  is  enveloped  by  the  persistent  calyx. 
The  corolla  is  more  or  less  labiate,  the  upper-lip  is  often  absent  as  in  the  ligu- 
late  corollas  of  the  Asterese  ;  stamens  4,  didynamous,  with  transversely  placed 
anthers  opening  by  one  transverse  cleft.  The  leaves  are  scattered,  simple, 
entire,  and  generally  form  a  rosette.  Globularia. 

Order7.  Stilbacese.  Heath-like  shrubs.  The  ovary  is  bilocular;  1  erect 
seed  in  each  loculus,  or  the  posterior  cell  is  empty.  Stilbe.  7  species.  S.  Africa. 

Family  32.    Contortse. 

Hypogynous,  regular,  g  >  gamopetalous  flowers  (Figs.  581,  582), 
which  are  generally  5-  or  4-merous,  with  5  or  4  stamens  (with 
the  exception  of  Oleacece  and  Jasminacece  which  have  only  2  sta- 
mens, alternating  with  the  carpels).  The  gynceceum  is  formed  of 

1  Those  which  are  officinal  are  indicated  by  t. 


542  DICOTYLEDONES. 

2  (nearly  always  median)  carpels.  The  corolla  very  frequently 
has  twisted  cestivation  (the  tipper  edges  of  the  petals  being  free  • 
Fig.  581  A),  and  hence  the  individual  lobes  of  the  corolla  are 
oblique,  but  the  flower  as  a  whole  is  regularly  actinomorphic. 
A  nectary,  in  the  form  of  a  honey- secreting  ri^g  or  glands,  is 
often  found  round  the  base  of  the  ovary. — The  leaves,  with  a  few 
exceptions,  are  opposite  and  without  stipules.  Endosperm  large 
(Fig.  581  (7),  except  in  Jasminacece  and  Asclepiadacece. 

The  ApocynaceaB  and  the  Asclepiadaceae,  on  account  of  the  free  ovaries, 
without  doubt  represent  a  more  primitive  form,  but  the  Asclepiadaceas  on  the 
other  hand  form  an  offshoot  on  account  of  their  peculiar  pollen-masses.  The 
Loganiacese  form  a  transition  to  the  Rubiaceae. 

The  orders  are  : — 

A.  STAMENS  5.    1,  Gentianacese  ;  2,  Apocynacese ;  3,  Asclepiadace®  ;  4,  Loga- 
niacese. 

B.  STAMENS  2.     5,  Oleaceae ;  6,  Jasminaceee  ;  7,  Salvadoraceae. 

Order  1.  Gentianaceae  (Gentians).  Glabrous  herbs,  with- 
out latex ;  the  opposite,  undivided  and  entire  leaves  are  often 
slightly  united  at  the  base ;  many  have  rosette-like  radical  leaves, 
Stipules  absent.  The  flowers  are  generally  borne  in  regular, 
dichotomously-branched  dichasia  (Figs.  580,  581^1),  which  finally 
become  transformed  into  uni  pared  scorpioid  cymes  ;  the  parts  of 

the  flower  are  4-5-merous  as  far  as 
the  gynoeeeum,  which  is  2-merous  ; 
the  calyx  frequently  is  almost  poly- 
sepalous  ;  the  corolla  has  distinctly 
twisted  aestivation  (the  upper  edges 
being  free)  (Fig.  581  ^4),  except 
Menyanthece.  The  carpels  are  en- 
tirely united,  and  most  frequently 
form  a  I-locular  ovary  with  2  pa- 
rietal placentae  bearing  many  ovules 
(often  in  several  rows,  Fig.  581  I), 
F).  Capsule,  2-valved,  with  septi- 
FIG.  580,-BrytJirao.  inflorescence,  cidal  dehiscence,  the  incurved  edges 
i,  2,  3,  etc.,  the  successive  shoot-  bearing  the  seeds  (Fig.  581  D,  F). 

I.  GENTIANEJ:. — Gentiana  (Gen- 
tian) has  most  frequently  a  tubular,  campanulate  or  funnel-shaped 
corolla,  sometimes  with  teeth  between  the  corolla-lobes  and  fringed 
in  the  throat  of  the  corolla;  G.  lutea  has  a  rotate,  yellow  corolla. — 
Swertia :  rotate  corolla ;  each  lobe  has  at  its  base  1-2  nectaries,  with  fringed  edges. 


CONTORTS. 


543 


Erythrcea  (Centaury,  Fig.  581) ;  corolla  most  frequently  salver- 
shaped.  The  anthers  ultimately  become  spirally  twisted  (E).  The 
style  prolonged,  deciduous.  The  flower  has  the  Lo&eZm-arrangement, 
i.e.  the  median  sepal  is  anterior;  the  corolla  is  rose-coloured  (in  the 
native  species).  The  capsule  is  semi-bilocular  (Fig.  581  F,  G). 
— Cicendia  has  a  low  creeping  stem,  fine  as  a  thread,  and  small,  yellow  flowers, 
4-merous  (without  twisted  anther). — Chlora  (Yellow-wort)  6-8-merous. 


FIG.  581.— Erythrasa  centaurium.  Inflorescence,  flower  and  fruit :  br1,  br2  floral-leaves  of 
the  1st  and  2nd  order  ;  Gf  a  valve  of  the  capsule  separated  from  its  fellow. 

2.  MENYANTHE^.  Menyanthes  (Buck-bean)  deviates  in  several 
respects  from  the  type  of  the  order.  The  leaves  are  scattered  and 
in  M.  trifoliata,  trifoliate ;  the  corolla  has  valvate  aestivation  ;  the 
testa  is  also  very  hard  (thin  in  the  true  Gentians).  They  are 
aquatic  plants  with  creeping  rhizome  ;  the  flowers  borne  in  racemes, 
with  terminal  flower,  heterostylous.  The  corolla  is  funnel-shaped 
with  a  very  hairy  throat.  —Limnanthemum  with  floating  leaves,  like  the 
Water-lilies. 

575  species;  distributed  over  the  entire  globe,  but  most  numerous  in  Alpine 
districts.  Neithei  poisonous  nor  nutritive  plants  are  found,  but  several  are 
used  in  medicine  on  account  of  the  bitter  properties  so  prevalent  amongst  them. 
OFFICINAL:  the  roots  of  Gentiana  lutea.  The  roots  of  other  species,  e.g.  G. 
purpurea,  punctata  and  pannonica  (Europe)  end  the  leaves  of  Menyanthes  tri- 
foliata  are  medicinal.  Some  are  grown  as  ornamental  plants  on  account  of 
the  pure  (often  deep  blue)  colour  of  the  flowers. 

Order  2.  Apocynaceae  (Periwinkles).  Trees  and  shrubs 
(also  lianes),  less  frequently  herbs,  generally  with  latex.  The 
leaves  are  opposite,  simple,  entire,  without  stipules ;  the  flowers  are 


W.  B. 


NN 


544  DICOTYLEDONES. 

regular;  corolla-lobes  oblique,  aestivation  twisted.  The  stamens 
are  individually  free,  and  the  pollen-grains  are  free  or  at  most 
united  in  fours  (see  Asclepiadaceae).  The  two  carpels  have  2-oo 
ovules,  in  all  cases  there  is  only  1  style  and  a  capitate  stigma, 
which  towards  the  base  is  widened  out  into  a  disc-like  table  (stigma- 
disc)  abstricted  in  the  centre  ;  but  the  carpels  in  most  of  the 
genera  (e.g.  those  mentioned  below)  are  entirely  separate,  and  the 
fruit  consists  of  two  follicles,  the  seeds  of  which  often  have  a  tuft 
of  woolly  hairs  projecting  from  the  micropyle,  less  frequently  of 
two  drupes.  In  some  other  genera  there  is  a  1-locular  (provided 
with  2  parietal  placentae)  or  a  2-locular  ovary  becoming  a  2- 
valved  capsule  or  a  berry.  Endosperm  abundant. 

Vinca  (Periwinkle)  has  a  salver-shaped  corolla,  which  is  twisted 
to  the  left  in  sestivation  (i.e.  the  left  edge  of  the  petals  is  free)  ; 
nectaries  2,  alternating  with  the  carpels;  the  summit  of  the  style 
is  hairy.  Follicles  ;  seeds  without  hairs.  Mostly  creeping,  perennial, 
evergreen  plants,  whose  large  flowers  are  apparently  axillary  ;  in  reality  they 
are  terminal,  but  by  the  development  of  the  bud  in  the  axil  of  one  of  the  two 
uppermost  leaves,  they  are  thus  displaced  over  the  other  leaf  of  the  pair 
{a  helicoid  sympodium  being  formed). — Plumeria,  Taberncemontana,  Cerbera 
(drupe).  Aspidosperma. 

Nerium  (Oleander).  The  leaves  are  in  whorls  of  3.  Corolla 
funnel-shaped,  in  aestivation  twisted  to  the  right,  and  with  a  corona 
resembling  that  of  Lychnis.  The  anthers  are  prolonged  at  the 
the  base  and  each  also  bears  at  the  apex  a  long,  linear,  hairy 
appendage ;  these  finally  become  spirally  twisted.  Follicles ; 
seeds  hairy.  Apocynum,  Echites,  etc.  Epigijnum  is  epigynous. 

124  genera,  1,000  species  ;  principally  in  the  Tropics.  Only  2  species  of 
Vinca  are  natives  of  this  country  ;  the  following  are  cultivated  as  ornamental 
plants : — Vinca  minor,  V.  major,  V.  (Lochnera)  rosea,  Amsonia  salicifolia, 
Nerium  oleander  (Eastern  Mediterranean).  The  latex  of  some  is  poisonous 
(Tanghinia  venemfera,  Cerbera).  Caoutchouc  is  obtained  from  others  (Han- 
kornia,  Landolphia,  Vahea,  etc.).  Tough  bast  is  frequently  developed.  The 
bark  of  Aspidosperma  quebracho  and  the  seeds  of  Strophanthus  hispidus  are 
used  in  medicine  (also  for  African  arrow-poison),  the  latter  is  officinal. 

Order  3.  Asclepiadaceae.  A  natural  and  easily  recognised 
order,  closely  allied  to  the  Apocynaceee,  having,  .like  it,  frequently 
a  poisonous  latex,  opposite,  single,  entire  leaves  and  fundamentally 
the  same  floral  diagram  and  floral  structure  (S5,  P5,  A5,  G2)  ; 
but  in  some  the  aestivation  of  the  corolla  is  valvate.  The  carpels 
here  also  have  free  ovaries,  but  are  united  for  some  distance  above 
into  a  large,  shield-like,  ^-angular  head,  having  on  its  under-side 


CONTORTS. 


548 


the  true  stigmas,  and  the  fruit  always  consists  of  2  follicles ;  seeds 

most  frequently  numerous  and  hairy  at  the  micropyle  ("vegetable 

silk");  endosperm  scanty. — The  order  is  distinguished  from  the 

Apocynacese  and  from  all  other  plants  also,  except  the  Orchids,  by 

having  all  the  pollen-grains  in  each  of  the  2  loculi  of  the  anthers 

(true  2-locular  anthers)  united  into  one  waxy,  club-shaped  pollen- 

?nass("  pollinium"),  for  the  purpose  of  pollination  by  insects.    These 

heavy  masses,  iri  order  to  secure  pollination  (as  in  the  case  of  the 

Orchids),  must  be  attached  to 

sticky  discs  (corpuscula) ;  there 

are  5  corpuscula,  one  at  each 

of  the  corners  of  the  5-angular 

stylar-head    (alternating  with 

the  anthers),  and    to  each  of 

these  are  attached  2  pollinia, 

one  from  each  of  the  anthers 

situated  on   either   side  (thus 

each    anther    gives    its    right 

pollinium  to  one  corpusculum 

and  its  left  to  another).  .  The 

stamens  are  frequently  united 

at  the  base,  and  each  bears  on 

the  back  a  variously  formed, 

petaloid     appendage,     termed 

a  "cucullus." 

A  peculiar  relative  position 
(and  therefore  a  good,  dis- 
tinctive characteristic)  is  often 
found  in  the  inflorescence,  which 
is  cymose  ;  it  is  placed  between 
the  two  leaves  of  a  whorl, 
nearer  to  one  than  to  the  other. 
The  leaf-pairs  are  placed  obliquely 
in  the  floral  region,  at  acute  and 
obtuse  angles,  and  not  at  right 
angles  (as  in  the  purely  vegetative 
parts) ;  the  inflorescences  are  placed 
in  two  rows  only  which  are  nearly 
90°  from  each  other,  and  the  two 
contiguous  to  one  another  are  anti- 
dromous;  they  are  in  reality  ter- 
minal, each  on  its  own  axis,  and  the 
entire  floral  portion  of  the  shoot  is 


FIG.  582.— Asclepias  cornuti.  A  An  open 
flower  with  the  calyx  (7c)  and  corolla  (c)  turned 
down ;  the  stamens  are  bent  together  and 
surround  the  gynceceurn.  B  The  androecium 
after  removal  of  the  sterile  part  (cucullus) 
of  the  anther,  which  functions  as  a  nec- 
tary: e  the  lateral  expansions  of  the  fertile 
portion  of  the  anthers;  /  the  slit  between 
the  expansions  of  two  contiguous  anthers, 
through  which  the  insect's  foot,  and  later  a 
pollinum  which  is  caught  by  it,  is  dragged, 
and  behind  which  the  only  receptive  part 
(stigma) is  hidden ;  above  the  slit  /  is  the  glnnd 
(r),  which  secretes  the  horny  corpusculum, 
which  is  split  at  its  base  and  joined  on  either 
side  with  a  pollinium  (this  is  more  distinctly 
seen  in  D  and  E).  When  the  foot  of  the  insect  is 
caught  in  the  slit  (/)  and  is  drawn  upwsmls, 
it  becomes  entrapped  in  the  slit  of  the  cor- 
pusculum, which  is  then  pulled  our,  together 
with  the  pollinia  firmly  attached  to  it.  In 
walking  over  the  flowers  the  insect  will  draw 
its  foot  through  other  slits  (/)  and  so  leave 
the  pollinia  on  the  stigmas.  C,  D  The  gjnce- 
ceuin  with  the  pollinia  hanging  freely, 
corpusculum  and  two  pollinia. 


E  A 


546  DICOTYLEDONES. 

a  unipared  scorpioid  cymose  sympodium  ;  in  addition,  complications  also  arise 
through  individual  parts  becoming  united. — Herbs  and  shrubs,  some  twining 
or  climbing. 

In  Asdepias  the  corolla  is  bent  back  and  there  is  a  cup-like 
cucullus,  from  the  base  of  which  protrudes  a  horn-shaped  body, 
bent  inwards. —  Vincetoxicum  has  a  rotate  corolla  and  a  ring-like, 
5-lobed  cucullus,  without  internal  prominences. — Stapelia  (especially 
from  S.  Africa)  is  remarkable  on  account  of  its  Cactus-like,  leafless  stems  and 
large,  brownish  flowers,  often  with  carrion-like  smell.  Periploca  has  more 
powdery  pollinia  (S.  Eur.,  etc.);  Hoy  a  carnosa  (Wax-flower;  Trop.  Asia)  is  a 
climber,  and  has  small,  annual,  flower-bearing  dwarf-branches.  Ceropegia. 

204  genera  with  1700  species,  distributed  over  all  tropical  countries ;  few 
outside  these  limits :  no  native  species.  Several  are  used  in  medicine  on 
account  of  the  pungent  properties  of  the  latex.  Condurango-bark  of  Gonolobus, 
condurango  is  medicinal.  Caoutchouc  is  obtained  from  the  latex  of  some  (e.g^ 
from  Cynanclmm).  The  seed-hairs,  which  are  most  frequently  shining,  silk- 
like,  and  white,  are  not  sufficiently  pliant  to  be  of  much  value.  Ornamental 
plants  in  our  gardens  :  Asclepias-species,  etc. 

Order  4.  Loganiaceae.  Ovary  single,  with  two  loculi,  in  structure  resem- 
bling the  Kubiacese,  but  superior.  360  species  are  included  in  this  order  ;  the 
majority  are  tree-like,  some  lianes  which  climb  by  tendril-like  branches.  The- 
inter{>etiolar  stipules  of  some  species  are  very  characteristic  (as  in  Rubiaceae,  to 
which  they  maybe  considered  to  be  closely  related).  The  fruit  is  a  capsule  or 
berry.  The  most  familiar  genus  is  Strychnos,  which  has  spherical  berries  with 
an  often  firm  external  layer,  and  compressed  seeds  with  shield-like  attachments; 
endosperm  abundant.  The  leaves  have  3-5  strong,  curved  nerves  proceeding 
from  the  base. — Spigelia. — They  have  no  lutex,  as  in  the  two  preceding  orders, 
but  many  are  very  poisonous  (containing  the  alkaloid  "  strychnine,"  etc.) ;  the 
South  American  arrow-poison,  urare  or  curare,  is  made  from  various  species 
of  Strychnos,  also  an  arrow-poison  in  the  East  Indian  Islands  (Java,  etc.) 
OFFICINAL,  the  seeds  of  Strychnos  nux  vomica  ("  Vomic  nut,"  Ind.).  The  seeds 
of  Strychnos  ignatii  (Ignatius-beans,  medicinal),  and  others  are  poisonous. 

Order  5.  Oleaceae.  The  leaves  are  always  opposite.  The 
inflorescences  are  racemes  or  panicles.  The  calyx  and  corolla  are 
4i-merotis,  more  or  less  united,  free  in  some  species  ;  the  corolla 
has  most  frequently  valvate  estivation.  All  four  forms  of  fruit 
occur  (see  the  genera).  Ovules  pendulous,  2  in  each  loculus 
(Fig.  583  0).  Endosperm  oily. — Syringa  (Lilac)  arid  Forsythia 
(anthers  somewhat  extrose)  have  capsules  with  loculicidal  de- 
hiscence  and  winged  seeds. — Fraxinus  (Ash)  has  winged  nuts 
(samara)  (Fig.  583  D)  ;  trees  with  most  frequently  imparipinnate 
leaves ;  the  flowers  are  naked  and  sometimes  unisexual  (poly- 
gamous), the  Manna  Ash  ( F.  ornus)  has  however  a  double  perianth 
with  4  free  petals  (Fig.  583  a)  ;  in  the  native  species,  F.  excelsior^. 


CONTORTS. 


547 


the  flowers  open  before  the  foliage  appears. — Ligustrum  (Privet)  has 
berries. — Olea  (0.  europcea ;  Olive)  has  drupes-,  the  pulp  and  seeds 
of  the  ellipsoidal  fruits  are  rich  in  oil.  The  lanceolate  leaves  are 
grey  on  the  under  surface,  being  covered  with  stellate  hairs.  In 
the  wild  state  it  is  thorny  (modified  branches). — Phillyrea; 
Chionanthus. — Few  species  of  Linociera  have  4  stamens. 


TIG.  633. — Fraxinus  ornus :  A  flower;  ca  calyx;  co  corolla;  B  gynoeceum  and  calyx; 
C  longitudinal  median  section  of  gynoeceum;  D  fruit. 

180  species  ;  chiefly  in  the  northern  temperate  zone.  The  Olive-tree  (')lea 
europcea)  has  been  an  important  cultivated  plant  from  ancient  times  (Olive  oil, 
Provence  oil,  "  Sweet  oil  ").  The  best  oil  is  extracted  from  the  fruit-pulp. 
The  fruits  are  edible.  Home:  Western  Asia,  Eastern  Mediterranean.  TIMBER : 
the  Ash  (Fr.  excelsior).  OFFICINAL  :  the  Manna  Ash  (Fr.  ornus),  cultivated  in 
the  Mediterranean  countries  for  the  sake  of  its  saccharine  juice,  which  flows 
out  and  coagulates  into  "  Manna." — The  following  are  ornamental  plants  : 
species  of  Ligustrum  and  Syringa  (introduced  in  the  16th  century,  from  S.E. 
Europe  and  Asia),  Forsythia  (China,  Japan  ;  the  large,  yellow  flowers  are  borne 
on  dwarf-branches  with  scale-like  leaves,  before  the  opening  of  the  foliage- 
leaves),  Chionanthus. 

Order  6,  Jasminaceae.  The  aestivation  of  the  corolla  is  imbricate  ;  the 
ovules  are  erect ;  seeds  almost  without  endosperm;  radicle  directed  downwards. 
The  number  of  lobes  in  the  calyx  and  corolla  is  not  4,  but  e.g.  5,  8,  10,  and 
variations  are  sometimes  found  in  the  same  individual.  The  fruit  is  a  berry  or 
capsule.  Many  species  are  twiners,  and  their  scattered  or  opposite  leaves  are 
most  frequently  imparipinnate.— 120  species ;  especially  in  Trop.  Asia  (K.  India). 
Some  Jasminum-syecies  are  cultivated  as  ornamental  shrubs  in  the  warmer  dis- 
tricts on  account  of  their  elegant  foliage,  and  beautiful,  sweet-scented  flowers, 
the  essential  oil  of  which  is  also  used  in  perfumery  ;  the  best  known  are :  J. 
sambac  and  grandiftvirtim.  Nyctanthes  arbor-tristis  opens  its  sweet-scented 
flowers  only  at  night  (E.  India). 

Order  7  (?).     Salvadoracese.     8-9  species;  Asia,  Africa. — Saluadora. 


548 


DICOTYLEDONES. 


b.     Tetracyclicse  with  epigynous  flowers. 
Family  33.     Rubiales. 

The  leaves  are  always  opposite  or  verticillate.     The  flower  is  epigy- 
nous, 2  j  o-  (or  4-)  mercras,  with  the  usual  sympetalous  diagram ; 


FIG.  581. — Cinchona  calisayo.    Flowering  branch. 

2-5  carpels.  The  inflorescences  are  frequently  dichasial.  The 
sepals  are  small,  reduced  to  teeth,  and  become  almost  entirely 
suppressed  in  the  higher  forms. — The  flower  is  regular  in  Rubiacece 


RUBIALES. 


549 


and  some  Caprifoliacece,  but  in  other  genera  of  this  latter  order 
(especially  of  Lonicerese)  it  is  unsymmetrical.  In  several  genera 
of  the  order  first  mentioned  the  loculi  of  the  ovary  contain  many 
ovules,  but  in  the  last  the  number  of  loculi  and  ovules  becomes 
reduced.  This  is  to  some  extent  connected  with  the  nature  of 
the  fruit  which  is  many-seeded  in  most  instances,  namely  a 
capsule  or  berry,  but  in  others  nut-like.  Endosperm  is  present. 

The  family  on  one  side  is  allied  to  the  Contortae  (not  only  through  the 
Loganiacece  but  also  through  the  Apocynacece),  and  may  be  regarded  as  an 
epigynous  continuation  of  this  family ;  on  the  other  side  it  is  allied  to  the 
Valerianaceaa  and  Dipsacaceae.  Many  points  of  agreement  with  the  Cornacece 
and  Araliacece  are  also  found,  and  in  fact  several  Caprifoliaceae  are  distin- 
guished from  these  by  hardly  any  other  feature  than  the  gamopetalous  corolla. 

Order  1.  Rubiaceae.  Leaves  opposite  (or  verticillate),  un- 
divided and  entire,  with  interpetiolar  stipules  (Fig.  586).  Flowers 


Fia.  585.— Cinchona  calisayu.    A  entire  flower;  B  after  removal  of  the  corolla; 
C  longitudinal  section  of  ovary;  D  fruit;  E  seed. 

epigynous  and  hermaphrodite,  regular,  4-  or  5-merous  with  the 
usual  arrangement  (Figs..  585,  588-590) ;  corolla  gamopetalous,  in 
aestivation  of  ten  valvate  ;  ovary  frequently  2-locular. 

There  are  no  external  characters  which  at  once  distinguish  this  exceedingly 
large  order,  as  in  many  other  natural  orders  (Composite,  Umbelliferae,  etc.). 
but  the  opposite  leaves  with  interpetiolar  stipules  form  an  excellent  mark  of 
recognition.  It  is  divided  into  many  sub-orders  and  groups,  especially  charac- 
terised by  the  nature  of  the  ovary  (1  or  several  ovules  in  each  loculus),  and  of 
the  fruit  (schizocarp,  berry,  drupe,  capsule). — The  corolla  is  bilabiate  in  4 
genera  ;  its  aestivation  in  some  is  twisted  ;  in  Capirona,  etc.,  the  filaments  are 
of  unequal  size.  The  ovary  is  semi -epigynous  in  Henriquezia,  etc.  In  Morindu. 
all  the  fleshy  fruits  coalesce  into  one  multiple  fruit. 


^ 


550  DICOTYLEDONES. 

1.  CINCHONEJ;.      The    fruit  is   a    2-valved  capsule,  with  many 
winged  seeds  (Fig.  585).     Cinchona  (Quinine,  Fig.  584).      Trees 
and  shrubs  with  the  foliage  and  inflorescence  somewhat  resembling 
Syringa  ;  the  corolla  also  being  of  a  lilac  colour,  more  or  less  salver- 
or  funnel-shaped,  and  frequently  edged  with  a  fringe  of  hairs  (Fig. 
585),  is  somewhat  similar  to  that  of  Menyanthes.    Their  home  is  the 
Andes  from  Bolivia  to  Venezuela,  varying  in  altitude  from  1-3000 
metres.     There  are  now  large  plantations  in  Java  and  E.  India. 
(The  name  "quinine"  is  of  Indian  origin;   that  of  the  genus  "  Cinchona,"  is 
from  the  Spanish  Duchess  Ciuchon,  who  in  1638  first  introduced  the  bark  into 
Europe.)     The  following  are  closely  allied  :  Cascaiilla,  Remijia,  Ladenleiyia, 
Manettia,  Bouvardia,  etc. 

2.  GAKDENIE^;.     Trees  and  shrubs,  frequently  having  a  many-locular  berry. 
Randia,  Gardenia,  Genipa,  Hamelia,  etc. 

3.  COFFEES.     Only  1  seed  in  each  of  the  two  loculi  of  the  ovary; 
the  fruit  is  a  drupe  with  2  stones.      Coffea  has  an  ellipsoidal  fruit 

about  the  size  and  colour 
of  a  cherry;  the  two  thin- 
shelled,  parchment-like 
stones  are  enclosed  by  a 
thin  layer  of  pulp  ;  the  two 
seeds  are  flat  on  the  side 
turned  to  one  another, 
which  has  also  a  deep, 
longitudinal  groove  curving 
to  the  sides.  The  endo- 
sperm is  hard,  horny  and 
greyish  (without  starch) ; 

FIG.  686.— Cephaelis  ipecacuanha.    Por*;or.  of  a          the     Small     embryo     lies     in 
branch  :8t  stipules.  the      lower     end      near     the 

circumference.  The  Coffee-plant  (C.  arabica)  is  a  small  tree,  or 
more  frequently,  and  especially  in  plantations,  a  shrub  with  large 
dark-green  leaves  and  scented,  white  flowers.  Its  home  is  in 
Tropical  Africa;  it  is  now  cultivated  in  many  tropical  countries. 
C.  liberica,  "W.  Africa. —  Cephaelis  ((7.  ipecacuanha,  Fig.  586;  the 
roots  are  officinal). — Psychotria,  Chiococca,  Ixora,  Hydnophytum,  Myr- 
mecodia,  etc. 

4.  SPEBMACOCE;E.      Chiefly    small  shrubs    and  herbs,  many  of   which  are 
weeds  in  tropical  countries.    The  stipular  sheaths  bear  numerous  bristles  at  the 
edge.     Spermacoce,  Borreria,  Diodia,  Richardsonia,  etc. 

5.     STELLATE.     Herbaceous  plants  with  verticillate  leaves  (Figs. 


RUB1ALES. 


551 


587,  588-590)  ;  the  stipules  are  large,  leaf-like,  and  resemble  the 
lamina  of  the  leaves,  so  that  the  leaves  appear  to  be  placed  several  in 
a  whorl,  while  in  reality  there  are  only  two  opposite  leaves,  the 
stipules  of  which  project  freely,  and  are  not  erect  (Fig.  587). 

In  some  cases  there  are  apparently  4  leaves  in  the  whorl,  and  then  2  of 
these  are  leaves,  and  the  other  two  are    their  interpetiolar  stipules.      When 


Fia.  687. — Rubia  tmctorum. 

there  are  apparently  6  leaves,  then  the  two  of  these  which  are  opposite  each 
other  are  leaves,  and  the  other  four  are  stipules ;  if  there  are  several  members 
in  the  whorl,  then  a  division  of  the  stipules  has  taken  place.  The  proof  of  this 
theory  is  founded  upon  the  fact  that  not  more  than  2  of  the  leaves  of  the  whorl 
ever  support  buds  (which,  in  addition,  are  seldom  of  equal  vigour),  and  also 
that  the  whorls  do  not  alternate  with  each  other,  which,  according  to  the  rules 
of  the  position  of  the  leaves,  they  should  do  if  all  the  members  of  a  whorl  had 


552 


DICOTYLEDONES. 


equal  value.  If  there  are,  for  instance,  4  members  in  two  succes>ive  whorls, 
they  stand  right  above  one  another,  and  do  not  alternate.  The  development 
and  anatomical  relations  (the  branching  of  the  vascular  bundles)  also  point  to 
the  same  conclusion. — All  the  other  groups  of  the  order  have  only 
2  small  scale-like  interpetiolar  stipules,  or  they  form,  at  the  base  of 
the  leaf-stalks  an  interpetiolar  sheath,  having  often  a  toothed  edge 
(Fig.  586). — Another  characteristic  feature  in  this  group  is  that 
the  calyx  is  rudimentary,  the  corolla  valvate  (Fig.  588),  and  that 
each  of  the  two  loculi  of  the  ovary  has  only  1  ovule.  The  fruit  is  a 
schizocarp  dividing  into  2  fruitlets  (Fig.  590).  The  forms  of  the  fruit, 
as  well  as  many  other  characters,  as,  for  example,  the  epigynous  flower,  the 
rudimentary  calyx,  the  two  free  or  almost  free  styles,  present  interesting 
analogous  resemblances  to  the  polypetalous  order  of  the  Umbelliferffl.  This 
group  has  its  home  chiefly  in  the  temperate  regions  of  the  northern 
hemisphere,  especially  about  the  Mediterranean  ;  it  is  the  only 
group  which  occurs  in  this  country,  represented  by  4  genera. 


658. 


5:0. 


589. 

FIGS.  558-590.— Rubia  tinctorum. 

FIG.  588.— Diagram.  FIG.  589.— Longitudinal  section  of  flower. 

FIG.  590. — Longitudinal  section  of  fruit  (|). 

Galium  (Cleavers)  is  almost  destitute  of  a  calyx  ;  it  has  a  small 
^-partite,  rotate  corolla,  4  stamens,  and  2  free  styles.  The  fruitlets 
are  nut-like.  The  inflorescence  is  a  paniculate  dichasium  passing 
into  helicoid  cymes. — Asperula  (Woodruff)  is  distinguished  from 
the  above  by  its  salver-  or  funnel-shaped  corolla.  1  style. — Rubia 
(Madder,  Figs.  587-590)  has  almost  the  same  form  of  corolla  as 
Galium,  bub  (most  frequently)  a  b-merous  flower,  and  the  fruitlets 
are  "drupes."  Sherardia  (Field  Madder) ;  the  flowers  are  clustered  in  closely 
arranged  cymes  surrounded  by  an  involucre ;  the  calyx  has  6  distinct  teeth, 
while  the  number  of  petals  and  stamens  is  4.  The  corolla  is  funnel-shaped. 
— Vaillantia.  Crucianella. 

The  DISTBIBUTION  OF  SEEDS,  in  some  instances,  is  promoted  by  hooked  append- 
ages on  the  fruitlets  (e.g.  Galium  aparinc). 

The  small  flowers  of  the  Stellatse  are  frequently  collected  in  compact  inflor- 
escences, and  are  therefore  rendered  more  conspicuous;  slight  protandry  is 


RUBIALES.  553 

found  in  some,  self-pollination  in  the  species  which  are  less  conspicuous. 
Many  species  are  heterostylous.  Myrmecodia,  Hydnophytum,  and  other  genera 
have  large  tubers  (hypocotyledonous  stems),  whose  labyrinthine  cavities  and 
passages  are  inhabited  by  ants. 

About  4,500  species;  tropical  or  sub-tropical  except  tbe  Stellatae ;  especially 
American.  Tbe  tropical  ones  are  mostly  trees. —  Several  are  OFFICINAL  on  ac- 
count of  the  large  amount  of  alkaloids  and  glycosides  which  they  contain.  The 
most  important  are  the  Cinchonas  (Cinchona  calisaya,  C.  succirubra,  C.  officinalis, 
C.  micrantha,  etc  ),  whose  bark  contains  the  well-known  febrifuge  and  tonic, 
Quinine,  Cinchonin,  etc. ;  Quinine  is  also  found  in  Exostemma,  Ladenbergia, 
and  Remijia.  The  root  '•  Ipecacuanha  "  (an  emetic)  from  Cephaelis  ipecacuanha 
(Brazils).  Caffeine  is  officinal.  The  use  of  the  seeds  of  the  coffee  plant  ("  the 
beans  ")  was  first  known  in  Europe  in  1583. — There  are  only  a  few  which  con- 
tain aromatic  properties,  principally  among  the  Stellataa  (coumarin  in  Asperula 
odorata,  the  Woodruff),  in  which  group  colouring  materials  are  also  found.  The 
root  and  root-stalks  of  Eubia  tinctorum,  the  Madder  (S.  Eur.,  Orient.,  Fig. 
587),  were  formerly  largely  used  for  dyeing,  but  are  now  superseded  by  the 
analine  colours.  Ked  dyes  are  also  obtained  from  the  roots  of  species  of 
Asperula  and  Galium.  Gambier  is  a  splendid  colouring  material,  obtained  from 
Uncaria  gambir  (S.E.  Asia),  which  is  used  in  dyeing  and  tanning. — The  order 
does  not  furnish  many  ornamental  flowers. 

Order  2.  Caprifoliaceae.  This  order  agrees  with  the  Rubiaceae 
in  having  opposite  leaves  and  an  epigynous  flower,  most  frequently 
5-merous  •  with  the  ordinary  tetracyclic  diagram,  but  in  some 
species  it  is  zygomorphic ;  the  corolla  has  imbricate  aestivation. 
carpels  3-5,  most  frequently  3  (not  2,  which  is  the  most  usual 
number  in  the  Rubiaceae).  The  fruit  is  generally  a  berry  or  a 
drupe,  but  the  most  important,  and  in  any  case  most  easily  recog- 
nisable feature,  is  the  absence  of  stipules;  in  exceptional  cases, 
where  they  are  present,  they  are  not  interpetiolar,  and  are  most 
frequently  small. — The  majority  of  plants  belonging  to  this  order  are  shrubs 
or  trees.  Compound  leaves  sometimes  occur.  Stipules  only  appear  in  a  few 
species  of  Lonicera,  Sambucus  and  Viburnum;  in  the  common  Elder  (Sambucus 
nigra]  they  are  in  some  instances  glandular  and  small,  but  in  other  cases  larger 
and  more  leaf-like  (upon  long,  well-developed  shoots) ;  in  the  Dwarf  Elder  (S. 
ebulus)  they  have  the  normal  leaf -like  form  ;  in  Viburnum  opnlus  they  are  pre- 
sent as  narrow  lobes  at  the  base  of  the  petiole;  in  others  tbey  are  completely 
absent.  The  leaves  are  frequently  penninerved,  rarely  palmiuerved.  The 
calyx,  as  in  the  Stellatae  and  Aggregatae,  is  often  very  insignificant. 

I.  LONICERE^:,  HONEYSUCKLE  GROUP.  This  has  campanulate  or 
tubular  corollas  which  are  often  zygomorphic ;  in  connection  with 
the  length' of  the  corolla  the  style  is  long,  filamentous,  and  most  fre- 
quently has  a  large,  capitate  stigma.  There  are  several  ovules  in 
the  loculi  of  the  ovary,  and  the  fruit  is  most  frequently  a  berry. 


554  DICOTYLEDONES. 

Lonicera  (Honeysuckle).  Shrubs,  sometimes  twiners.  The 
corolla  in  some  species  is  considerably  bilabiate  (Fig.  591),  with  4 
lobes  in  the  upper  lip,  and  1  in  the  under  lip,  but  in  others  more 
regular,  tubular,  or  campanulate.  The  flowers  are  either  borne  in 
capitate  inflorescences,  which  are  compound  and  formed  of  closely 
compressed  3-flowered  dichasia  (sect.  Oapnfolium),  or  in  dichasia 
with  2  flowers  (the  terminal  flower  is  wanting).  The  ovaries  and 


FIG.  591. — Lonicera. 

fruits  coalesce  in  some  (sect..Xylosteum).  The  opposite  leaves  in  some 
spec:es  unite  with  each  other  and  form  a  broad  collar  encircling  the  stem  (Fig. 
591).  Above  the  primary  bud  1-2  accessory  buds  are  often  found  in  the 
leaf-axils. — Diervilla  (IVeigelia) ;  with  a  2-locular,  2-valved  capsule. — Symphori- 
carpus  (Snowberry)  has  an  almost  regular,  funnel-shaped  corolla ;  a  peculiar 
feature  is  found  in  the  ovary  which  has  4  loculi,  the  2  median  having  many 
ovules  in  2  rows,  all  of  which  are  aborted ;  the  2  lateral  ones,  on  the  other 
hand,  each  have  only  1  ovule  which  is  developed.  Different  forms  of  leaves 
are  frequently  found  on  the  same  branch  ;  they  are  entire  or  lobed. 


RUBIALES. 


555 


2.  SAMBUCE x,  ELDER  GROUP  (Fig.  592) .  This  has  a  rotate,  regular 
corolla,  extrorse  anthers,  a  very  short  and  thick  (or  almost  absent) 
style,  with  tripartite  stigmas,  and  only  1  pendulous  ovule  in  each 
of  the  3  (-5)  loculi  of  the  ovary.  The  fruit  is  a  "  drupe  "  with  1-3 
(-5)  stones.  The  inflorescence  is  made  up  of  cymes  grouped  in  an 
umbel-like  arrangement. 

Sambucus  (Elder,  Fig.  592)  has  imparipinnate  leaves  and  a 
"drupe  "  with  3  (-5)  stones.  Between  the  calyx  and  the  style  a  disc 
remains  on  the  apex  of  the  fruit.  S.  nigra  with  black  fruit ;  S.  racemosa 
with  red  fruit;  S.  ebulus  is  a  perennial  herb;  the  others  are  woody. —  Vibur- 
num (Gruelder-rose)  has  simple  leaves  (penninerved  or  palmi- 
nerved,  entire,  dentate  or  lobed),  and  a  "drupe"  with  only  1  stone, 
which  is  compressed,  cartilaginous,  and  parchment-like;  2  of  the 
loculi  of  the  ovary  are  aborted.  (In  V.  opulus  the  marginal  flowers  of 
the  inflorescence  are  barren,  and  in  that  case  their  corollas  are  generally 
specially  large ;  the  cultivated  Viburnum  has  only  barren  flowers,  with  large 
corollas.) 


FIG.  592. — Sambucus  nigra :  cor  corolla;  s  calyx. 

3.  LiNN^EE.33.  Linncea  borealis  (the  only  species)  is  an  extreme  form  of  the 
•order  ;  it  has  a  2-flowered  dichasium,  funnel-shaped,  slightly  bilabiate  corollas 
(§) ;  4  didynamous  stamens.  Two  of  the  3  loculi  of  the  ovary  have  several 
ovules  which  are  not  developed,  while  the  third  has  only  1  ovule,  which  de- 
velopes  into  a  seed.  The  fruit  is  a  nut,  which  is  enveloped  by  the  two  large 
bracteoles,  which  are  covered  by  sticky,  glandular  hairs,  and  serve  as  a  means 
-of  distribution.  It  is  a  small  under-shrub. 

[Adoxa,  which  was  formerly  classed  in  this  order,  appears,  according  to  recent 
investigations,  to  be  more  properly  placed  among  the  Saxifraginee.] 

In  cases  where  the  flowers  are  small,  as  in  Sambucus  and  Viburnum  opulus, 
•they  are  rendered  conspicuous  by  being  arranged  in  closely-packed  inflores- 
cences; they  are  massed  together  and  form  large  surfaces,  and  in  the  lasi 
•named  are  still  more  conspicuous  on  account  of  the  barren,  but  large  ray- 
flowers,  which  are  of  service  in  this  respect.  Honey  is  secreted  in  the  nectaries 
<at  the  base  of  the  styles.  In  the  genera  with  rotate  flowers,  as  Viburnum  and 


556  D1COTYLEDONBS. 

other  Sambuceae,  the  honey  lies  so  exposed  and  in  such  a  thin  layer,  that  only 
flies  and  insects  with  short  probosces  can  procure  it ;  bees,  however,  visit  these 
flowers  for  the  sake  of  the  pollen.  There  is  hardly  any  nectar  in  the  Elder  ; 
self-pollination  frequently  takes  place.  The  flowers  of  the  Caprifoliaceas, 
which,  with  their  long  corolla-tube  are  adapted  for  evening-  and  night-flying 
insects  with  long  probosces,  open  in  the  evening,  and  at  that  time  give  off  their 
strongest  scent. 

DISTRIBUTION.  230  species ;  especially  outside  the  Tropics  in  the  Northern 
Hemisphere.  In  this  country  they  are  found  especially  in  hedges  and  as 
under-shrubs  — OFFICINAL  :  the  flowers  and  fresh  fruits  of  the  Elder  (S.  nifjra), 
the  fruits  ("  berries  ")  being  also  used  in  the  household.  ORNAMENTAL  SHRUBS  : 
species  of  Lnnicera,  Symphoricarpus,  Dier villa,  which  are  chiefly  from  N.  Am., 
Abelia  and  Viburnum. 

Family  34.     Dipsacales. 

The  leaves  are  opposite  and  without  stipules.  The  flower  (Figs. 
593,  595,  598,  599,  600)  is  epigynous,  zygomorphic  or  asymmetrical, 
5-merous  with  S5,  P5,  stamens  typically  5,  but  by  suppression 
never  more  than  4,  sometimes  less,  carpels  3-2.  The  calyx  is  more 
or  less  insignificant,  and  almost  suppressed  in  the  extreme  forms. 
The  ovary  has  3-1  loculi,  but  only  one  loculus  has  an  ovule,  which 
is  pendulous  with  the  micropyle  turned  upwards  (Fig.  594).  Fruit 
a  nut.  Embryo  straight,  with  the  radicle  pointing  upwards  (Fig. 
597),  without  or  with  endosperm. 

The  inflorescences  are  distinct  dichasia  in  Valerianacere,  but  in 
Dipsacacese  and  CalyceraceaB  they  are  crowded  together  into 
capitnla. 

This  family  is  closely  allied  to  the  Bubiales  through  the  Valerianaceaa.  which 
have  almost  the  same  structure  as  many  of  the  Caprifoliaceaa.  It  attains  the 
'highest  development  in  the  Dipsacaceae,  which  are  composite  plants,  but  differs 
from  Composite  in  the  position  of  the  ovule,  etc. 

Order  1.  Valerianacese.  Herbaceous  plants  or  under-shrubs 
with  opposite  leaves,  often  pinnate ;  stipules  absent.  The 
flowers  are  borne  in  dichasia  and  in  scorpioid  cijmose  inflor- 
escences and  are  entirely  without  any  plane  of  symmetry  (Fig.  593). 
The  calyx  and  corolla  are  5-merous,  but  the  calyx  is  frequently 
very  insignificant  and  ultimately  a  pappus,  as  in  Composite; 
the  corolla  is  frequently  saccate  or  produced  into  a  spur  at  the 
base.  Most  frequently,  only  3  (4r-l)  of  the  5  stamens  are  de- 
veloped ;  these  are  free.  Carpels  3,  which  form  an  inferior  ovary, 
often  with  3  loculi,  but  only  1  of  the  loculi  contains  1  pendidous, 
anatropous  ovule  (Figs.  593,594^1),  the  other  loculi  are  empty 


DIPSACALES. 


557 


and  shrink  up  more  or  less  completely.  (Compare  Fig.  593  A, 
B}.  Style  1,  stigma  tripartite.  Endosperm  absent;  embryo 
straight,  with  the  radicle  directed  upwards. 

The  inflorescences  are  dichasia,  or  unipared  scorpioid  cymes  with  the 
branches  developed  in  the  axil  of  the  second  bracteole,  Both  the  bracteoles  are 
generally  present  and  frequently  form  4  very  regular,  longitudinal  rows  on 
the  branches  of  the  inflorescence. — 5  stamens  do  not  occur  (except  perhaps  in 
Patrinia).  The  suppression  of  stamens  and  carpels  takes  place  most  readily 
on  the  anterior  side  of  the  flower  and  that  turned  towards  the  first  bracteole 

(a)  (Fig.  593),  whose  branch  is  suppressed  in  the  dichasium ;  after  this  the  pos- 
terior median  stamen  is  next  suppressed. 

By  the  vegetative  characters  as  well  as  • 

by  the  inflorescence  and  the  flower,  the 
order  is  allied  to  the  Caprifoliaceae  and 
especially  to  the  Sambuceae. 

In  the  least  modified  (oldest) 
forms,  Patrinia  and  Nardostachi/s, 
there  is  an  almost  regular  flower, 
a  5-merous  calyx,  4  stamens,  and 
3  loculi  in  the  ovary,  2  of  which 
however  are  barren.  The  stamens 
in  Voter ianella  are  reduced  to  3,  in 
F&dia  to  2  (posterior),  and  the 
calyx  is  less  distinctly  5-dentate  ; 
the  2  empty  loculi  in  the  ovary 
are  still  visible.  Fedia  has  a  small 
spur  at  the  base  of  the  corolla. 
Valeriana  has  a  very  reduced,  hair- 
like  calyx  (pappus),  an  unsymmet- 
rical,  salver-shaped  corolla  with  a 
sac-like,  nectariferous  spur  at  the 
base,  3  stamens  and  only  1  loculus 
in  the  ovary  (Figs.  594,  593).  Centranthus  (Fig.  593)  is  still 
further  reduced.  The  corolla  has  a  spur  and  only  1  stamen ; 
unipared  scorpioid  cymes  with  4  rows  of  bracteoles.  In  the  last  two  genera 
there  is  a  peculiar  wall  in  the  corolla-tube,  which  divides  it  longitudinally 
into  two  compartments  (indicated  by  a  dotted  line  in  Fig.  593),  one  of  which 
encloses  the  style.  This  wall  is  low  in  Valeriana,  but  in  Centranthus  it  reaches 
as  far  as  the  throat. — The  rays  of  the  pappus  are  pinnately  branched  and 
rolled  up  before  the  ripening  of  the  fruit.  12-20  in  number  (Fig.  594  A,  B). 

Val.  officinalis  and  others  are  protandrous :  in  the  first  period  the  stamens 
project  from  the  centre  of  the  flower  (Fig.  595  a),  the  stigma's  in  the  second 

(b)  when"  the  stamens  have  become  bent  backwards.      (V.  dioica  is  dioecious 


FIQ.  593.— A    Diagram   of    Valerian 
officinalis.    B  Diagram  of  Centranthus. 


558 


DICOTYLEDONES. 


with  large  $-  and  small  ?  -flowers).— 275  species ;  especially  from  the  temperate 
and  colder  parts  of  the  northern  hemisphere  of  the  Old  World,  Western  North 
America  and  the  Andes.—  Bitter  properties  are  characteristic,  such  for  instance 
as  the  volatile  acid  and  volatile  oil  of  Valeriana ;  these  occur  especially  in  the 


FIG.  594. — Valeriana  :   A  oVary  (longitudinal 
section) ;  B  ripe  fruit. 


FIG.  595. — Valeriann:  a  flower 
in  the  c?  stage ;  b  in  the  ? . 


rhizomes.  OFFICINAL  ;  the  rhizomes  of  V.  officinalis.  —  The  true  Indian 
"  Nardus,"  an  important  medicine  and  perfume  in  India,  is  extracted  from 
Nardos tacky s  (Himalaya).  A  variety  of  Valerianella  olitoria  is  sometimes 
used  as  salad. 

Order  2.  Dipsacaceae  (Teasels).  Herbs  with  oppose  leaves 
without  stipules.  The  flowers  are  situated  in  compact  capitula 
each  with  an  involucre.  A  characteristic  feature  of  the  order 
is  that  each  flower  of  the  capitulum  has  a  gamophyllous  "  epicalyx" 


FIG.  596.  —  Centranthus  riiber. 
Flower,  its  lowermost  portion  (the 
ovary  and  spur)  in  longitudinal 
section.  (Mag ) 


FIG.  697.— Scabiosa  atropurp'irea.  Fruitin lon- 
gitudinal section.  Inside  the  "  epicalyx  "  may 
be  seen  thp  fruit  drawn  out  into  a  beak,  with 
straight  embryo  and  radicle  directed  upwards. 


DIPSACALES.  559 

(Figs.  597,  599,  600),  which  envelopes  the  inferior  ovary.  The 
flowers  (Figs.  599,  600)  are  J  >  5-merous  (S5,  P5,  stamens  typi- 
cally 5,  G2),  but  the  calyx  often  expands  at  the  edge  into 
a  membrane  with  5,  or  an  indefinite  number  of  bristles  or  teeth 
(pappus.  Figs.  597,  600),  and  the  zygomorphic,  funnel-shaped  corolla 
is  sometimes  5-lobed  and  bilabiate  (f),  but  most  frequently  4- 
partite  (Fig.  599),  the  two  lobes  of  the  upper  lip  coalescing  into 
one  lobe,  as  in  certain  Labiates,  Veronica  and  Plantago;  the 
cestivation  is  imbricate. 


5yd.  59tt.  600. 

FIG.  598  -600.— Dtpsocus  /ullonum. 

FIG.  598.— Inflorescence  (the  flowers  in  a  zone  below  the  apex  commence  to  flower  first), 
FIG.  599. — Flower  (f).        FIG.  600. — The  same  in  longitudinal  section. 

The  stamens  are  never  more  than  4,  the  posterior  one  remaining 
undeveloped ;  they  usually  have  free  anthers  which  generally  project 
considerably  (Fig.  599).  The  ovary  is  unilocular  with  1  pen- 
dulous ovule  and  bears  1  undivided  style ;  fruit  a  nut  with  1  seed,, 
containing  endosperm  and  with  the  radicle  turned  upwards  (Fig.  597). 

The  flowers  do  not  always  open  in  centripetal  order,  a  fact  which  may  be 
observed  especially  in  the  Dipsacace®,  in  which  a  zone  of  flowers  round  the 
centre  of  the  capititlum  opens  first,  and  the  flowering  then  proceeds  both  up- 
wards and  downwards  (Fig.  598).  This  has  probably  some  connection  with  the 
fact  that  the  capitulurn  has  arisen  from  the  coalescence  of  several  dichasial 
inflorescences.  In  species  of  Scabiosa  the  flowers  open  simultaneously  at  the  cir- 
cumference, or  in  a  zone  at  the  centre. — The  morphological  explanation  of  the- 
W.  B.  00 


560  DICOTYLEDCKNES. 

*'  epicalyx"  is  not  quite  certain  ;  in  all  probability  it  is  formed  from  two  united 
bracteoles,  for  an  "epicalyx"  is  distinctly  formed  in  this  way  in  one  of  the 
Valerianaceas,  Phyllactis. — The  ray-flowers  are  larger  and  more  irregular,  labiate 
or  ligulate,  than  the  disc-flowers,  yet  not  in  so  high  a  degree  as  in  the  Com- 
positae. 

A.  A  scarious  bract  to  each  flower.  Scabiosa  has  a  5-lobed 
corolla;  the  "epicalyx"  has  a  dry,  scarious,  often  finally  large 
collar,  and  the  true  calyx  is  formed  of  long  bristles  (generally  5) 
(Fig.  597).  Succisa  pratemis  (Devil's-bit)  has  a  4-lobed  corolla,  the 
collar  of  the  "  epicalyx  "  is  herbaceous  ;  the  calyx  as  in  the  preced- 
ing.— Pterocephalus. — Dipsacus  (Teasel) ;  large,  spiny  and  stiff- 
haired  herbs  with  capitula,  or  short,  thick  spikes  on  which  both 
the  involucral-leaves  and  bracts  project  considerably,  and  are 
stiff  and  spinose  (Fig.  598).  The  "  epicalyx  "  has  short  teeth,  or 
is  almost  entire.  The  leaves  of  the  stem  unite  together  in  pairs,  so  that 
shallow  cups  are  formed  round  the  stems  in  which  rain-water  may  collect. — 
Cephalaria. — Marina  :  the  flowers  are  falsely  verticillate  as  in  the  Labiatae ; 
the  calyx  has  2  laterally-placed,  entire,  or  emarginate  lobes ;  2  stamens,  or  2 
large  and  2  small  ones. 

E.  Bristles,  but  no  true  bract  to  each  flower.  Knautia ;  the 
corolla  is  4-partite,  the  calyx  cup-like,  with  many  bristles  or 
teeth  on  the  edge. 

POLLINA.TION  is  in  many  species  effected  by  insects.  The  honey  is  secreted 
by  a  ring  round  the  base  of  the  style.  The  flowers  in  our  native  species  are 
considerably  protandrous.  Gynodioecious  flowers  also  occur. — 150  species ; 
especially  in  the  Mediterranean  and  the  Orient ;  the  order  is  not  represented  in 
the  South  Sea  Islands,  Australia  and  America.— The  heads  of  the  true  Teasel 
{Dips,  fullonum)  are  used  for  carding  wool,  on  account  of  the  elastic  bracts, 
which  are  hooked  at  the  point.  The  order  has  bitter  properties  ;  tanin,  etc. ; 
but  no  species  are  used  in  medicine  or  the  household. — Scabiosa  atropurpurea, 
•etc.,  are  used  as  ornamental  plants. 

Order  3.  Calyceraceae.  This  order  resembles  the  Composite  in  the  valvate 
.aestivation  of  the  corolla  and  the  more  or  less  united  stamens,  and  the  Dipsa- 
caceae  in  the  undivided  style,  pendulous  ovule  and  endosperm.  The  calyx  is 
frequently  composed  of  5  distinct  scales.  An  "  epicalyx  "  is  wanting. — 20 
species ;  America. 

Family  35.     Campanulinae. 

The  flower  is  epigynous,  perfect,  with  5  sepals,  5  petals,  and  5 
stamens  in  regular  alternation,  and  3  (2-5)  carpels.  The  sepals 
in  all  cases  are  distinct,  but  narrow  and  pointed,  so  that  the 
aestivation  is  open.  The  corolla  is  gamopetalous  with  (as  in  the 
Composite)  valvate,  or  slightly  infolded- valvate  aestivation.  The 


CAMPANDLINJB.  561 

stamens  are  nearly  always  situated  on  the  torus  without  being 
united  to  the  corolla  (Figs.  601,  604).  The  anthers  adhere  or 
unite  and  form  a  tube  with  introrse  anthers  from  which  the  pollen 
is  swept  out  by  the  projecting,  brush-like  hairs  on  the  style  (as  in 
the  Composite).  The  ovary  is  3- (2-5)  locular,  many  ovules  in 
each  loculus.  The  fruit  is  generally  a  many-seeded  capsule  (or 
berry).  Embryo  in  the  centre  of  a  fleshy  endosperm. — The 
majority  are  herbs  with  scattered  leaves,  without  stipules.  The 
presence  of  latex  and  inulin,  together  with  the'  tubular  formation 
of  the  anthers,  the  pollination,  etc.,  indicate  a  relationship  with 
the  Composites. 

The  Cucurbitace.(B  are  by  some  authorities  placed  in  this  family  as  being 
most  closely  related  to  the  Campanulaceas.  Although  the  corolla  is  most  fre- 
quently gamopetalous,  and  other  similarities  to  the  Carnpanulaceae  are  present, 
yet  on  account  of  the  structure  of  the  ovule,  and  for  other  reasons,  the  Cucur- 
bitaceae  are  here  placed  in  the  Choripetalae.  The  Campanulinse  without  doubt 
proceed  upwards  to  the  Compositae,  with  which,  in  addition  to  the  occurrence 
of  inulin  and  laticiferous  vessels  (Cichoriese),  there  are  many  corresponding 
features  both  in  the  structural  and  biological  relations  (epigyny,  valvate  aesti- 
vation of  the  corolla,  tendency  of  the  anthers 'to  adhere  or  unite,  protandry 
with  a  stylar-brush,  etc.)  The  inflorescence  of  Jasione  is  almost  identical  with 
that  of  the  Compositae. 

Order  1.  Campanulaceae  (Campanulas).  The  flowers  are 
regular  and  in  some  only  semi-epigynous,  5-merous,  except  in  the 
gynceceum  which  is  3-merous  (the  unpaired,  median  carpel  being 
generally  posterior),  more  rarely  2-5-merous,  and  has  a  corre- 
sponding number  of  stigmas  and  loculi  in  the  ovary ;  the  placen- 
tation  is  axile  with  a  large  number  of  ovules.  The  median  sepal 
is  posterior.  The  stamens  frequently  have  broad,  free  bases  (Fig. 
601  H)  which  cover  the  nectariferous  upper  surface  of  the  ovary; 
the  anthers  only  fit  loosely  together,  and  become  separated  as 
soon  as  the  pollen  is  shed  (Fig.  601  G).  1  long  style,  which  is 
studded  by  sweeping- hairs  (stylar-brush),  which  ultimately  be- 
come invaginated;  the  stigmas  do  not  unfold  until  the  stamens 
have  shed  the  pollen  (Fig.  601  Ey  G).  Fruit  a  capsule. — Herbs, 
more  rarely  under-shrubs  or  shrubs,  with  latex  and  scattered, 
undivided  leaves  without  stipules.  The  inflorescence  is  most  fre- 
quently a  raceme  or  spike  with  terminal  flower. 

A.  Capsule  opening  at  the  side  by  pores  and  small  valves, 
Campanula  (Canterbury-bell)  ;  the  corolla  is  bell-shaped,  rarely 
almost  rotate  ;  capsule  obconical.  The  pores  of  the  capsule  are  found 
near  the  top  of  the  fruit  when  it  is  erect,  and  near  the  base  when  it  is  pendu- 


562 


DICOTYLEDONES. 


lous,  so  that  the  seeds  are  not  liberated  unless  the  capsule  is  forcibly  shaken, 
and  they  are  thus  ejected  to  a  considerable  distance.  —  Phytcuma  (Ram- 
pion)  has  free  petals,  which  for  a  long  time  adhere  at  the  apex  and 
form  a  tube  round  the  stamens  (Fig.  601);  inflorescence  compact, 
spike-like  or  capitate,  in  the  latter  case  resembling  that  of  the 
Composites,  and  frequently  with  an  involucre  similar  to  the  one 
possessed  by  this  order.  Specularia  (rotate  corolla,  prismatic  capsule), 
Michauxia  (flower  8-merous). — Symphyandra  has  syngenesious  anthers. 

B.  Capsule  with  valves  at  the  apex,  loculicidal  dehiscence  : 
Jasione ;  the  petals  are  almost  free.  The  anthers  are  united  at 
the  base  (syngenesious).  The  flowers  are  situated  in  capitate 
umbels  with  involucres. —  Wahlenbergia  ;  Platycodon. 


FIG.  601.— Phyteuma  spicatum.    Flowers  and  parts  of  flowers  in  various  stages  of 
development. 

C.    Berry :  Canarina ;  flower,  6-merous  ;  leaves  opposite. 

Protandry  is  general  (Fig.  601).  510  species ;  principally  in  temperate 
countries.  Several  genera  furnish  ornamental  plants,  but  are  of  little  use  for 
other  purposes.  The  roots  of  some  Campanula-  and  Phyteuma-species  are 
large  and  may  serve  as  pot-herbs  (C.  rapunculiis,  P.  spicatum). 

Order  2.  Cyphiaceae.  In  this  order  the  corolla  is  zygomorphic  and  the 
stamens  free,  hence  it  is  intermediate  between  orders  1  and  3. — About  24 
species;  Africa. 

Order  3.  Lobeliaceae  (Lobelias).  This  order  may  briefly  be- 
described  as  Campanulacese  with  zygomorphic  flowers  and  anthers 
united  into  a  tube,  in  most  cases  slightly  bent ;  generally  2  carpels 
and  an  inverted  position  of  the  flower,  i.e.  the  median  sepal  is 


CAMPANULIN.E. 


563 


tnrued  anteriorly  (Fig.  602)  (a  position  which  is  found  to  occur 
within  the  Campanulacege).  A  twisting  of  the  peduncle  takes 
place  even  before  flowering  (as  in  the  Orchids)  so  that  the  ordin- 
ary position  of  the  5-merous  Dicotyledons  appears  to  be  restored. 
The  zygomorphy  of  the  flower  is  especially  present  in  the  corolla, 
which  has  a  bipartite  under-lip  and  a  tripartite  upper-lip,  and  is, 
in  Lobelia,  anteriorly  (apparently  posteriorly)  deeply  cleft  (Fig.  602). 
There  is  1  style,  but  the  stigma  is  capitate  and  bilobed  and  sur- 
rounded at  its  base  by  a  whorl  of  hairs,  which  assists  in  pollination 
(as  a  stylar-brush)  in  the  same  manner  as  the  sweeping-hairs  in  the 
Campanulacese  and  Composite.  There  is  no  terminal  floiver  in  the 
spicate,  or  racemose  inflorescences. — Lobelia  has  a  capsule,  several 
others  have  berries.  Isotoma  (regular  flower);  Heterotoma  has  a  spur; 


FIG.  602. — Diagram  of  Lobelia 

fnlgens.  FIGS.  603,  601.— lobelia  syphilitica. 

FIG.  603.— Flower  (Y).      FIG.  601. — Longitudinal  section 
of  the  same. 

Siplwcampylos ;  Lysipoma  (pyxidium) ;  Clintonia  (1-locular  fruit).  Metzleria 
(all  the  petals  are  free). 

Entomophilous  and  protandrous.  About  500  species,  especially  in  the 
Tropics;  in  this  country,  L.  dortmanna  (margin  of  lakes). — Several  are  culti- 
vated in  gardens  and  conservatories  as  ornamental  plants  (Lobelia  bicolor, 
£rimts,fulgens,  etc.,  Siphocampylos,  Centropogori).  The  latex  of  several  species 
of  '1  upa  is  poisonous ;  caoutchouc  is  also  obtained  from  them.  OFFICINAL  : 
"herba  Lobelia"  (the  alkaloid  lobeline)  from  the  poisonous  L.  inftata,  (N. 
Am.). 

Order  4.  Goodeniaceae.  Chiefly  Australian  (200  species),  closely  related  to 
Orders  3  and  5,  but  without  latex.  The  style  is  provided  with  a  "  collecting- 
cup  "  which  receives  the  pollen  before  the  flower  opens  ;  it  has  a  small,  hairy 
aperture  through  which  the  pollen  is  forced  out  by  the  stigmas,  and  through 
which  they  emerge  when  the  pollen  is  shed  ;  it  is  sensitive  and  exhibits  move- 


564  DICOTYLEDONES. 

ments  when  touched. — Herbs,  under-shrubs,  less  frequently  shrubs.     Goodenia, 
Leschenaultia,  Sccevola. 

Order  5.  Stylidiacese  (or  Candolleaceae) ;  100  species,  the  majority 
Australian  ;  zygomorphic  flowers,  but  with  the  ordinary  position.  The  anterior 
petal  is  very  small.  The  chief  characteristic  feature  is  the  presence  of  only  2 
stamens  (with  extrorse  anthers)  which  are  united  with  the  style  and  form  a 
stylar-column ;  this  is  bent  like  a  knee  and  sensitive  at  the  bend  to  such  a 
degree  that  when  toucbed  it  jerks  violently  across  the  flower  to  the  opposite 
side  and  then  loses  its  sensitiveness. — Herbs,  less  frequently  under-shrubs. 
Stylidium  (Candollca}. 


Family  36.     Aggregatse. 

The  flowers,  which  are  borne  in  "capitula"  (Figs.  605,  610), 
are  epigynous  (Fig.  605  0,  D),  5-merous  in  the  calyx,  corolla  and 
androecium,  the  corolla  is  valvate  in  aestivation,  with  2  carpels 
(S5,  P5,  A5,  G2).  The  anthers  are  united  into  a  tube  (syngene- 
sious)  (except  Ambrosiece]  which  surrounds  the  bifid  style.  There 
is  never  more  than  1  loculus  in  the  ovary,  with  1  erect,  anatropous 
ovule.  The  fruit  is  a  1-seeded  nut  (cypsela),  with  thin  pericarp, 
the  calyx  generally  persists  as  a  tuft  of  hairs  (pappus}  (Fig.  606) 
on  the  summit  of  the  fruit.  Embryo  without  endosperm ;  the 
radicle  directed  downwards. 

Only  1  Order :  Compositor. 

With  respect  to  the  inflorescence  and  the  development  of  the 
individual  flowers,  there  is  a  very  close  resemblance  to  the  Dipsa- 
caceae,  which  stand  on  the  same  plane  of  progression  as  the 
Composite.  But  while  the  latter  are  allied  to  Campanulinae  as 
the  last  stage  in  the  process  of  evolution,  the  Dipsacacese  form  the 
final  stage  of  the  Rubiales-Dipsacales. 

Order  Compositae.  (For  the  principal  characteristics  com- 
pare those  of  the  family.)  The  Compositae  are  chiefly  herbs, 
but  trees  and  shrubs  also  occur  in  tropical  countries.  The  leaves 
may  be  scattered  or  opposite,  but  have  no  stipules.  The  outer 
leaves  of  the  involucre  as  a  rule  are  barren,  especially  when 
numerous  and  imbricate,  while  the  innermost  ones  support  the  ray- 
flowers  of  the  capitulum  ;  in  a  few  instances  all  are  fertile  (e.g. 
Tragopogon,  Tagetes).  The  CAPITULA  are  many-flowered,  with  the 
exception,  e.g.  of  Echinops,  which  has  1-flowered  capitula  (see  page 
570).  The  capitula  are  again  arranged  in  inflorescences,  most  fre- 
quently corymbose  with  centrifugal  order  of  development.  The 
form  of  the  receptacle  is  an  important  character  for  the  division  of 
the  genera  (flat,  convex,  conical),  and  also  the  presence  of  scales; 


AGGREGATE. 


565 


these  may  be  one  scale  (bract)  for  each  flower  (Fig.  610  br),  or  a 
large  number  of  bristles,  which  do  not  each  correspond  to  a  leaf,  or 
the  receptacle  may  be  entirely  without  covering  (naked).  The  flowers- 
open  in  acropetal  order  in  each  capitulum.  All  the  flowers  in  a 
capitulum  may  be  of  the  same  sex,  and  their  form  and  colour  are  in 
that  case  the  same,  or  the  sexes  may  be  different,  in  which  case  the 
form  and  colour  are  also  most  frequently  different :  the  ray -flowers- 
have  projecting  labiate  or  ligulate  corollas,  while  the  disc-flowers 
have  tubular  corollas.  As  a  rule  in  the  latter  case  the  $  -flowers 


A.FAGUET.dtl 


FIG.  605.  —  Calendula  arvensis:     A  capitulum;    -B  capitulum  in   longitudinal   section; 
C  ?•  flower;  D  £ -flower;  E  the  stamens;  ^capitulum  with  ripe  fruits;  G  ripe  fruit. 

are  at  the  circumference,  and  the  £  in  the  centre,  less  frequently 
$ -flowers  at  the  edge  and  (J -flowers  in  the  centre.  The  ray- 
flowers  in  some  genera  are  neuter  (e.g.  Centaur  ea}.  Some  are 
dioecious. 

There  is  no  trace  of  an  epicalyx  (in  contrast  to  the  Dipsacaceae, 
which  they  generally  so  resemble).  The  formation  of  the  CALYX 
is  very  varied.  The  calyx  always  consists  of  a  very  small 
cushion -like  structure,  most  frequently  developed  later  than  the 
corolla  ;  the  5  corners,  which  correspond  to  the  5  sepals,  in  a  few 


566 


DICOTYLEDONES. 


instances  are  raised  as  5  large,  flat,  membranous  bodies,  e.g.  in 
species  of  Xcranthemum,  Catananche,  Sphenogyne,  etc. ;  in  otlier 
instances  each  of  these  bears  a  shorter  or  longer  bristle  on  its 
apex,  followed  by  others  in  rather  uncertain  numbers  and  with 
but  slight  indications  of  order,  on  the  edge  and  on  the  outer  side 
of  the  calyx  between  the  5  points ;  in  other  instances,  again,  the 
calyx  is  covered  with*  bristles  and  hairs  without  any  indication  of 
order  or  definite  number  (Fig.  606  a,  b)  ;  finally  instances  occur 
in  which  the  edge  is  raised  as  a  membranous  collar,  irregularly 
toothed  and  'notched,  or  divided  into  small  scales.  There  are 
naturally  differences  in  the  means  of  distribution  corresponding  to 


FIG.  630. — a  Fruit  of  Taraxacum;  b  of  Senecio;  c  of  B/flcns. 

the  differences  in  structure  of  the  calyx.  The  fruits  a  and  &  re- 
presented in  Fig.  606  are  distributed  by  the  wind,  those  like  c,  on 
the  other  hand,  by  attaching  themselves  to  animals  and  human 
beings.  The  rays  of  the  pappus  are  termed  rough  when  special 
cells  project  a  little  beyond  the  surface,  but  if  the>e  grow  out,  arid 
are  hair-like,  the  pappus  is  said  to  be  feathery.  In  some  genera 
the  pappus  is  raised  011  a  long  stalk,  which  is  developed  from  the 
upper  part  of  the  fruit,  and  termed  a  beak  (Fig.  606  a).  The 
pappus  does  not  attain  its  full  development  till  the  ripening  of 
the  fruit,  i.e.  until  it  is  about  to  be  of  use. 

The  COEOLLA  has  various  forms  :  (a)  tubular  (Fig.  605  D),  with  a 


AGGREGATE. 


567 


shorter  or  longer  tube,  not  always  of  the  same  bore  throughout 
and  especially  slightly  widened  at  the  top  to  form  a  bell-shaped 
opening,  with  5  regular  teeth :  (6)  labiate  after  -f ,  i.e.  with  2 
petals  in  the  upper  and  3  in  the  under  lip  :  (c)  ligulate,  i.e.  the 
corolla  is  split  for  a  considerable  distance  on  the  posterior  side  (as 
in  the  Labiate  genus  Teucrium)  and  prolonged  into  a  long,  strap- 
like  portion  (Fig.  609  A},  which  projects  upwards.  A  distinction 
must,  however,  be  drawn  between  the  true  and  false  ligulate 
corolla.  In  the  first  case  the  corolla  has  5  teeth  at  the  apex  (Fig. 
609  A}  and  is  made  up  of  all  the  petals  of  the  corolla  united 
together ;  this  is  the  usual  condition  in  the  Ligulate-floivered.  In 
the  latter  case  (Fig.  605  G)  the  tongue  has  only  3  teeth  (or  is 
more  irregularly  2-3-dentate),  and  is  only  formed  of  3  petals  ; 
the  corolla  is  then  truly  bilabiate,  the  tongue  is  the  large  under  lip, 
and  the  upper  lip  is  very  slightly  developed,  or  even  at  an  early 
stage  quite  suppressed.  This  false  "ligulate"  corolla  is  found 
among  the  ray-flowers ;  sometimes  the  upper  lip  is  seen  quite 
plainly,  e.g.  in  Tagetes,  especially  in  the  double  capitula.  The 
VENATION  of  the  corolla  is  peculiar ;  there  are  always  commisural  veins  which 
branch  dichotomously  at  the  angles  between  the  teeth  of  the  corolla,  and  send 
a  branch  into  the  edge  of  the  two  nearest  teeth.  The  midrib  is  frequently 
absent,  but  may  be  present,  and  then  it  has  sometimes  no  connection  with  the 
other  veins  of  the  corolla. 

The  STAMENS  are  attached  to  the  corolla,  and  have  free 
filaments  (Silybum  has  united  filaments),  but  the  anthers,  which  at 
first  are  free,  adhere  together  and  form  a  tube  (Fig.  605  E  :  only 
Ambroneze  have  free  anthers).  The  connective  is  generally  pro- 
longed, and  protrudes  above  the  anthers 
as  a  thin,  brown  membrane  of  various 
forms  (Fig.  605  E)  ;  appendages  of 
various  forms  may  also  be  found  at  the 
base  of  the  anthers.  The  anthers  open 
introrsely,  and  the  pollen  must  be  carried 
out  at  the  top  of  the  tube  by  upward 
growth  of  the  style,  and  by  means  of  the 
"stylar-brush"  (Figs.  607,  608,  609) ;  the 
filaments  are  sometimes  sensitive  {e.g.  in 
the  Corn-flower,  Fig.  607),  and  shorten 
on  being  touched,  so  that  the  anther- 
tube  is  pulled  downwards,  and  the  pollen 
swept  out  at  the  top  (Figs.  607,  608 
A,  B). 


FIQ.  607. — Centaurea  cyanus: 
A  the  anther-tube  (*t)  with  the 
crescentic  curved  filament  be- 
fore irritation;  g  the  style;  fc 
the  base  of  the  corolla;  .B  the 
same  after  irritation,  the  an- 
thers are  drawn  further  down. 


568 


DICOTYLEDONES. 


The  STYLE  divides  at  the  apex  into  two  branches  (Figs.  609,  610)r 
both  of  which  generally  bear  on  the  inner  surface  two  lines  of 


Fm.  608. — Cirsium  arvense  :  A  the  upper 
portion  of  a  flower,  the  pollen  (e)  is  being 
ejected ;  B  part  of  the  upper  portion  of  the 
style  with  stylar-brush(b,  c)  and  the  stigmatic 
papilla?  (d). 


FIG.  609. — Le&ntodon  autumnale  :  A 
ligulate  flower;  B  extremity  of  the- 
style  with  stylar-brush  (a),  stigma  (b) 
and  pollen-grains  (c).  C  Centaurea- 
cyanus. 


stigmatic  papillae  (Fig.  610  B,  C)  and  being  in  shape,  etc.,  very 
varied,  are  therefore  employed  as  systematic  characters. — The  most 
important  types  are  :  A.  The  style  is  uniformly  cylindrical ;  its  branches  are 
semi-cylindrical,  long,  and  with  long  hairs,  and  finally  bend  backwards;  the 
stylar  branches  bear  slightly  projecting  stigmatic  papillae  on  the  inner  side. 


FIG.  610.—  Achillea  millefolium. 


This  form  is  characteristic  of  the  Cichoncte  (Fig.  609  A  B).     B.  The  style  i& 
uniformly  cylindrical ;  the  branches  are  long,  cylindrical  or  club-like,   short. 


AGGREGATE.  5G9 

not  rolled  back,  with  fine  hairs  externally ;  the  stigmatic  lines  do  not  reach 
beyond  the  centre,  and  do  not  meet  together.  Characteristic  of  Eupatorium, 
Petasites,  Tussilago.  C.  The  style  is  thickened  beneath  the  stigmatic  branches 
in  the  form  of  a  knob,  or  very  hairy  (Fig.  609  C) ;  the  stigmatic  lines  reach  as 
far  as  the  apex  of  the  branches  and  then  converge  ;  sometimes  the  stigmatic 
branches  are  united  as  far  as  the  apex.  Characteristic  of  the  Cynarece.  D. 
The  stylar  branches  are  lanceolate,  or  linear,  pointed  ;  externally  flat  and 
thickly  covered  with  hairs  in  the  tipper  portion ;  the  stigmatic  lines  cease 
where  the  hairs  commence  externally.  Characteristic  of  Aster,  Bellis,  Inula, 
Dahlia,  etc.  E.  The  stylar  branches  are  linear,  with  long,  brush-like  hairs  at 
the  apex,  where  they  are  either  abruptly  cut  off  or  prolonged  into  a  very  hairy, 
conical  appendage ;  the  stigmatic  lines  are  broad,  reach  as  far  as  the  brush-like 
hairs,  and  do  not  meet  together  (Fig.  610).  Characteristic  of  Senecio,  Helian- 
thus,  Xaiithium,  Gnaphalium,  Artemisia,  Anthemis,  and  others  related  to  these. 

A  ring-like  nectary  is  found  round  the  base  of  the  style. 

The  thin-walled  cypsela  (Fig.  606),  with  seeds  fitting  closely  to 
the  pericarp,  ha-s  many  different  forms  (smooth,  ribbed,  spined, 
etc.) ;  its  point  of  attachment  generally  lies  at  the  lowest  end 
but  sometimes  it  is  drawn  obliquely  up  the  side  (Centaurea,  etc.). 
The  calyx,  persistent  on  the  apex  of  the  fruit,  has  been  described 
above.  Some  genera  have  two  or  three  different  forms  of  fruits  in 
each  capitulum. — The  embryo  is  straight,  with  the  radicle  turned 
downwards,  and  without  endosperm,  but  is  rich  in  oil. 

The  variously  flowered  capitula,  whose  normal  tubular  disc- 
flowers  have  been  changed  to  ligulate  flowers,  may  be  termed 
"  double  flowers." 

The  relationship  of  the  Composite  to  the  Campannlinse  has  been  described 
above  (page  561).  The  alliance  with  the  Dipsacaceas  is  more  apparent  than 
real.  Similar  capitate  inflorescences  also  occur  as  the  final  stage  in  other  lines 
of  descent,  as  in  Eryngium  among  the  Umbelli ferae. 

I.  Cynareae,  Thistle  Group.  Flowers  all  $ ,  regular,  with 
tubular  corollas.  The  receptacle  is  covered  with  numerous  bristles, 
which  surround  the  flowers  without  any  definite  order,  or  the 
edges  of  the  grooves  in  which  these  are  placed  have  a  well-marked 
fringe.  The  involucral  leaves  are  numerous,  imbricate,  and  are 
either  prolonged  into  a  thorn  or  terminate  with  a  membranous 
edge.  The  style  has  been  described  on  page  568  (Fig.  609  C). 
Nearly  all  have  a  hairy  or  feathery  pappus.  The  filaments  are 
sensitive. 

Carduus  (Thistle)  ;  capitula  ovoid ;  involucral  leaves  compact, 
imbricate,  with  thorny  points  ;  the  pappus-rays  are  hair-like  and 
united  at  the  base  by  a  ring  (i.e.  the  calyx),  and  fall  off  together. 
—Cirsium  (Fig.  608)  has  a  feathery  pappus,  in  other  respects  it 
is  like  Carduus.  C.  arvense  reproduces  and  passes  the  winter  by  means  of 


570  DICOTYLEDONES. 

suckers. — Cynara  (Artichoke)  has  a  feathery  pappus  and  large, 
solitary  capitulum,  with  broad  involucral  leaves ;  these  have  a 
fleshy  base  like  the  receptacle  (edible). — Silylum  has  united  fila- 
ments. S.  marianum  (Milk  -  thistle),  has  leaves  with  numerous  white  spots. 
Onopordon  (Cotton-thistle).  Cnicus  (C.  benedictus)  has  a  large,  many-spined 
thorn  on  the  involucral  leaves;  pappus  trimorphic. — Lappa  (Burdock)  is 
easily  recognized  by  the  hooked  involucral  leaves,  which  assist  in  the 
distribution  of  the  fruit ;  in  this  respect  it  differs  from  the  other 
inflorescences,  and  also  in  the  fact  that  the  pappus  is  short,  and 
quickly  falls  off,  without  serving  as  a  means  of  distribution. — 
Carlina  ;  the  external  involucral  leaves  are  leafy,  thorny,  with  branched  or  un- 
branched  spines  standing  straight  out  or  bent  backwards ;  the  internal  ones  are 
dry,  and  prolonged  as  dry,  coloured,  radiating  scales.  The  well-developed 
bristles  on  the  receptacle  and  edge  of  the  calyx  are  deeply  cleft  and  lobcd. 
— Centaurea  (Knap-weed,  Fig.  607).  The  ray-flowers  are  neuter, 
and  generally  larger  than  the  disc-flowers ;  the  involucral  leaves 
are  regularly  imbricate,  but  are  frequently  provided  at  the  apex 
with  a  dry,  chaffy,  often  lobed,  fringed  appendage.  The  attach- 
ment of  the  fruit  is  lateral.  Serratula  (Saw-wort). — Carthamus,  the 
outer  and  inner  involucral  leaves  differ  very  much. — Echinops  (Globe- thistle) 
is  characterised  by  having  "  compound  capitula,"  i.e.  there  is  only 
one  flower  in  each  capitulum,  but  many  such  capitula  are  collected 
into  a  spherical  head,  which  at  the  base  may  also  have  a  few 
involucral  leaves.  The  individual  capitula  have  narrow,  linear 
involucral  leaves.  (There  are  altogether  about  150  species  of  Compositae 
with  1-flowered  capitula,  all  from  warm  countries.) — Xeranthemum,  IStaelie- 
lina,  Jurinea,  Saussurea,  etc. 

2.  Mutisieae,  Labiate-flowered  Group.  Tropical  (S.  American)  forms 
whose  zygomorphic  flowers  have  a  bilabiate  corolla  (|).  *The  involucre  is 
nearly  the  same  as  in  the  Thistles. 

3.  Cichorieae,  Chicory  Group  (or  LIGULJFLOR^).  The  flowers 
are  all  §  and  have  a  ligulate,  ^-dentate  corolla.  The  stylar  branches 
are  thin  and  prolonged  (Fig.  609  J5).  Laticiferous  vessels  occur  in 
the  majority  (in  this  feature  they  resemble  the  Lobeliacene  and 
Campanulacea3). 

A.  The  pappus  is  wanting,  or  it  is  scale-like,  but  not  long  and 
hairy. — Cichorium  (Chicory)  ;  capitula  with  blue  flowers,  borne 
singly  or  a  few  together  in  the  leaf-axil  ;  there  are  two  whorls  of 
involucral  leaves,  an  outer  one  of  short  and  radiating,  an  inner 
of  more  numerous,  longer  and  erect  leaves  ;  pappus,  scale-like. — 
Lapsana  (Nipplewort).  The  few  involucral  leaves  are  nearly 
of  the  samesize,  and  persist  forming  a  sort  of  capsule  round  the 


AGGREGATE.  571 

fruits,  which  are  entirely  without  a  pappus.  There  are  only  a  few 
flowers  in  the  small  capitula. — Arnoseris  (Swine's  -  succory )y 
Catananche,  etc. 

B.  The  pappus  is  long  and  liairy    (not  branched),  generally 
fine  and  snowy- white.     There  are  no  scales  on  the  receptacle.     The 
two  genera  first  considered  have  beaked  fruits. — Taraxacum  (Dan- 
delion) (Fig.  606  a) ;    the  capitula  are  many -flowered,   and  borne 
singly  on  the  top  of  a  leafless,  hollow  stalk. — Lactuca  (Lettuce) 
has  many  small,  few-flowered  capitula  borne  in  panicles. — Crepis 
(Hawksbeard). — Hieracium    (Hawk- weed)    has    many    imbricate 
involucral  leaves,  and  a  stiff,  brittle,  brownish  pappus. — Sonchus 
(Sow-thistle) ;  the  capitula,  when  a  little  old,  have  a  broad  base,, 
and  are  abstricted  above  in  the  form  of  a  jug;  involucral  leaves- 
imbricate  ;  the  fruit  is  compressed,  without  a  beak,  ridged.     The- 
soft,  white  pappus  falls  off  collectively. 

C.  The  pappus  is  feathery  and  branched;  no  scales  on  the  recep- 
tacle.— Tragopogon  (Goat's-beard)  generally  has  8  involucral  leaves 
in  one  whorl.     The  fruit  has  a  long  beak;  the  rays  of  the  pappus 
are  interwoven  in  the  form  of  an  umbrella. — Scorzonera  has  fruits 
like  the  preceding, but  almost  without  any  beak;  involucral  leaves 
many,    imbricate. — Leontodon    (Hawkbit)   has  a  slightly  feathery 
pappus,  rays  not  interwoven  ;  beak  absent. — Picris. 

D.  Long,  chaff -like,  deciduous  scales  on  the  receptacle  ;  pappus 
feathery. — Hypochceris  (Cat's-ear). 

4.  Eupatorieae,  Hemp-agrimony  Group.     All  the  flowers 
are    most  frequently    §  '•>  corollas  tubular  and  regular ;    the  in- 
volucral   leaves    are   not   stiff    and   spiny;    the   receptacle  is  not 
covered  with  stiff  bristles.     The  stylar  branches  are  long,  club- 
like,  or    gradually   tapering.     There   is   no    swelling    below   the 
stigma. 

Eupatorium  (Hemp-agrimony)  ;  all  the  flowers  are  5  • — Petasites 
(Butterbur)  ;  ray-flowers  ?  ,  disc-flowers  $  or  6*  '•>  sometimes 
dioecious.  Capitula  in  racemes1  or  panicles.  The  leaves  develop 
after  the  flowering. — Tussilago(T.  farfara,  Colt's-foot)  has  a  soli- 
tary capitulum  borne  on  a  scaly,  scape-like  stem;  the  ray- 
flowers  are  $  with  ligulate  corollas,  disc-flowers  $ .  The  leaves 
unfold  after  the  flowering.  Ageratum,  Mikania,  Vernonia. 

5.  Asterese,  Aster  Group  (or  RADIATE,  Ray-flowered).    The 
flowers    are    of    two   forms    and   different   sexes;    the  ray-flowers 
are    ?    (sometimes  neuter),  most  frequently  with  irregular,  falsely 
ligulate,  radiating  corollas  ;  the  disc-flowers  are  ?  ,  regular,  with: 


572  DICOTYLEDONES. 

tubular  corollas  (Fig.  6.10).  Sometimes  only  tub  alar  flowers  are 
present,  as  e.g.  in  Senecio  vulgar  is  (Groundsel),  and  the  exterior 
of  the  capitulum  is  then  as  in  the  Eupatoriese.  The  stylar 
branches  are  straight,  more  or  less  flat  and  short  (Fig.  610). 

A.  ANTHEMIDEJ:.     Involucral  leaves  imbricate,  generally  mem- 
branous at  the  edge ;    pappus  wanting,  or  at  most  a  membranous 
margin  to  the  calyx,  but  without  hairs. 

f.  Chaff-like  bracts  on  the  receptacle  are  found  in  Anthemis 
(Chamomile),  Anacyclus  (A.  officinarum) ,  Achillea  (Milfoil,  Fig.  610), 
Sa'idolina,  etc. 

ft-  A  'naked  receptacle  is  found  in  the  following :  Bellis 
(Daisy)  has  solitary  capitula  on  leafless  stalks  with  white  ray- 
flowers. — Matricaria  (Wild  Chamomile)  has  a  conical  receptacle. 
(M.  chamom ilia  has  a  very  high,  hollow  receptacle  ;  M.  inodora  has  large,  odour- 
less capitula,  and  the  receptacle  is  not  hollow.)  —  Chrysanthemum  (Ox- 
eye)  most  frequently  large,  solitary  capitula ;  flat  receptacle. — 
Pyrethrum ;  pappus  scanty. — With  these  are  classed  Tanacetum 
(Tansy)  and  Artemisia  (W^ormwood)  with  tubular  corollas  only. 

B.  HELIANTHE.E.     Most  frequently  a  bract  to  each  flower  is 
found  on  the  receptacle.     The  pappus  is  never  exactly  hairy,  but 
-consists  of  scales,  spines,  etc.,  and  the  fruits  are  most  frequently 
•compressed   (Fig.    606  c). — Helianthus  (Sun-flower)  ;  H.   tuberosus 
(Jerusalem  Artichoke)  has  tuberous  underground  stems.     Dahlia 
has  tuberous  roots  (Am.).     Bidens  (Bur-marigold,  Fig.  606  c) ;  the 
fruits    are    compressed  with    2    (or  more)    spines    provided  with 
ireflexed  barbs. — Calliopsis;  Rudbeckia;  Zinnia;  Tagetes  has  united 
involucral  leaves,  and  yellow,  transparent  oil-glands.     Spilanthes, 
•Galinsoga,    Melampodium,    Silphium    (Compass-pla.nt),     Helenium, 
•Gaillardia. 

C.  CALENDULE^  have  1-2  rows  of  involucral  leaves,  a  naked 
receptacle,  and  large,  crescent-shaped,   irregularly  warted  fruits, 
-of   different  forms  in  the  same  capitulum;    pappus  absent  (Fig. 
605). — Calendula  (Marigold);  ray-flowers   ?,  disc-flowers  <^. 

D.  SENECIONE.E,  have  a  fine,  hairy,  white  pappus;  no  bracts, 
•otherwise    as    in   Anthemideae.     The    involucral  leaves   are  most 
frequently  in    1-2    rows. — Senecio   (Groundsel)    has    two    whorls 
-of  involucral  leaves,  which  most  frequently  have  black  tips,  the 
external  being  much  shorter  than  the   internal  ones  ($.  vulgaris 
has   all   flowers    $    an(^   alike). —  Cacalia,    Doronicum,     Cineraria, 
Jjigularia,  Arnica  {A.  montana  ;  large,  long-stalked  capitula  ;  leaves 
opposite,  forming  a  kind  of  rosette). 


AGGREGATE.  573 

E.  AsTEREa:  have  a  bristle-like,  unbranched  pappus,  often  of  a 
dingy  brown  ;  receptacle  naked ;  involucral  leaves  numerous,  im- 
bricate.— Solidago  (Golden-rod)  ;  capitula  small,  yellow-flowered, 
borne  in    panicles.     Aster;   disc-flowers   most  frequently   yellow, 
ray-flowers  violet;    Callistephus ;    Erigeron  (Flea-bane) — Inula. — 
All  the  corollas  are  tubular  in  :   Gnaphalium  (Cud-weed)  ;  involu- 
cral leaves   dry,  rattling,  often  coloured;  the  foliage-leaves   and 
stem  often  white  with  woolly  hairs  ;  ray-flowers   ?  ,  with  narrow, 
tubular   corolla;  disc-flowers    $     (few).     Antennaria  (Cat's-foot ; 
dioecious),  Filago,  Helichrysum,  Ammobium,  Rliodanthe  and  others. 
Leontopodium  (L.  alpinum,  "  Edelweiss  "'). 

F.  AMBKOSIE^E,  a  very  reduced  type  of  Composite,  differing  from  the  others 
in  having  free  anthers  ;  the-capitula  are  generally  unisexual,  monoecious,  the  $ 
borne  in  a  terminal  inflorescence,  the  ?  in  the  leaf-axils.     In  other  respects 
they  are  most  closely  related  to  Helianthece. — Xanlhium.     In  the  $  -capitula 
there  are  many  flowers  without  calyx,  but  with  tubular  corolla  and  free  invo- 
lucral leaves.    In  the  $  -capitula  there  are  only  2  flowers,  which  are  entirely 
destitute  of  both  calyx  and  corolla  ;  involucral  leaves  2-spined,  united  to  form 
an  ovoid,  bilocular  envelope,  each  compartment  containing  one  flower.     The 
envelope  of  involucral  leaves  unites  with  the  fruits,  enclosing  them  at  maturity 
with  a  hard  covering  from  which  numerous  hook -like  spines  project,  assisting 
very  greatly  in  the  distribution  of  the  fruit.     The  whole  structure  thus  finally 
becomes  a  1-  or  2-serded  false  nut. — Ambrosia,  the  ?  capitulum  1-flowered. 

POLLINATION.  The  flowers  are  somewhat  insignificant,  but  become  very  con- 
spicuous owing  to  a  number  being  crowded  together  in  one  inflorescence.  The 
corollas  of  the  ray-flowers,  being  often  very  large  (Asterece ;  Centaurea),  fre- 
quently render  the  capitula  still  more  conspicuous.  The  capitula  display  many 
biological  phenomena  similar  to  those  often  shown  by  the  individual  flowers  in 
other  orders,  e.g.  by  periodically  opening  aud  closing,  in  which  the  involucral 
leaves  resemble  the  calyx  in  their  action.  (The  name  "  Compositae  "  originates 
from  the  term  "  flos  compositus,"  composite  flower).  An  abundance  of  honey 
is  formed,  which  to  some  extent  fills  up  the  corolla-tube,  and  since  insects  may 
visit  a  number  of  flowers  in  the  course  of  a  short  period  they  are  very  fre- 
quently visited,  especially  by  butterflies  and  bees.  The  pollination  has  been 
described  on  page  567.  Protandry  is  universal.  In  the  bud  the  tips  of  the 
styles,  covered  by  the  sweeping-hairs,  lie  closely  enveloped  by  the  anther-tube ; 
in  the  next  stage  the  style  grows  through  the  tube  and  sweeps  out  the  pollen  as 
it  proceeds ;  ultimately  the  stylar  branches  expand  and  the  stigma  is  then  pre- 
pared to  receive  the  pollen.  In  many,  the  sensitiveness  of  the  filaments  assists 
in  sweeping  out  the  pollen  at  the  exact  moment  of  the  insect  visit.  Regular 
self-pollination  is  found  e.g.  in  Senecio  vulgaris  j  wind-pollination  e.g.  in 
Artemisia  and  the  plants  related  to  it. 

This  extremely  natural  and  well-defined  order  is  the  largest  (and  no  doubt 
one  of  the  youngest?) ;  it  embraces  10-12,000  known  species  (in  770  genera),  or 
about  one-tenth  of  all  Flowering-plants.  They  are  distributed  over  the  whole 
globe,  but  are  most  numerous  in  temperate  countries ;  the  majority  prefer  open 


574  D1COTYLEDONES. 

spaces ;  a  smaller  number  are  forest-forms.     They  abound  especially  in  open 
districts  in  America. 

Among  the  substances  frequently  found  may  be  mentioned :  INULIN  (especially 
in  the  subterranean  parts),  BITTER  materials,  Tannin,  volatile  .oils,  fatty  oils  in 
the  fruits.  MEDICINAL  :  *  "  Herba  "  of  Artemisia  absinthium  (Wormwood)  and 
maritima]  (Sea-wormwood),  A  chilleamille folium;  the  leaves  of  Cnicus  benedictus 
and  Tussilago  farfara ;  the  unopened  capitula  of  Artemisia  maritima,  var.  stech- 
manniana;  the  capitula  of  Tanacetum,Matricariachamomilla^  (wildChamomile), 
Anthemis  nobilis  f  (common  Chamomile) ;  the  separate  flowers  of  Arnica  •  the 
roots  of  Arnica  montana^,  Taraxacum  ojficinale^,  Anacyclus  officinarum^,  Lappa 
major,  minor,  nemorosa  and  tomentosa,  Inula  helenium  and  Artemisia  vulgar  is  -T 
the  latex  of  Lactuca  virosaf.  The  following  are  cultivated  for  food : — Lactuca 
sativa  (Lettuce),  Cichorium  endivia  (from  E.  Asia,  for  salads),  Cynara  scolymus 
(Artichoke,  Mediterranean),  Scorzonera  hispanica  (S.  Eur.),  Helianthus  tuberosus 
(Jerusalem  Artichoke,  from  N.  Am.,  introduced  into  Europe  1616),  Cichorium 
intybus  (roots  as  "  chicory,")  Tragopoyott  ponifolium  (Salsafy),  Artemisia  dra- 
cunculus.  OIL  is  extracted  from  the  following  (the  seeds) :  Helianthus  annuus 
(Peru),  Madia  sativa  (Chili),  Guizotia  oleifera  (Abyssinia).  DYES  from  :  Car- 
thamu*  tinctorim  (Safflower,  used  in  the  preparation  of  rouge;  Egypt),  Serratula 
tinctoria.  INSECT-POWDER  from:  Pyrethrum  cinerariifolium  (Dalmatia)  and 
roseum  (Persia,  Caucasus).  The  following  are  cultivated  in  houses  and  gardens 
for  the  sake  of  their  scented  leaves  :  —  Tanacetum  balsamita  (Balsam),  Arte- 
muia  abrotanum  (Southernwood)  and  A.  argentea.  A  great  many  of  the  genera 
enumerated  are  cultivated  in  dwelling  houses  for  the  sake  of  the  flowers  ; 
e.g.  Pericallis  cruenta  (generally  termed  "  Cineraria  ").  Asteriscus  pygmceus  is 
supposed  to  be  the  genuine  "  Hose  of  Jericho  "  ;  the  involucral  leaves  envelop 
the  fruits  after  their  ripening  and  keep  them  enclosed  for  8-10  months  until 
rain  occurs. 

*  Those  marked  with  a  +  are  officinal. 


APPENDIX   ON   THE    CLASSIFICATION   OF 
PLANTS. 

BY  M.  0.  POTTER. 

THE  earliest  systems  of  classification  were  derived  from  the 
properties  and  uses  of  plants ;  and  it  was  not  until  some  two 
centuries  ago  that  any  scientific  grouping  of  plants  was  attempted. 
Aristotle  and  Theophrastus  had  adopted  the  groups  of  Trees, 
Shrubs  and  Herbs  as  the  chief  divisions  of  the  Vegetable  King- 
dom, a  system  which  persisted  and  was  employed  by  Tournefort 
and  Ray  as  late  as  the  end  of  the  17th  century.  The  arrange- 
ment by  which  these  three  divisions  were  separated  into  smaller 
divisions  was  often  founded  upon  a  single  character,  such  as  the 
formation  of  the  corolla,  the  form  of  fruit,  that  of  the  calyx  and 
corolla,  etc.  All  these  systems  of  classification  which  brought 
into  close  proximity  plants  distinguished  by  some  one  character 
alone,  could  only  be  considered  as  artificial,  since  plants  related  to 
one  another  would  not  necessarily  be  included  in  the  same  group. 
As  the  knowledge  of  the  morphology,  physiology,  and  reproduction 
of  plants  increased,  such  systems  were  recognised  as  unscientific, 
and  it  became  the  aim  of  botanists  to  establish  a  natural  system, 
founded  upon  mutual  relationships,  which  would  associate  together 
only  those  plants  which  are  truly  allied. 

The  following  are  some  of  the  chief  systems  of  classification 
which  will  show  the  gradual  development  of  the  natural  system, 
and  may  be  of  service  to  students  making  use  of  this  text-book.1 

System  of  JOHN  RAY  (1703). 
I.  Herbse. 

A.  IMPERFECTS  (Flowerless) 

B.  PERFECTS  (Flowering). 

Dicotyledones. 
Monocotyledones. 
II.  Arbores. 

A.  Monocotyledones. 

B.  Dicotyledones. 

Ray  was  the  first  botanist  who  recognised  the  importance  of  the 
one  or  two  seed-leaves  of  the  embryo,  and  initiated  the  division  of 
the  Flowering-plants  into  Monocotyledons  and  Dicotyledons. 

1  For  further  reference  see  Sachs,  History  of  Botany;  Lindley,  Vegetable 
Kinydom;  Le  Maout  and  Decaisne,  General  System  of  Botany,  etc. 

576 
W.  B.  P  P 


576 


APPENPIX   ON   THE    CLASSIFICATION    OF   PLANTS. 


System  of  LINN^US  (1733). 

In  his  well  known  artificial  system  Linnaeus  divided  the  Vege- 
table Kingdom  into  twenty-four  classes,  based  upon  the  number, 
relative  position  and  union  of  the  stamens  with  regard  to  each 
other,  and  also  to  the  gynoeceum. 

Class 


I. 

MONANDRIA. 

Flowers  with 

1  stamen. 

II. 

DlANDRIA. 

»             i> 

2  stamens. 

III. 

TRIANDRIA. 

n             ii 

3 

IV. 

TETRANDRIA. 

,,             Jt 

4 

V. 

PENTANDRIA. 

rt                  >» 

5         „ 

VI. 

HEXANDRIA. 

ii                  ii 

6        „ 

VII. 

HEPTANDRIA. 

»                  »» 

7 

VIII. 

OCTANDRIA. 

»                  » 

8 

IX. 

ENNEANDRIA. 

ii                  it 

9        ,, 

X. 

DECANDRIA. 

i*                  i) 

10 

XI. 

DODECANDRIA. 

ii                  ii 

11  to  19  star 

XII. 

ICOSANDRIA. 

it                  ii 

20  or  more 

the  calyx. 
XIII.    POLYANDRIA.       ,,  ,,20  or  more  stamens  inserted  on 

the  receptacle. 

DIDYNAMIA.        Stamens  didynamous. 
TETRADYNAMIA.        „          tetradynamous. 
MONADELPHIA.     Filaments  united  into  1  bundle 
DIADELPHIA.  „  „  ,,     2  bundles. 

POLYADELPHIA.  „     several  bundles. 


XIV. 
XV. 

XVI. 

XVII. 
XVIII. 

XIX. 
XX. 

XXI. 

XXII. 
XXIII. 
XXIV. 


SYNGENESIA.  Anthers  united  together: 

GYNANKRIA.  Stamens  and  pistil  united. 

MONCECIA.  Flowers  diclinous,  $  and  $  on  the  same  plant. 

DICECIA.  „            ,,          $  and  $  on  different  plants. 

POLYGAMIA.  $  -,  ?  -,  and  §  -flowers  on  the  same  plant. 

CRYPTOGAMIA.  Flowerless  plants  (Ferns,  Mosses,  Algae,  Fungi). 

These  classes  were  further  divided  into  orders,  according  to  the  number  of 
styles,  as  Monogynia,  flowers  with  1  style;  Digynia,  with  2  styles,  etc.  Thus 
a  Dock  (Rumex),  having  6  stamens  and  3  styles,  would  be  placed  in  Class  VI., 
HEXANDRIA,  and  Order  III.,  Trigynia. 

Class  XIV.  was  divided  into  two  orders.  Order  I.,  Gymnospermia,  with  seeds 
apparently  naked,  comprising  the  Labiatse  ;  and  Order  II.,  Angiospermia,  with 
the  seeds  enclosed  in  a  capsule  (Bartsia,  Rhinanthus). 

Class  XV.  was  divided  into  two  orders :  Order  I.,  Siliculosa,  fruit  a  silicula 
{Capsella} ;  and  Order  II.,  Siliquosa,  fruit  a  siliqua  (Brassica). 

Class  XIX.  was  divided  into  Order  I.,  .ZEqualis,  all  the  flowers  perfect 
(Sonchus) ;  Order  II.,  Superflua,  flowers  in  the  centre  perfect,  those  at  the 
circumference  with  pistils  only  (seemingly  superfluous),  e.g.  Aster  ;  Order  III., 
Frustranea,  flowers  in  the  centre  perfect,  those  at  the  circumference  neuter, 
e.g.  Centaurea. 

"Fragments"  of  a  natural  system  have  also  come  down  to  us  from  Linnaeus, 
who  himself  always  recognised  the  imperfection  of  his  artificial  system. 


APPENDIX   ON    THE    CLASSIFICATION   OF   PLANTS.  577 

System  of  ANTOINE  LAURENT  DE  JUSSIEU  (1789). 

Acotyledones.     Plants  without  cotyledons :  Fungi,  Ferns,  Mosses, 

Algas,  Naiades         ......  I. 

Monoctyledones.     Plants  with  one  cotyledon  : — 

1.  Stamens  hypogynous II. 

2.  „         perigynous            . HI. 

3.  „         epigynous             IV. 

Dicotyledones.     Plants  with  two  cotyledons : — 

f  Stamens  epigynous             .....  V. 

1.  APETAL^:                 ..,         perigynous VI. 

^       „        hypogynous          .                 ...  VII. 


2.  MONOPETAL.E 


3.    POLYPETAL^E 


Corolla  hypogynous VIII. 

„      perigynotfs IX. 

anthers  connate           .        .  X. 

free  XL 


epigynous,  j ' 


Stamens  epigynous  XII. 

,,         hypogynous          .         .         .         .         .     XIII. 
„        perigynous  .         .         .        .         .     XIV. 


4.  DICLINES  IRTCEGULARES,  male  and  female  flowers  on  different  plants, 
corolla  generally  absent. 


System  of  A.  P.  DE  CANDOLLK  (1819). 

I.  Vasculares.     Plants  with  vascular  bundles. 

1.  EXOGEN^E.     Vascular  bundles  arranged  in  a  ring. 

A.  Diplochlamydece.     Calyx  and  corolla  present. 

a.  Thalamiflorje.     Corolla  polypetalous  and  hypogynous. 
6.  Calycifloras.     Corolla  perigynous  or  epigynous  ;  stamens 

inserted  on  the  calyx. 
c.  Corolliflorffi.      Corolla  gamopetalous ;  stamens  inserted 

on  the  corolla. 

B.  Monochlamydea.     Perianth  simple. 

2.  ENDOGEN.E.     Vascular  bundles    scattered,  the  youngest  in  the 

centre. 

A.  Phanerogams.     Flowers  present. 

B.  Cryptogamce.     Flowers  absent. 

II.  Cellulares.     Vascular  bundles  absent. 

1.  FOLIACE^E.    Leaves  present. 

2.  APHYLL^E.     Leafless. 


578  APPENDIX   ON   THE   CLASSIFICATION    OF   PLANTS. 

KOBERT  BROWN  published  in  1827  his  discovery  of  the  gymno- 
spermy  of  the  ovules  of  the  Coniferse  and  Cycadece,  and  showed 
that  the  Gyrnnosperms,  which  had  previously  been  classed  with 
the  Dicotyledons,  must  be  regarded  as  an  independent  group. 


System  of  STEPHEN  ENDLICHER  (1836-40). 

I.  Thallophyta.     No  differentiation  into  stem  and  root. 

1.  PKOTOPHYTA.     Class  I.,  Algae ;  Class  II.,  Lichenes. 

2.  HYSTEROPHYTA.     Class  III.,  Fungi. 

II.  Cormophyta.     Differentiated  into  stem  and  root. 

1.  ACROBBYA.     Stem  growing  at  the  point. 

Anophyta  (Hepaticae,  Musci). 
Protophyta  (Filices,  etc.). 
Hysterophyta  (BalanophoreeB,  etc.). 

2.  AMPHIBRYA.      Stem  growing  at  the  circumference  (Monocotyle- 

dons). 

3.  ACBAMPHIBRYA.      Stem  growing  both  at  the  point  and  circum- 

ference. 

Gymnosperma  (Conifers). 
Apetula.     Perianth  single  or  absent. 
Gamopetala.     Petals  gamopetalous. 
Dialypetala.     Petals  polypetalous. 


System  of  A.  BRONGNIART  (1843). 

I.  Cryptogamae.     Plants  without  flowers. 

1.  AMPHIGEN.K.    Not  differentiated  into  stem  or  leaf  (Algse,  Fungi, 

Lichenes). 

2.  ACBOGEN^:.    Plants  with  stem  and  leaf  (Muscineas,  Jb  ilium®). 
II.  Phanerogamae.     Plants  with  flowers. 

3.  MONOCOTYLEDONES. 

a.  Albuminosaa.     Seeds  with  endosperm. 

b.  Exalbuminosse.     Seeds  without  endosperm. 

4.  DlCOTYLEDONES. 

a.  Angioppermae. 

a.  Gamopetalffl. 

0.  Dialypetalse. 
6.  Gymnospermse. 


APPENDIX   ON    THE    CLASSIFICATION   OF    PLANTS.  579 

System  of  JOHN  LINDLEY  (Vegetable  Kingdom,  1845). 
Asexual,  or  Flowerless  Plants. 

Stem  and  leaves  undistinguishable  ....      I.  Thallogens. 

Stem  and  leaves  distinguishable II.  Acrogens. 

Sexual,  or  Flowering  Plants. 

Fructification  springing  from  a  thallus    ....  III.  Rhizogens. 
Fructification  springing  from  a  stem. 

Wood  of  stem  youngest  in  the  centre ;   cotyledon 
single.  Leaves  parallel- veined,  permanent;  wood 

of  stem  always  confused IV.  Endogens. 

Leaves  net-veined,  deciduous ;   wood   of  the   stem, 
when   perennial,   arranged  in   a   circle  with  a 

central  pith V.   Dictyogens. 

Wood  of  stem  youngest  at  the  circumference,  always 
concentric  ;  cotyledons,  2  or  more. 

Seeds  quite  naked VI.   Gymnogens. 

Seeds  inclosed  in  seed-vessels VII.  Exogens. 


System  of  ALEXANDER  BRAUN  (1864). 

I.  Bryophyta. 

1.  THALLODEA  (Algae,  Fungi,  Lichenes). 

2.  THALLOPBYLLODEA  (Charas,  Mosses). 
II.  Cormophyta. 

1.  PHYLLOPTERIDES  (Ferns,  Equisetums). 

2.  MASCHALOPTEKIDES  (Lycopods). 

3.  HYDROPTERIDES  (Water-ferns), 
III.  Anthophyta. 

GYHNOSPERM.E. 

1.  Frondoscc  (Cycadeae). 

2.  Acerosce  (Coniferae). 
ANGIOSPEKMJE. 

1.  Monocotyledones. 

2.  bicotyledones. 

Apetalae. 

Sympetalae. 

Eleutheropetal®. 


W.  HOFMEISTER  published  from  1849  to  1851  his  researches  upon 
the  embryology  of  the  Phanerogams,  and  upon  the  embryology 
and  life-history  of  the  Vascular  Cryptogams,  and  established  the 
phylogenetic  connection  existing  between  the  Mosses,  Vascular 
Cryptogams  and  Phanerogams. 


580  APPENDIX   ON   THE    CLASSIFICATION    OF    PLANTS. 


System  of  HOOKER  and  BENTHAM  (Genera  plantarum,  1862-1883). 

DICOTYLEDONES. 

I.  POLYPETAL^E. 

Series  I.  Thalamiflorae.  Calyx  most  often  free  from  the  ovary.  Petals 
uniseriate  or  often  2-Oo-seriate.  Stamens  oo  or  definite,  inserted  on  the  recep- 
tacle, often  small,  or  raised,  or  stipitate.  Ovary  most  frequently  free. 

Cohort  I.  BANALES.  Stamens  oo,  or  if  definite  the  perianth  is  3-  oo-seriate. 
Carpels  apocarpous,  or  immersed  in  the  receptacle.  Endosperm  usually  abun- 
dant, fleshv. 


Order  1.  Banunculaceae. 
,,      2.  Dilleniaceaa. 
„      3.  Calycanthacese. 
,,      4.  Magnoliaceee. 


Order  5.  Anonaceae. 
,,      6.  Menispermaceffl. 
„      7.  Berberideas. 
,,      8.  Nymphaeacese. 


Cohort  II.  PAKIETALES.  Stamens  oo  or  definite.  Ovary  unilocular,  or  divided 
into  loculi  by  spurious  dissepiments,  with  parietal  placentation.  Endosperm 
absent  or  fleshy. 


Order  9.  Sarraceniaceaa. 
,,    10.  Papaveraceae. 
„    11.  Cruciferae. 
,,    12.  Capparidero. 
,    13.  Besedaceae. 


Order  14.  Cistineae. 
,,      15.  Violarieae. 
„      16.  Canellaceaa. 
,,      17.  Bixineae. 


Cohort  III.  POLYGALIN^;.  Stamens  definite.  Ovary  usually  perfectly  or  im- 
perfectly bilocular.  Micropyle  often  superior.  Fruit  very  often  compressed 
laterally.  Endosperm  very  often  abundant  and  fleshy. 

Order  18.  Pittosporeae.  I     Order  20.    Polygaleae. 

,,      19.  Tremandreae.  |        ,,      20a.  Vochysiaceae. 

Cohort  IV.  CAKYOPHYLLINE.E.  Stamens  definite,  or  rarely  oo.  Ovary  unilo- 
cular, or  imperfectly  septate.  Placenta  central,  more  rarely  parietal.  Micropyle 
inferior.  Embryo  curved,  rarely  straight.  Endosperm  farinaceous. 

Order  21.  Frankeniacese.  I     Order  23.  Portulaceae. 

„      22.  Caryopbyllese.  |        ,,      24.  Tamariscinese. 

Cohort  V.  GUTTIFE RALES.  Sepals  inbricate.  Stamens  usually  oc.  Ovary 
septate,  placentas  on  the  inner  angles  of  the  loculi.  Endosperm  absent  or 
fleshy. 


Order  25.  Elatineaa. 
,,      21).  HypericincBa. 
27.  Guttiferse. 


Order  28.  TernstroemiacezB. 

,,      29.  Diptcrocarpeee. 

30.  Chlaanaceffi. 


APPENDIX   ON   THE    CLASSIFICATION   OF   PLANTS.  581 

Cohort  VI.  MALVALES.  Sepals  valvate.  Stamens  usually  GO  or  mona- 
delphous.  Ovary  septate,  placentae  on  the  inner  angles  of  the  loculi.  Endo- 
sperm absent  or  fleshy. 

Order  31.  Malvaceae.  |    Order  33.  Tiliacese. 

%,,      32.  Sterculiaceaa. 

Series  II.  Disciflorae.  Calyx  usually  free  from  the  ovary.  Petals  uniseriate. 
Stamens  usually  definite,  inserted  within,  or  upon,  or  around  the  receptacle, 
which  is  more  often  expanded  as  a  disc.  Ovary  usually  free,  or  embedded  in 
the  disc. 

COHORT  VII.  GEKANIALES.  Disc  usually  as  a  ring  between  the  stamens,  or 
adnate  to  the  staminal  tube,  or  reduced  to  glands  alternating  with  the  petals, 
more  rarely  absent.  Gynoaceum  entire,  or  more  often  lobed,  or  sub- apocarpous. 
Ovules  most  often  1-2  in  each  loculus,  pendulous,  raphe  ventral.  Leaves 
various. 


Order  34.  Lineaa. 


35.  Humiriaceae. 

36.  Malpighiaceaa. 

37.  Zygophylleaa. 

38.  Geraniaceae. 

39.  Kutaceae. 


Order  40.  Simarubeae. 

„      41.  Ochnaceae. 

„      42.  Burseraceffi. 

„      43.  Meliaceas. 

44.  Chailletiaceoa. 


Cohort  VIII.  OLACALES.  Disc  cupular  or  annular,  free,  or  bearing  the  stamens 
and  petals  on  its  edge.  Gynoeceum  entire.  Ovules  1-3  in  the  unilocular 
ovaries,  or  1-2  in  each  loculus,  pendulous,  raphe  dorsal.  Leaves  simple. 

Order  45.  Olacinese.  |     Order  46.  Ilicineae. 

Cohort  IX.  CELAST RALES.  Disc  tumid,  adnate  to  the  calyx,  or  covering  its 
base.  Stamens  inserted  round  the  disc  or  affixed  to  its  margin.  Gynoeceum 
usually  entire.  Ovules  most  often  two  in  each  loculus,  erect,  raphe  ventral. 
Leaves  simple,  or  rarely  compound. 

Order  47.  Celastrineas.  I     Order  49.  Rhamneae. 

„      48.  Stackhousiea).  „      50.  Ampelideae. 

Cohort  X.  SAPINDALES.  Disc  various.  Stamens  variously  inserted  on  the 
disc.  Gynoeceum  entire,  or  more  often  lobed,  or  sub-apocarpous.  Ovules  more 
often  1-2  in  each  loculus,  ascending  with  ventral  raphe,  or  reversed,  or  solitary 
and  pendulous  from  an  ascending  funicle,  or  rarely  oo  horizontal.  Leaves 
pinnate,  or  more  rarely  simple  or  digitate. 

Order  51.  Sapindaceas.  I    Order  53.  Anacardiaceae. 

„      52.  Sabiaceae. 

Anomalous  orders,  or  rather  genera, — 

Order  54.  Coriarieae.  |     Order  55.  Moringeae. 

Series  III.  Calyciflorae.  Calyx-tube  usually  surrounding  the  ovary,  or 
adnate  to  it.  Petals  uniseriate,  inserted  on  the  calyx-tube.  Stamens  oo  or 
definite,  inserted  on  the  calyx-tube,  or  most  often  on  the  disc  lining  the  calyx- 
tube.  Ovary  often  enclosed  by  the  calyx-tube,  or  inferior. 


582  APPENDIX   ON   THE    CLASSIFICATION   OF   PLANTS. 

Cohort  XI.  ROSALES.  Carpels  solitary,  or  free,  or  united  at  the  base,  more 
rarely  at  the  apex;  styles  distinct,  or  very  rarely  united  into  a  column,  and 
easily  separated. 


Order  56.  Connaraceaa. 
„      57.  Leguminosae. 
„      58.  Kosaceae. 
„      59.  Saxifrageae. 
60.  Crassulaceas. 


Order  61.  Droseraceaa. 
,,      62.  Hamamelideae. 
,,      63.  Bruniaceae. 
„      64.  Halorageaa. 


Cohort  XII.  MYRTALES.  Ovary  syncarpous,  inferior,  or  enclosed  in  the 
calyx-tube,  usually  divided  into  loculi ;  style  undivided.  Ovules  2-ao  in  the 
loculi. 


Order  65.  Bhizophorese. 
,,  66.  Combretaceae. 
,,  67.  Myrtaceae. 


Order  68.  Melastomaceae. 
,,      69.  Lythrarieaa. 
,,      70.  Onagrarieae. 


Cohort  XIII.  PASSIFLORALES.  Ovary  syncarpous,. inferior  or  superior,  en- 
closed in  the  calyx-tube  or  exserted,  unilocular  with  parietal  placentation,  or 
divided  into  loculi ;  styles  distinct,  one  style  divided,  or  undivided. 


Order   71.  Samydaceit. 
,,      72.  Loaseae. 
„      73.  Turneraceae. 
74.  Passifloreae. 


Order  75.  Cucurbitaceaa. 
„      76.  Begoniaceae. 
,,      77.  Datisceaa. 


Cohort  XIV.  FICOIDALES.  Ovary  syncarpous,  inferior  or  superior,  divided 
into  loculi  with  sub-basilar  placentae,  or  more  rarely  unilocular  with  parietal 
placentae.  Styles  distinct,  or  divided  at  the  apex.  Embryo  curved  or  excentric. 

Order  78.  Cacteae.  |        Order  79.  Ficoideae. 

Cohort  XV.  UMBELLALES.  Ovary  syncarpous,  inferior,  crowned  by  the  disc, 
divided  into  loculi,  or  unicarpellate.  Styles  distinct  or  divided  at  the  apex. 
Ovules  solitary  and  pendulous  in  the  loculi. 

Order  80.  Umbelliferaa.  |         Order  82.  Cornaceae. 

,,      81.  Araliaceae. 


II.     GAMOPETAL^E. 

Series  I.  Inferse.  Ovary  inferior.  Stamens  equal  to  the  lobes  of  the 
corolla,  rarely  fewer. 

Cohort  I.  RCBIALES.  Stamens  adnate  to  the  corolla.  Ovary  2-oc-locular, 
loculi  1-oc-ovuled. 

Order  83.  Caprifoliaceaa.  |        Order  84.  Rubiaceae. 

Cohort  II.  ASTEBALES.  Stamens  adnate  to  the  corolla.  Ovary  formed  of 
2  carpels,  unilocular  and  1-ovuled. 

Order  85.  Valerianeae.  I        Order  87.  Calycereae. 

,,      86.  Dipsaceae.  „      88.  Composite. 


APPENDIX    ON    THE    CLASSIFICATION    OF    PLANTS.  583 

Cohort  III.  CAMPANALES.  Stamens  generally  free  from  the  corolla.  Ovary 
2-6-locular,  loculi  most  o*ften  oo-ovuled. 

Order  89.  Stylidiese.  |       Order  91.  Campanulaceffi. 

„      90.  Goodenovieae. 

Series  II.  Heteromerae.  Ovary  most  often  superior.  Stamens  free  from 
the  corolla,  opposite  to,  or  double  the  lobes  of  the  corolla,  or  oo,  or  if  epipeta- 
lous.  equal  and  alternating  with  them.  Carpels  more  than  2. 

Cohort  IV.     ERICALES.    Stamens  double  the  lobes  of  the  corolla,  or  alternating 
with  them.     Ovary  2-oc-locular.     Seeds  small,  frequently  minute. 
Order  92.  Vacciniaceae.  Order  95.  Epacrideae. 


93.  Ericaceae. 

94.  Monotropeae. 


96.  Diapensiaceae. 

97.  Lennoaceae. 


Cohort  V.  PRIMULALES.  Stamens  equal  to  and  opposite  the  lobes  of  the 
corolla.  Ovary  unilocular,  with  a  free  central  placenta,  1-^  ovules. 

Order  98.  Plumbagineae.  I         Order  100.  Myrsineae. 

,,      99.  Primulaceje. 

Cohort  VI.  EBENALES.  Stamens  equal  to  and  opposite  the  lobes  of  the 
corolla,  or  double,  or  oo.  Ovary  2-oo-locular.  Seeds  few  and  large.  Trees  or 
shrubs. 

Order  101.  Sapotaceae.  |        Order  103.  Styraceae. 

„      102.  Ebenaceae. 

Series  III.  Bicarpellatae.  Ovary  most  often  superior.  Stamens  equal,  or 
fewer  than  the  lobes  of  the  corolla,  and  alternating  with  them.  Carpels  2, 
rarely  1  or  3. 

Cohort  VII.  GENTIANALES.  Corolla  regular.  Stamens  equal  to  the  lobes  of 
the  corolla,  or  if  fewer,  usually  alternating  with  the  carpels.  Leaves  generally 
opposite. 


Order  104.  Oleaceae. 
„      105.  Salvadoracese. 
,,      106.  Apocynaceae. 


Order  107.  Asclepiadeas. 
„      108.  Loganiaceae. 
109.  Gentianeae. 


Cohort  VIII.     POLEMONIALES.     Corolla  regular.     Stamens  equal  to  the  lobes 
of  the  corolla.     Leaves  generally  alternate. 


Order  110.  Polemoniaceaa. 
„      111.  Hydrophylhiceae. 


Order  113.  Convolvulacese. 
114.  Solanaceae. 


„      112.  Boragineae. 

Cohort  IX.  PERSONALES.  Corolla  most  often  irregular  or  oblique.  Posterior 
stamen  less  than  the  others,  more  often  reduced  to  a  staminode,  or  altogether 
absent.  Ovary  oo-ovuled,  or  2-ovuled. 


Order  115.  Scrophularineae. 
„      116.  Orobanchaceae. 
„      117.  Lentibularieae. 
118.  Columelliaceae. 


Order  119.  Gesner  iceae. 
„      120.  Bif>noniaceae. 
„      121    Pedalineaa. 
122.  Acanthaceae. 


584  APPENDIX    ON    THE    CLASSIFICATION    OF    PLANTS. 

Cohort  X.  LAMIALES.  Corolla  most  often  irregular  or  oblique.  Posterior 
stamen  less  than  the  others,  most  frequently  reduced  4o  a  staminode  or  absent. 
Carpels  1-ovuled  or  with  2  collateral  ovules.  Fruit  enclosed  in  the  persistent 
calyx,  indehiscent,  and  with  one  seed,  or  dehiscing  into  2  or  4,  rarely  oo, 
1-seeded  nuts. 

Order  123.  Myoporineaa.  I         Order  125.  Verbenaceaa. 

„      124.  Selagineaa.  „      120.  Labiateaa. 

Anomalous  Order  127.    Plantagineae. 

III.     MONOCHLAMYDE-flB. 

Perianth  simple,  lobes  or  segments  1-2-seriate  and  often  sepaloid,  or  small, 
or  wanting. 

Series  I.  Curvembryeae.  Endosperm  frequently  farinaceous.  Embryo 
curved,  excentric,  lateral  or  peripheral,  rarely  straight.  Ovules  most  frequently 
1  in  the  ovary,  or  1  in  each  loculus.  Flowers  $  ,  in  some  genera  unisexual  or 
polygamous.  Petals  very  rare.  Stamens  equal  to  the  segments  of  the  peri- 
anth, rarely  fewer  or  more. 


Order  128.  Nyctagineaa. 
,,      129.  Illecebraceaa. 
130.  Amarantaceaa. 


Order  132.  Phytolaccaceaa. 
„      133.  Batideae. 
„      134.  Polygonaceaa. 


„      131.  Chenopodiaceaa. 

Series  II.  Multiovulatse  Aquaticse.  Aquatic  herbs,  submerged.  Ovary 
syncarpous  ;  ovules  numerous  in  each  loculus  or  on  each  placenta. 

Order  135.  Podostemaceae. 

Series  III.  Multiovulatse  Terrestres.  Terrestrial  trees  or  shrubs.  Ovary 
syncarpous  ;  ovules  numerous  in  each  loculus  or  on  each  placenta. 

Order  136.  Nepenthaceaa.  I         Order  138.  Aristolochiaceae. 

„      137.  Cytinaceae. 

Series  IV.  Micrembryeae.  Ovary  syncarpous,  monocarpous,  or  apocarpous. 
Ovules  generally  solitary  in  each  carpel,  rarely  2  or  few.  Endosperm  copious, 
fleshy,  or  rarely  farinaceous.  Embryo  very  minute. 


Order  139.  Piperaceae. 

140.  Chlorantbaceae. 


Order  141.  Myristiceae. 
142.  Monimiaceaa. 


Series  V.  Daphnales.  Ovary  monocarpous,  very  rarely  syncarpous,  with 
2-4  loculi ;  ovules  in  the  ovary  or  in  each  loculus,  solitary,  or  in  pairs.  Trees 
or  shrubs,  very  rarely  herbs  ;  flowers  generally  £  .  Perianth  perfect,  sepaloid, 
1-2  seriate.  Stamens  perigynous,  equal  to  the  lobes  of  the  perianth,  or  double 
unless  fewer. 


Order  143.  Laurineae. 
144.  Proteaceaa. 


Order  146.  Penaaceaa. 
,,     147.  Elaaaguaceaa. 


,,     145.  Thymelaaaceae. 

Series  VI.     Achlamydosporeae.      Ovary  unilocular,   1-3   ovu'es.     Ovules 
most    frequently  poorly   developed  before  flowering.       Seeds   endospermous, 


APPENDIX   ON   THE    CLASSIFICATION   OF   PLANTS. 


585 


but  without  testa,  either  free  in  the  pericarp  or  attached  to  its  walls, 
generally  perfect,  sepaloid  or  petaloid. 


Perianth 


Order  148.  Loranthaceae. 
149.  Santalacese. 


Order  150.  Balanophorese. 


Series  VII.  Unisexuales.  Flowers  unisexual.  Ovary  syncarpous  or  mono- 
carpous,  ovules  in  the  ovary  or  in  each  loculus,  solitary,  or  in  pairs.  Endosperm 
copious,  fleshy,  or  scanty,  or  absent.  Trees  or  shrubs,  rarely  herbs.  Stipules 
generally  present.  Perianth  sepaloid,  or  minute,  or  absent.  Styles  equal  in 
number  to  the  carpels,  not  rarely  bifid. 


Order  151.  Euphorbiaceao. 

,,     152.  Balanopseaa. 

„     153.  Urticaceae. 

„     154.  Platanaceae. 

,     155.  Leitnerieae. 


Order  156.  Juglandese. 
„     157.  Myricaceae. 
„     158.  Casuarineas. 
„     159.  Cupuliferaa. 


Series  VIII.     Ordines  Anomali.     Anomalous  Orders. 


Order  160.  Salicinese. 

161.  Lacistemacese. 


Order  162.  Empetraceae. 
,,     163.  Ceratophylleae. 


Order  164.  Gnetaceae. 
165.  Coniferae. 


GYMNOSPERME^.. 

I  Order  166.  Cycadaceffi. 


MONOCOTYLEDONES. 

Series  I.  Microspermae.  At  least  the  inner  series  of  the  perianth  petaloid. 
Ovary  inferior,  unilocular,  with  3  parietal  placentas,  or  rarely  3-locular,  with 
axile  placentation.  Seeds  minute,  numerous,  without  endosperm. 


Order  167.  Hydrocharideae. 
,     168.  Burmanniaceae. 


Order  169.  Orchideaa. 


Series  II.     Epigynae.     At  least  the  inner  series  of  the  perianth  petaloid. 
Ovary  most  often  inferior.     Endosperm  copious. 

Order  174.  Amaryllideae. 
„     175.  Taccaceae. 
,     176.  Dioscoreaceaa. 


Order  170.  Scitamineae. 
,,     171.  Bromeliaceae. 
„     172.  Hasrnodoraceae. 
173.  Irideae. 


Series  III.     Coronarieae.     At  least  the  inner   series  of  perianth  petaloid. 
Ovary  free,  very  rarely  slightly  adnate  at  the  base.     Endosperm  copious. 


Order  177.  Roxburghiaceae. 
,,     178.  Liliace*. 
„     179.  Pontederiaceae. 
„     180.  Philydraceae. 


Order  181.  Xyrideae. 
„     182.  Mayacesa. 
„     183.  Commelinaceaa. 
„     184.  Eapateaceaa. 


586 


APPENDIX   ON   THE    CLASSIFICATION   OF    PLANTS. 


Series  IV.    Calycinae.    Perianth  sepaloid,  small,  rigid,  or  herbaceous  (inner 
series  subpetaloid  or  small).     Ovary  free.     Endosperm  copious. 


Order  185.  Flagellariea?. 
186.  Juncacese. 


Order  187.  Palmse. 


Series  V.  Nudiflorae.  Perianth  absent,  or  reduced  to  hairs  or  scales. 
Ovary  superior,  carpel  solitary,  or  if  many,  syncarpous,  1-oo-ovuled.  Endosperm 
most  frequently  present. 


Order  188.  Pandaneaa. 
„     189.  Cyclanthaceffi. 
„     190.  Typhacese. 


Order  191.  Aroideae. 
192.  Lemnaceae. 


Series  VI.  Apocarpse.  Perianth  1-2-seriate,  or  absent.  Carpels  superior, 
solitary,  or  if  more,  apocarpous.  Endosperm  absent. 

Order  193.  Triuridese.  Order  195.  Naiadaceae. 

,,     194.  Alismaceae. 

Series  VII.  Glumacese.  Flowers  solitary,  sessile  in  the  axils  of  bracts  and 
arranged  in  capitula  or  spikelets  with  bracts.  Segments  of  perianth  small, 
scale-like,  glumaceous  or  absent.  Ovary  1-ovuled,  or  divided  into  1-ovuled 
loculi.  Endosperm  present. 


Order  196.  Eriocauleaa. 
,,     197.  Centrolepideae. 
,,     198.  Bestiaceae. 


Order  199.  Cyperaceae. 
,     200.  Gramineffl. 


Classification  of  the  Thallophytes  proposed  by  SACHS  (Text-Book 
of  Botamj,  English  Edition,  1882). 

THALLOPHYTES. 

Containing  chlorophyll.  \      Not  containing  chlorophyll. 

Class  I.     Protophyta. 


Cyanopbyceae. 
Palmellaceae  (in  part). 


Schizomycetes. 
Saccharomycetcs. 

Class  II.     Zygosporeae. 
Conjugating  cells  motile. 

Myxomycetes. 


Pandorineaa. 
(HydrodicfcyeaB). 

Conjugating  cells  stationary. 
Conjugate  (including  Diatomacese).     |      Zygomycetes. 

Class  III.     Oosporeae. 
Spliaaroplea. 

Vaucheria    .     .     .     (Caeloblastce) 
Volvocineae. 
(Edogoniese. 
Fuco'deee. 


rSaprolegniesB. 
iPeronosporeae. 


APPENDIX   ON   THE    CLASSIFICATION   OF    PLANTS.  587 

Class  IV.     Carposporeae. 


Coleochaeteae. 

Florideae. 

Characete. 


Ascomycetes  (including  Lichens). 
.ZEcidiomycetes  (Urediuete). 
Basidiomycetes. 


System  of  A.  W.  EICHLEE  (1883). 
A.  Cryptogamae. 
I.  Thallophyta. 

1.  Class.     ALG^;. 

1  Group.   Cyanophycem. 

2  ,,        Diatomere. 

3  „        Chlorophycese. 

1  Series.   Conjugate. 

2  „       Zoosporeos. 

3  ,,       Characese. 

4  Group.   Pheeophycece. 

5  ,,       Rhodopliycera. 

2.  Class.     FONGI. 

1  Group.     Schizomycetes. 

2  ,,          Eumycetes. 

1  Series.    Phycomycetes. 


2       „ 
3       „ 

4      „ 
5       „ 
3  Group. 

II.  Bryophyta. 
1  Group. 

2      „ 

Ustilagineee. 
JEcidiomycetes. 
Ascomycetes. 
Basidiomjcetes. 
Lichenes. 

HepaticoB. 
Musci. 

III.  Pteridophyta. 

1  Class.    EQUISETINJE. 

2  ,,         LYCOPODIN^:. 

3  ,, 


B.  Phanerogamas. 

1.  Gyinnospermro. 

2.  Angiospermse. 

The  subdivisions  of  the  Phanerogams  have  with  little  variation  been  adopted 
in  this  book. 


588  APPENDIX   ON   THE    CLASSIFICATION   OF   PLANTS. 

Classification  of  the  THALLOPHYTES,  adopted  in  the  3rd  Danish 
Edition  (1891).  [Algse  by  Wille;  Fungi  by  Rostrup  (after 
Zcpf).} 

I.  DIVISION.     THALLOPHYTA. 

I.  Sub-division.     Algae. 

1  Class.     CHLOROPHYCE^E  (GREEN  ALG.E). 

1  Family.    Conjugates. 

2  , ,  Protoc,occoidea9. 

3  ,,  Confervoideee. 

4  ,,  Siphonese. 

5  „  Gyrophycese. 

2  Class.       PHjEOPHYCE^E  (BROWN  ALGJE). 

1  Family.     Syngeneticse. 

2  „  Dinoflagellata. 

3  „  Pyritophycese  (Diatomea). 

4  ,,  Phasosporeee. 

5  „  Cyclosporeae. 

6  ,,  Dictyoteffl. 

3  Class.     ACILIAT^E. 

A.  Sub-class.     Schizophycea. 

1  Family.     Myxophyce®  (Blue-Green  Algae). 

2  ,,         Bacteria. 

B.  Sub-class.     Rhodophycea. 

1  Family.     Bangioidese. 

2  ,,         Florideas. 

II.  Sub-division.     Myxomycetes. 

III.  „  Fungi. 

A.  Phycomycetes. 

1  ClaSS.       OOMYCETES. 

2  „          ZYGOMYCETES. 

B.  Mycomycetes. 

3  Class.     BASIDIOMYCETES. 

A.  Sub-class.     Protobasidiomycetes. 

B.  ,,  Autobasidiomycetes. 

1  Family.     Hymenomycetes. 

2  „         Gasteromycetes. 

3  „         Basidiolichenes. 

4  Class.    ASCOMYCETES. 

1  Family.     Gymnoasci. 

2  ,,         Perisporiese. 

3  ,,         Pyrenomycetes. 

4  ,,         Discomycetes. 

5  Ascolichenes. 


APPENDIX    ON   THE    CLASSIFICATION   OF   PLANTS.  589 

System  of  A.  ENGLER  (Syllabus  der  Vorlesungen,  etc.,  1892). 

I.  DIVISION.     MYXOTHALLOPHYTA. 

Snb-division.     Myxomycetes. 

1  Class.     ACRASIE^:. 

2  „         PLASMODIOPHOKALES. 

3  ,,          MYXOGASTERES. 

1  Series.     Ectospores. 

2  „         Endosporeae. 

II.  DIVISION.     EUTHALLOPHYTA. 

I.  Sub-division.     Schizophyta. 

1  ClaSS.       ScHIZOPHYCEjE. 

2  ,,  SCHIZOMYCETES. 

II.  Sub-division.     Dinoflagellata. 
Class.     DINOFLAGELLATA. 

1  Series.     Adinida. 

2  ,,         Dinifera. 

III.  Sub-division.     Bacillariales. 

Class.     BACILLARIALES. 

IV.  Sub-division.     Gamophyceae. 

1  Class.     CONJUGATE. 

2  ,,          CHLOROPHYCEJE. 

1  Sub-class.    Protococcales. 

2  „  Confervales. 

3  „  Siphonece. 

3  Class.     CHARALES. 

4  ,,         PH^EOPHYCE^E. 

1  Sub-class.    Phceosporece. 

2  ,,  Cyclusporea. 

5  Class.     DICTYOTALES. 

6  ,,        KHODOPHYCE.E. 

1  Sub-class.     Bangiales. 

2  ,,          Floridea.  .    . 

1  Series.     Nemalionales. 

2  ,,         Gigartinales. 

3  „  Ehod^meniales. 

4  ,,  Cryptonemiales. 
V.   Sub-division.  Fungi. 

1  Class.     PHYCOMYCETES. 

1  Series.     Zygomycetes. 

2  ,,          Oomycetes. 

1  Sub- series.     Chytridiales. 

2  ,,  Mycosiphonales. 

2  Class.     MESOMYCETES. 

1  Sub-class.     Hemiasci. 

2  Hemibasidii. 


590  APPENDIX   ON   THE    CLASSIFICATION   OF    PLANTS. 

3  Class.    MYCOMYCETES. 

1  Sub-class.     Ascomycetes. 

1  Series.     Exoasci. 

2  ,,         Carpoasci. 

1  Sub- series.     Gymnoascales. 

2  „  Perisporiales. 

3  „  Pyrenomycetes. 
Appended.     Pyrenolichenes. 

4  Sub- series.     Hysteriales. 

5  „  Discomycetes. 
Appended.     Discolichenes. 

2  Sub-class.     Basidiomycetes. 

1  Series.     Protobasidiomycetes. 

1  Sub-series.   Uredinales. 

2  „  Auriculariales. 

3  „  Tremellinales. 

4  ,,  Pilacrales. 

2  Series.  Autobasidiomycetes. 

1  Sub-series.     Dacryomycetes. 

2  ,,  Hymenom.ycetes. 
Appended.     Hymenolichenes. 

3  Sub-series.    Phalloidese. 

4  „  Gasteromycetes. 
Appended.     Gasterolichenes. 

Fungi  imperfecti. 

III.  DIVISION.     EMBRYOPHYTA   ZOIDIOGAMA    (Archego- 

niatae). 

I.  Sub-division.     Bryophyta  (Muscinei). 
1  Class.     HEPATICJE. 

1  Series.     Marchantiales. 

2  ,,         Anthocerotales. 

3  „         Jungermanniales. 

1  Sub-series.     Anacrogynae. 

2  „  Acrogynaa. 
.     2  Class.     Musci. 

1  Sub-class.     Sphagnale.s. 

2  ,,  Andre  (Bales. 

3  ,,  Archidiales. 

4  ,,  Bryales. 

1  Series.     Cleistocarpae. 

2  ,,         Stegocarpae. 

1  Sub-series.    AcrocarpaB. 

2  ,,  Pleurocarpa. 
II.  Sub  division.     Pteridophyta. 

1  Class.    FILICALES. 

1  Sub-class.     Filices. 

1  Series.     Plauithalloste. 

2  „         TuberitballossB. 

2  Sub-class.     Hydropterides. 


APPENDIX   ON   THE    CLASSIFICATION   OF   PLANTS. 


591 


2  Class.     EQUISETALES. 

1  Sub -class.     Jaosporce. 

2  ,,  He.terospnrce, 

3  Class.     SPHENOPHYLLALES. 

4  „        LYCOPODICELES. 

1  Sub-class.     Isoaporcr, 

2  ,,  Heterosporce. 

IV.  DIVISION.     EMBRYOPHYTA  SIPHONOGAMA. 

(Siphonogamaa,  Phanerogamaj). 
I.  Sub-division.     Gymnospermae. 

1  Class.    CYCADALES. 

2  ,,        CORDAITALES. 

3  ,,        BENNETTITALES. 

4  ,,          CONIFERS. 

5  ,,         GNETALES. 

II.  Sub-division.     Angiospermss. 

1  Class.     CHALAZOGAM^:. 

Series.     Verticillatae. 

2  Class.     ACROGAM^E. 

1  Sub-class.     Monocotyledon  ecf. 


1  Series.  Pandanales. 


6  Series.  Spathiflorre. 


2       „ 

Helobiae. 

7 

Farinosa?. 

3       „ 

Glumifloraa. 

8       ,, 

Liliifloree. 

4       „ 

Principes. 

9      „ 

ScitaniineaB. 

5      „ 

Synauthse.                                   10      „ 

Microspermaa. 

2.  Sub-class  Dicotylrdonea. 

1  Group  of  Series.     ArchicbJamydeae. 

1  Series. 

Piperales. 

13  Series 

Sarraceniales. 

2       „ 

Juglandales. 

14       „ 

Eosales. 

3       „ 

Salicales. 

15       „ 

Geraniales. 

4       „ 

Fagales. 

16      „ 

Sapindales. 

5       ,, 

Urticales. 

17      „ 

Ehamnales. 

6      ,, 

Proteales. 

18      „ 

Malvales. 

7      „ 

Santalales. 

19      „ 

Parietales. 

8       „ 

Aristolochiales. 

20       „ 

Opuntiales. 

9 

Polygonales. 

21       „ 

Thymelaeales. 

10       „ 

Centrosperiiia3. 

22       „ 

Myrtifloraa. 

11       >, 

Eanales. 

23       „ 

Umbellitloraa. 

12       „ 

Ehosadales. 

2  Group  of  Series.     Sympetalaa. 

1  Series. 

Ericales.                                          6  Series. 

Plantaginales. 

2       „ 

Primulales. 

7       „ 

Eubiales. 

3       „ 

Ebenales. 

8       „ 

Aggregatse. 

4       „ 

Contoi'tae. 

9      „ 

Campaiiulatse. 

5      „ 

Tubiflora). 

W.  B. 

QQ 

TABLE    OF    ABBREVIATIONS. 


S  = 
P  =  Petals. 
Pr=  Perianth. 
A  =  Androecium. 
G  =  Gynoeceum. 

<J    =  Male. 

?    =  Female. 

£    =  Hermaphrodite. 

oo  =  Indefinite. 

Names  of  continents  and  countries  have  sometimes  been  abbreviated,  for 
example: — Am. —America  ;  As.=Asia;  Af.=Africa;  Ind.=India,  etc.  N.,  S., 
E.,  W.,=North,  South,  East,  West;  Temp. = Temperate  Kegions;  Trop.= 
Tropics. 


59? 


INDEX. 


Abelia,  556. 

Abies,  124,  129,  130,  132, 
133,  148,  155,  165, 
246,  264,  265,  266. 

Abietaceae,  255,  263,  272. 

Abrus,  470,  473. 

Abutilon,  427. 

Acacia,  473,  474,  475. 
False,  470. 

Acalypha,  434. 

Acanthaceae,  518,  529,  530. 

Acanthus,  530. 

Acer,  122,  441,  442. 

Aceraceae,  441. 

Aceranthus,  390. 

Acetabularia,  12,  63. 

Achillea,  568,  572,  574. 

Achimenes,  528. 

Achlya,  107,  108. 

Achnantheae,  21. 

Achras,  511. 

Aeinetffi,  68,  72. 

Aconitum,  37t>,  383. 

Acorin,  306. 

Acorus,  303,  304,  306. 

Acrasieae,  6. 

Acrocarpi,  196. 

Acrocomia,  301. 

Acrogynae,  192. 

Acrospermaceas,  132. 

Acrostichum,  213. 

Acrotonous,  331. 

Acrotylacea3,  83. 

Acrotylus,  83. 

Actffia,  379,  380,  382. 

Actiuidia,  415. 

Adansonia,  427. 

Adder's  tongue,  211. 

Adenanthera,  475. 

Adiantum,  201,  206,  213. 

Adiuida,  17. 

Adlumia,  395. 

Adonis,  379,  383. 

Adoxa,  453,  555. 

Aerobic,  31. 

Mchmea,  319,  320. 

^Ecidiospores,  147. 

JScidium,  147,  148,  150, 
155. 


Jllgiceras,  513. 
^Egilops,  296. 
.'Egopodium,  494. 
^Eschynanthus,  528. 
^Esculinae,  439. 
^Esculus,  440. 
yEthalium,  8. 
^thusa,  495,  498. 
Affonsea,  466. 
Agapanthus,  312,  314. 
Agar-Agar,  33,  84. 
Agaricaceae,  166. 
Agaricinei,  171. 
Agathis,  263. 
Agave,  318. 
A^aveae,  318. 
Ageratum,  571. 
Aggregatae,  505,  564. 
Agrapbis,  312. 
Agiiinonia,  459,  460. 
Agrimoniese,  459. 
Agrimony,  459. 
Agropyrum,  113,  295. 
Agrostemuaa,  365,  367. 
Agrostideas,  294. 
Agrostis,  294. 
Ahnfeltia,  83. 
Ailanthus,  439. 
Aira,  294. 
Aizoaceae,  374. 
Aizoideae,  874. 
Aizoon,  375. 
Ajuga,  47,  537. 
Ajugeae,  537. 
Akebia,  390. 
Akinetes,  10. 
Alaria,  7i,  72. 
Albugo,  107. 
Albumen,  246. 
Albuminous,  249. 
Albumose,  473. 
Alchemilla.  460. 
Alchornea,  432. 
Alcoholic     fermentation, 

97. 

Alder,  8,  118,  341. 
Aldrovaiidia,  408,  409. 
Aleurites.  434. 
Algae,  1,  4,  8. 

593 


Algal-Fungi,  95,  96. 
Albagi,  472. 
Alisma,  281,  282. 
Alismaceae,  278,  281. 
Alismese,  281. 
Alkanet,  534. 
Alkanna,  534,  535. 
Alliariinae,  404. 
Allieae,  312. 
Allium,  312,  313,  314. 
Alloplectus,  528. 
Allosorus,  213. 
Almeidea,  437. 
Almond,  461,  462. 
Alnus,   8,   117,   118,  341, 

342. 

Alocasia,  306. 
Aloe,  274,  312,  313,  314. 
Aloineffi,  312. 
Alonsoa,  525. 
Alopecurus,  290,  294,  296. 
Alpine  Violet,  513. 
Alpinia,  326. 
Alsine,  364,  366. 
Alsineee,  365. 
Alsodeia,  411. 
Alsophila,  214,  215. 
Alstroemeria,  318. 
Alstrremeriese,  318. 
Alternanthera,  369. 
Althaea,    426,     428,    429, 

430. 

Althenia,  279. 
Alyssinae,  404 
Alyssum,  400. 
Amanita,  167,  171. 
Ainarautaceae,  3ii4,  368. 
Amarant-tree,  468. 
Amarantus,  368,  369. 
Amarylleae,  317. 
Amaryllidaceffi,  310,  316. 
Amaryllis,  317,  318. 
Amber,  267. 
Ambrosia,  573. 
Ambrosieaj,  564,  567,  573. 
Ambrosinia,  305. 
Amelanchier.  464,  465. 
Amentaceoe.  337. 
Amherstia,  468. 


594 


INDEX. 


Ammannia,  483. 
Ainmi,  494. 
Ammiese,  494. 
Ammobium,  573. 
Ammoniac-gum,  498. 
Ammophila,  295. 
Amomis,  488. 
Amorpba,  470. 
Ampelidaceee,  445. 
Ampelopsis,  445,  447. 
Amphidinium,  16. 
Amphigastria,  181,  188. 
Amphipleureaa,  21. 
Amphispbeeriaceaa,  130. 
Aiuphithecium,  186. 
A  mphitropideaa,  21. 
Arophorese,  21. 
Amsonia,  544. 
Amygdalaceffi,  461,  466. 
Amygdalin,  462. 
Amygdalus,  461,  462. 
Amjris,  438. 
Anabsena,  25,  219. 
Anacampseros,  373. 
Anacamptis,  332. 
Anacamptodon,  197. 
Anacardiaceee,  439. 
Anacardium,  439. 
Anacrogynse,  192. 
Anacyclus,  572,  574. 
Anadyomene,  62. 
Anaerobic,  31. 
Anagallis,  513. 
Atiamirta,  390. 
Ananassa,  319,  320. 
Ana^tatica,  401. 
Anathyllis,  471. 
Anatropous,  242,  243. 
Ancbusa,    150,    531,   532, 

534,  535. 

AneylistaceoB,  104. 
Ancylonema,  44. 
Andira,  472,  473. 
Andresea,    185,    187,   188, 

195. 

Andrcecium,  239. 
Androgenesis,  14. 
Andromeda,  161,  508. 
Andrornedeae,  508. 
Andropogon,  289,  293, 296. 
Andropogoneae,  293,  296. 
Androsace,  512,  513. 
Androspore,  57. 
Aneimia,  215. 
Anelaterese,  192. 
Anemone,  379,  384. 
Anemonese,  384. 
Anemonopsis,  379. 
Auethum,  496,  498. 
Aneura,  191,  192. 
Angelica,  496,  498. 


Angiopteris,  212. 
Angiospermee,  3,  234,  239, 

250,  273. 
Angiosperms,     237,     245, 

248. 

Angiosporese,  82. 
Angosturee,  Cortex,  437. 
Anguliferre,  21. 
Anise,  498. 
Anlage,  90. 
Aimatto,  412. 
Auuularia,  225. 
Aimularise,  225. 
Aunulus,  195,  209. 
inferus,  167. 
superus,  ]68. 
Anoda,  428,  429. 
Anodic,  480. 
Anomodon,  197. 
Anona,  388. 
Anonacese,  388. 
Autennaria,  124,  573. 
Anthemideffi,  572 
Antbemis,  569,  572,  574. 
Anther,  237,  238. 

Fibrous  layer  of,  241. 
Structure  of,  239. 
Anthericese,  312. 
Anthericum,  312,  313. 
Antberidiura,  13,  100, 198. 
Antherozoid,  13. 
Anthocarp,  374. 
Anthoceros,  25,  186,  187, 

188,  189,  191. 
Anthocerotese,  191. 
Antbolyza,  321. 
Anthostema,  432,  433. 
Anthoxanthum,  295,  296. 
Anthrax  bacillus,  31,  39, 

40. 

Anthriscus,  493,  495,  498. 
Anthurium,  304. 
Anthyllis,  471. 
Antiaris,  356. 
Antipodal  cells,  248. 
Antirrhineas,  523. 
Antirrhinum,  524,  527. 
Antisepsis,  32. 
Ai>iba,  424,  425. 
Apetalse,  336,  337. 
Aphanizomenon,  25. 
Aphanocapsa,  24. 
Aphanocbsete,  54. 
Aphtha,  180. 
Aphyllantbes,  312. 
Apiocystis,  51. 
Apios,  471. 
Apiosporium,  124. 
Apium,  494,  493. 
Aplanogametangium,  12. 
Aplanogainetes,  12. 


Aplanospores,  10. 
Apocuiaceas,  542,  543,549. 
Apocvnum,  5  ±4. 
Apogamy,  203. 
Aponogeton,  281. 
Aponogetonacese,  281. 
Apospory,  188. 
Apostasia,  329. 
Apostasieas,  328,  329. 
Apotbecium,  118,  132. 
Apple,  127,  130,  4(54,  465. 
Apricot,  121,  461,  462. 
Aquifoliaceaa,  444. 
Aquilegia,  378,   379,  381, 

382. 

Arabis,  402. 
Araceffi,  276,  278,  303. 
Arachis,  469,  472,  473. 
Aralia,  491. 

Araliacese,  454,  491,  549. 
Araucaria,  237,  263. 
Araucariaceaa,    257,    263, 

272. 

Arbor  vitae,  267. 
Arbutete,  508. 
Arbutus,  508. 
Archangelica,  496,  498. 
Archegoniata,  3,  185. 
Archegonium,  3, 184,  198. 

Development  of,  201. 
Archesporium,  186,  202. 
Archicarp,  1*20. 
Archidiurn,  193,  195. 
Arctostaphylos,  161,  508. 
Arcyria,  7,  8. 
Ardisia,  513. 
Areca,  301,  302. 
Areca-palm,  302. 
Arecineae,  301. 
Arenaria,  366. 
Arenga,  301. 
Argemone,  395. 
Aria,  152. 
Aril,  255,  258. 
Arineae,  305. 
Arisarum,  305. 
Aristida,  295. 
Aristolochia,  499,  500. 
Aristolochiacea3,  499. 
Aristolochiales,  499. 
Aristotelia,  425. 
Armeniaca,  461. 
Armeria,  514. 
Armillaria,  117, 169,  170. 
Arnebia,  533. 
Arnica,  572,  574. 
Arnoseris,  571. 
Aronia,  464. 
Arrack,  2%,  301. 
Arrow-head,  282. 
Arrow-poison,  544,  546. 


INDEX. 


595 


Arrow-root,  327,  434. 
Artabotrys,  388. 
Artemisia,  569,  572,  573, 

574. 

Arthonia,  134. 
Arthoniaceffi,  134. 
Artbrosporous,  29. 
Arthrotaxis,  267. 
Artichoke,  570,  574. 

Jerusalem,  572,  574. 
Artocarpete,  354. 
Artocarpus,  356. 
Arum,  303,  304,  305,  306. 
Arurido.  294,  296. 
Asafoetida,  498. 
Asaruin,  499,  500. 
Asclepiadaceas,   238,  542, 

544. 

Asclepias,  545,  546. 
Ascobolaceae,  135. 
Ascobolus,  136. 
Ascocarps,  88. 
Ascocorticium,  116,  117. 
Ascogone,  120. 
Ascoidea,  108. 
Ascoideaceae,  108. 
Ascolichenes,  95, 116, 136. 
Ascomycetes,  95,  114, 116. 
Ascophyllum,  73,  75. 
Ascospore,  88. 
Ascus,  88. 
Aseroe,  173. 
Asexual  rep  reductive  cells, 

10. 

Ash,  127,  546,  547. 
Asimina,  388. 
Asparageaa,  314. 
Asparagus,  314,  316. 
Aspen,  152,  338. 
Aspergillus,  122. 
Asperifoliee,  532. 
Asperococcus,  70. 
Asperugo,  534. 
Asperula,  552,  553. 
Asphodelus.  312,  313,  314. 
Aspidistra,  314. 
Aspidium,  203,  204,  207, 

213,  214. 

Aspidosperma.  3i4. 
Asplenium,  213,  214. 
Astelia,  316. 
Aster,  569,  571,  573. 
Asterese,  571,  573. 
Asteriscus,  574. 
Asterocystis,  78. 
Asterophylliteee,  225. 
Asteropbyllites,  225. 
Astragaleee,  470. 
Astragalus,  114,  470,  473. 
Astrantia,  493. 
Astrocarpus,  407. 


Atherurus,  305. 
Athyrium,  204,  207,  213. 
Atragene,  379,  385. 
Atraphaxis,  360. 
•Atriplex,  371,  372. 
Atriplicesa,  371. 
Atropa,     519,    521,     522, 

523. 

Atropine,  522. 
Attalea,  297,  301. 
Attar  of  Hoses,  460. 
Aubrietia,  400. 
Aucuba,  491. 
Aulacomnium,  197. 
Aurantieae,  437. 
Auricula,  156. 
Auricularia,  156. 
Auriculariaceae,  145,  155. 
Austrian  Pine,  267. 
Autobasidia,  144. 
Autobasidiomycetes,     96, 

145,  157. 
Autoacious,  148. 
Autoxenous,  148. 
Auxiliary  cells,  81. 
Auxospore,  19. 
Avena,  294,  296. 
Aveneae,  294. 
Avens,  458. 
Averrhoa,  416. 
Avicennia,  535. 
Avignon  grain,  448. 
Awl  wort,  401. 
Awn,  288,  290. 
Azalea,  508. 
Azolla,  25,  219. 


"Bablah,"  475. 

Bacillus,   26,  28,   30,  31, 

35,  36,  37. 
antbracis,  39. 
diphtheria,    leprae, 
mallei,  tetani,  tuber- 
culosis, typbosus,  40. 

Bacteria,  4,  5,  8,  9, 10,  22, 
26. 

Bacterium,  26,  28,  30,  35, 
39. 

Bactris,  301. 

Bseomyces,  140,  142. 

Balanopbora,  504. 

Balanophoraceae,  504. 

Ballota,  53d. 

Balsaminaceae,  420. 

Balsamodendron,  438. 

Balsam  of  Copaiba,  468. 
of  Peru,  473. 

Damboo,    289,   291,    292, 
293.  • 

Banibusa,  289,  291,  293. 


BambusesG,  293,  296. 
Banana,  324,  H25. 
Baneberry,  382. 
Bangia,  77,  78. 
Bangioideae,  77. 
Banksia,  450. 
Baobab,  427. 
Barbaceoia,  318. 
Barbarea,  402. 
Barberries,  389. 
Barbula,  196. 
Bark-canker,  169. 
Barley,  113,  292,  296. 
Barosma,  436. 
Barringtonia,  489. 
Bartonia,  476. 
Bartramia,  197. 
Bartsia,  526. 
Basella,  371. 
Baselleae,  371. 
Basidial-layer,  89. 
Basidiocarp,  89. 
Basidiolichenes,   96,   145, 

176. 
Basidiomycetes,   96,   114, 

144,  145. 
Basidiospore,  88. 
Basidium,  89,  144,  146. 
Basitonous,  331. 
Bassia,  511. 
Bast,  251,  425,  430. 
Bastardia,  428. 
Batatas,  516,  517. 
Batidacese,  372. 
Batis,  372. 
Batracbium,  383. 
Batrachospermum,  80,  83. 
Bauhinia,  467. 
Bay  berry-tree,  490. 
'•  Bay-rum,"  489. 
Beaked  parsley,  495. 
Beak-rush,  286. 
Bear-berry,  508. 
Beard  lichen,  143. 
Beech,  127,  134,  164,  1G5, 

526. 

Beef-steak  fungus,  166. 
Beer-yeast,  177,  178. 
Beet,  369. 
Beet-root,  372. 
Beggiatoa,  26,  28,  37. 
Begonia,  477,  478. 
Begoniacese,  475,  477. 
Bellis,  569,  572. 
Beniucasa,  481. 
Berberidacese,  238,  389. 
Berberis,  149,  389,  390, 
Bergamot,  438. 
Bergia,  413. 
Berteroa,  400. 
Bertholletia,  489. 


596 


INDEX. 


Feta,  369,  370,  372. 

Betal,  363. 

Betonica,  538. 

Betony,  538. 

Betula,  342. 

Betulaceje,  341. 

Biarum,  305. 

Biatorella,  134. 

Bicornes,    33t5,   451,  505, 
506. 

Biddulphiese,  21. 

Bidens,  566,  572. 

Biebersteinia,  419. 

Bignonia,  529. 

Bignoniaceae,  518,  529. 

Bilberry,  509. 

Billardiera,  455. 

Billbergia,  320. 

Bindweed,  515,  516. 

Biota,  268. 

Birch,  117,  135,  165,  342. 

Bird-cherry,  461,  462. 

Bird-lime,  501.  504. 

Bird's-foot,  472. 

Bird's-foot-trefoil,  471. 

Biscutella,  401. 

Bitter-cress,  402. 

Bitter-sweet,  522. 

Bixa,  412. 

Bixaceae,  412. 

Blackberry,  461. 

Black-boy,  312. 

Black-currant,  153,  455. 

Black-mustard,  401,  405. 

Black-pepper,  363. 

Blackthorn,  462. 

"Bladder"  plums,  117. 

Bladder-senna,  470. 

Bladder-wort,  528. 

Blasia,  25,  191,  192. 

Blattiaceas,  483. 

Bleclmum,  209,214,  254. 

Bletia,  332. 

Blight,  132. 

Blindia,  196. 

Blinks,  373. 

Blitum,  369. 

Blood-red  Currant,  455. 

Blue-green  Algoe,  5,  22. 

Bocconia,  395. 

Boehmeria,  353. 

Boerhaavia,  374. 
!  Bog-mosses,  193. 
'  Bog-myrtle,  351. 

Bog  Wortleberry,  509. 

Boisduvalia,  485. 

Boletus,  166. 

Bomarea,  318. 

Bombaceae,  427. 

Bombax,  427. 

Bonuemaisonia,  83. 


Bonnemaisoniaceae,  83. 
Borage,  533. 
Boragese,  532,  533. 
Boraginaceae,     515,     531, 

532,  537. 
Borago,  533,  534. 
Borassinae,  301. 
Borassus,301. 
Borderea,  323. 
Boronieae,  436. 
Borreria,  550. 
Boschia,  190. 
Bossiaea,  472. 
Boswellia,  438. 
Bo-tree,  356. 

Botrychium,202, 210,  211. 
Botrydiaceae,  47,  59. 
Botrydium,  59. 
Botrytis,  128,  134,  135. 
Bottle-gourd,  481. 
Bouchea,  535. 
Bougainvillea,  374. 
Boussingaultia,  371. 
Bouvardia,  550. 
Bovista,  174. 
Bowenia,  253,  254. 
Bowiea,  312. 
Box,  434. 

Brachypodium,  294. 
Brachythecium,  197. 
Bracken-fern,  207,  213. 
Bract,  235. 

Bracteole,  235,  275,  334. 
Bradypus,  8,  54,  356. 
Brahea,  300. 
Bramble,  458. 
Branching  of  Palm,  298. 
Brand-fungi,  95, 108, 109. 
Brand-spores,  91. 
Brasenia,  386. 
Brassica,  399,  400,  401. 
Brassicinae,  404. 
Bray  era,  460. 
Brazil-nuts,  489. 
Breadfruit,  356. 
Briza,  290,  294,  296. 
Brome,  296. 
Bromeliaceae,     308,     309, 

310,  318. 
Bromus,    287,    289,    290, 

293,  296. 
Brook-weed,  513. 
Broom,  472. 
Broom-rape,  528. 
Brosimum,  356. 
Broussonetia,  354. 
Browallia,  521. 
Brown  Algaa,  1. 
Brownea,  468. 
Brownian  movement,  28. 
Brugmansia,  504. 


Brunfelsia,  521. 
Bryaceae,  197. 
Bryonia,  481. 
Bryophyllum,  451,  452. 
Bryophyta,  1,  234. 
Bryopsidacese,  47,  60. 
Bryopsis,  60,  62. 
Bryum,  197. 
Buchu,  436. 
Buck-bean,  543. 
Buckthorn,  4  48. 
Buckwheat,  361. 
Buettneria,  422. 
Buettneriaceae,  422. 
Bugle,  537. 
Bulbine,  312. 
Bulbochaate,  55,  56. 
Bulbocodium,  310. 
Bulbophyllum,  332. 
Bulgaria,  134. 
Bulgariaceee,  134. 
Bullace,  461,  462. 
Bulliarda,  452. 
Bull-rush,  303. 
Bumelia,  511. 
Bunchosia,  442. 
Bunias,  400,  403. 
Bupleurum,  491,  494. 
Burdock,  570. 
Burmanniaceae,  328. 
Bur-marigold,  572. 
Bur  Parsley,  497. 
Bur-reed,  302. 
Burseraceaa,  438. 
Butcher's  broom,  316. 
Butomeae,  281. 
Butornus,  281,  282. 
Butterbur,  571. 
Butter-tree,  414. 
Butter-wort,  528. 
Butyric-acid-bacillus,  38. 
Buxacese,  434. 
Buxbaumia,  197. 
Buxbaumiaceae,  197. 
Buxus,  434. 

Cabbage,  401. 
Cabomba,  386. 
Cabombeae,  386. 
Cacalia,  572. 
Cactaceae,  375. 
Cacti,  375. 
Cactiflorae,  375. 
Cseoma,  147, 148,  152. 
Caesalpim'a,  468. 
Caesalpiniaceae,  466,  470. 
Caffeine,  441,  553. 
Cajanus,  471. 
Cajeput-oil,  489. 
Cajophora,  476. 
Cakile,  403. 


INDEX. 


597 


Calabar-bean,  471,  473. 
Calabash,  529. 
Caladium,  o06. 
Calamagrostis,  289,  294. 
Calamiutha,  540. 
Calamites,  224. 
Calamus,  298,  801,  303. 
Calamus-oil,  300. 
Calaudrinia,  373. 
Calathea.  327. 
Calceolaria,  525,  527. 
Calcocytacea?,  15. 
Calendula,  565,  572. 
Calendule®,  572. 
Caliciaceas,  134. 
Calicium,  134. 
Calla,  305,  307. 
Callea3.  305. 
Calliandra,  475. 
Callianthenmm,  379. 
Callicarpa,  535. 
Calligonum,  361. 
Calliopsis,  572. 
Callistemon,  489. 
Callistephus,  573. 
Callithamnion,  78,  79,  84. 
Callitrichaceas,  434, 
Callitriche,  434. 
Callitris,  269. 
Calloria,  134. 
Calluna,  507. 
Calocera,  158,  159. 
Calonyction,  516. 
Calophyllum,  414. 
Calothamnus,  489. 
Calothrix,  25. 
Caltha,  379,  380,  381,  382. 
Calycanthacese,  389. 
Calycantbus,  389. 
Calyceraceee,  556,  560. 
Calypogeia,  192. 
Calypso,  332. 
Calyptospora,  152. 
Calyptra,  186. 
Calystegia,  516. 
Calyx-stamens,  335. 
Camelina,  400,  401. 
Camellia,  414,  415. 
Campanula,  153,  561,  562. 
Campanulacese,  561,  563. 
Campanulinee,    505,    560, 

564,  569. 
Camphor,  392. 
Campion,  367. 
(Jampylopus,  196. 
Campylosperme8e,493,497. 
Campy lotropous,  242,  243. 
Canada-balsam,  266. 
Cananga,  388. 
Canarina,  562. 
Canary-grass,  295. 


Canavalia,  471. 
Candollea,  413,  564. 
Candolleaceee,  564. 
Cane,  298,  301. 
"  Canker,"  127. 
Canna,  326. 
Cannabacese,  356. 
Cannabis,  357,  358. 
Cannacese,  277,  326,  327. 
Canterbury-bell,  561. 
Cantharellei,  172. 
Cantharellus,  170,  172. 
Caoutchouc,  434,  544,  546, 

563. 
Capers,  405. 
Capillitium,  7,  174. 
Capirona,  549. 
Capnodium,  124 
Capparidacea?,  405. 
Capparis,  405,  406. 
Capraria,  525. 
Caprification,  355. 
Caprificus,  355. 
Caprifoliaceee,    454,    548, 

549,  553,  556,  557. 
Caprifolium,  554. 
Capsella,  400,  401,  402. 

Capsellinffi,  404. 

Capsicum,  521,  522. 

Capsosira,  26. 

Capsule,  186. 

Caragana,  470. 

Caraway,  494,  498. 

Cardaminas,  401. 

Cardamine,  400,  402, 404. 

Cardamom,  326. 

Cardiospermum,  441. 

Carduus,  569. 

Carex,  113,  151,  247,  286, 
287. 

Carica,  476. 

Caricese,  286. 

Caries  dentium,  38. 

Carlina,  570. 

Carludovica,  302. 

Carmichffilia,  470. 

Carnation,  367. 

Carnaiiba-wax,  301. 

Carob-bean,  466,  468. 

Carpels,  235,  238. 

Carpinus,  117,  344. 

Carpoasci,    95,   115,  116, 
118. 

Carpogonium,  77,  81. 

Carpophore,  91,  492. 

Carpospore,  77,  82. 

Carragen,  33,  84. 

Carrot,  496,  497,  498. 

Carthamus,  570,  574. 

Carum,  493,  494,  498. 

Carya,  350. 


Oaryophyllacero   336,  364. 
Daryopsis,  288. 
Daryota,  301. 
Dascara,  448. 
Dascarilla,  434,  550. 
Oashew-nut,  439. 
Cassandra,  508. 
Cassava,  434. 
Cassia,  467,  468. 
Cassine,  444. 
Cassiope,  508. 
Cassytha,  392. 
Castanea,  346. 
Castilloa,  356. 
Castor-oil,  431,  434. 
Casuarinaceee,  339. 
Casuarina,  273, 274. 
Casuarinifloree,  339. 
Cataba,  414. 
Catabrosa,  294. 
Catalpa,  529. 
Catananche,  566,  571. 
Catasetum,  332,  333. 
Catch-fly,  367. 
Catechu,  475. 
Catha,  444. 
Catharinea,  197. 
Cathartocarpus,  467,  468. 
Catmint,  539. 
Catodic,  480. 
Cat's-ear,  571. 
Cat's-foot,  573. 
Cat's-tail,  294. 

Cattle-beet,  372. 

Cattleya,  332. 

Caucalis,  497. 

Caudicle,  331,  332. 

Caulerpa,  10, 61,  62. 

Caulerpacerc,  47,  61. 

Cauliflower,  405. 

Cayenne-pepper,  522. 

Ceanothus,  448. 

Cecropia,  356. 

Cedar,  266. 

Cedrat,  438. 

Cedrcla,  436. 

Cedrus,  266. 

Celandine,  394. 

Celastractse,  444. 

Celastrus,  444. 

Celery,  494,  498. 

Celidium,  134. 

Cell,  Vegetative,  228. 

Celosia,  368,  369. 

Celsia,  525. 

CeltidesD,  352. 

Cenangiacese,  134. 

Cenangium,  134. 

Cenchrus,  295. 

Centaurea,  565,  567,  568, 
569,  570,  573. 


598 


INDEX. 


Centaury,  543. 
Centradenia,  484. 
"  Central  cell,"  185. 
Centranthus,  557,  558. 
Centrolepidaceee,  308,  b09. 
Centrolepis,  3U9. 
Centrolobium,  472. 
Centropogon,  563. 
Ceutunculus,  512,  513. 
Cephaelis,  550,  553. 
Cephalanthera,  328,  331. 
Cephalaria,  560. 
Cephalotaceae,  454. 
Cephalotaxeas.  259. 
Cephalotus,  453. 
Ceramiaceaa,  81. 
Ceraouum,  78,  80,  84. 
Cerastium,  364,  366. 
Cerasus,  462. 
•Ceratiomyxa,  8. 
Ceratium,  16,  17. 
Ceratocapnos,  396. 
Ceratodon,  196. 
Ceratonia,  468. 
Ceratophyllacese,  388. 
Ceratophyllum,  388. 
Ceratostomaceas,  130. 
Ceratozamia,     238,     253, 

254. 

Cerbera,  544. 
Cercis,  467,  468. 
Cereus,  375.  377. 
Cerinthe,  533  . 
Ceropegia,  546. 
Ceroxylon,  301. 
Cestre'ffi,  522. 
Cestrum,  522. 
Ceterach,  214. 
Oetraria,  138, 141,  142. 
Cheenorneles,  465. 
Cbseropbyllum,  495,  498. 
Chastangiaceee,  83. 
Cheetoceros,  20. 
Chaatocladiacese,  100. 
Cbaatocladium,  100. 
Cheetomiaceffi,  129. 
Cbffitomium,  129. 
Chsetomorpha,  58. 
Chsetopeltis,  54. 
Chsetophora,  54. 
Chsetopboraceae,  47,  54. 
Chaetopteris,  70. 
Chalaza,  242. 
Chalazogames,  273. 
Chalazogams,  273. 
Chamsecyparis,  268,  269. 
Chameedorea,  298,  301. 
Chameedoris,  02. 
Chameelaucieee,  489. 
Chamaelaucium,  489. 
Chamsenerium,  484. 


Chamserops,  298,  300,  301, 

302. 

Chamffisiplion,  22,  24,  25. 
Chamaasiphonaceae,  24,  25. 
Chamomile,  572,  574. 
Chantransia,  83. 
Chara,  65,  66,  «7. 
Characere,  1,  4,  10,  14,  64. 
Characium,  47,  51. 
Cbarese,  67. 
Charlock,  404. 
Cbeilanthes,  213. 
Cbeiranthus,     399,     400, 

402. 

Cheirostemon,  427. 
Cbelidonium,  394,  395. 
Cbelone,  525. 
Chenopodiacese,  364,  369. 
Chenopodiese,  369. 
Chenopodina,  371,  372. 
Chenopodium,  369,  372. 
Cberry,  117,  156,  461,  462. 
Cherry-laurel,  462. 
Chervil,  495,  498. 
Chervil-root,  498. 
Chick-pea,  470. 
Chickweed,  366. 
Chicory,  570,  574. 
Cbilies,  522. 
Cbimaphila,  505. 
Chimonanthus,  389. 
China-grass,  353. 
Chinese  galls,  439. 
Chiococca,  550. 
Chionanthus,  547. 
Chionodoxa,  312. 
Chive,  312. 
Cblamydomonas,  4S. 
Chlamydoinoneae,  14. 
Chlamydomucor,  97,  98. 
Chlamydospore,  90. 
Chlora,  47,  543. 
Chlorangium,  51. 
Chloranthaceog,  363. 
Chloranthus,  363. 
Cblorideee,  295. 
Chloris,  295. 
Chlorochytrium,  47,  51. 
Chlorococcum,  51. 
Chlorocystis,  51. 
Chlorophycese,  1,  14,  46. 
Chlorophytuin,  312. 
Chlorosphsera,  51. 
Chlorosphseracese,  47,  51. 
Chlorospleuium,  135. 
Chlorotylium,  54. 
Choanephora,  100. 
Choanepboraceee,  100. 
Chocho,  481. 
Choiromyces,  124. 
Choisya,  436. 


Chondrus,  79,  83,  84, 
Chorda,  72. 
Chordaria,  71. 
Chordariaceas,  71. 
Choripetalaa,  336,337,  505, 

561. 

Chorisia,  427. 
Choristocarpacese,  70. 
Choristocarpus,  70. 
Chromaceaa,  15. 
Chromulina,  15. 
Cbroococcaceae,  24. 
Chroococcus,  24,  176. 
Chrysalis  Fungus,  127. 
Chrysanthemum,  572. 
Chry&arobin,  473. 
Chrysobalanaceoe,  462, 466. 
Cbrysobalanus,  462. 
Chrysomonadinacese,     15, 

17. 
Chrysomyxa,147, 148, 153, 

155. 

Chrysophyllum,  511. 
Chrysopyxaceee,  15. 
Chrysopyxis,  15. 
Chrysosplenmm,  452,  454. 
Chylocladia,  83. 
Chysis,  333. 
Chytridiales,  95,  102. 
Chytridium,  103. 
Cibotium,  214,  215. 
Cicely,  495. 
^icendia,  543. 
Cicer,  470. 

Cichorieae,  561,  568,  570. 
Cichorium,  570,  574. 
Oicinnobolus,  ]20. 
Cicuta,  494,  498. 
Cilioflagellata,  17. 
Cimaruoli,  355 
Cimicifuga,  383. 
iJmchona,  548,  549,   550, 

553. 

Cinchonese,  550. 
Cinchouin,  553. 
Cinclidotus,  ]97 
Jineraria,  572,  574. 
Cinnamon,  392. 
Cinnamomuin,  391,  392. 
Cinquefoil,  458. 
Cipnra,  321 
Cimea,  485,  486. 
Circinate,  208. 
Cirsium,  151,  568,  569. 
Cissampelos,  390. 
Cissus,  445,  504. 
(Jistaceae,  412. 
Oistifloree,  406,  451. 
Cistus,  412,  503. 
Citharexylon,  535. 
Citriobatus,  455. 


INDIiX. 


;99 


Citron,  438. 
Citronella  oil,  296. 
Citrullus,  479,  480,  481. 
Citrus,  437,  438. 
Cladium,  286. 
Cladochytriurn,  103. 
Cladonia,    139,   140,    141, 

142,  143. 

Cladophora,  11,  58. 
Cladopboraceee,  47,  58. 
Cladopporium,  124. 
Cladothrix,  27,  33,  34,  35. 
Clamp-connections,  86. 
Clarkia,  485. 
Clastidium,  25. 
Clathrus,  173. 
Clavaria,  159,  161. 
Clavariacese,  161. 
Claviceps,  125,  126,  127. 
Clavija,  513. 
Claytonia,  373. 
Cleavers,  552. 
Cleistocai'peas,  195. 
Clematidese,  385. 
Clematis,   378,   379,   380, 

385. 

Cleome,  406. 
Clerodendron,  535. 
Clethra,  509. 
Climacium,  197. 
Clinopodium,  540. 
Clintonia,  563. 
Clitocybe,  171. 
Clitoria,  471. 
Clivia,  317,  318. 
Closterium,  43,  44. 
Clostridium,  31. 
Cloudberry,  461. 
Clover,  135,  471. 
Cloves,  489. 

Club-mosses,  2,  205,  226. 
Club-rush,  285. 
Clusia,  414. 
Clusiacese,  414. 
Cluster-cups,  150. 
Clypeospbaeriaceee,  130. 
Cnicus,  570,  574. 
Cnidium,  495. 
Cobasa,  515. 
Coca,  442. 
Cocaine,  442. 
Cocci,  26. 

Cojccochromaticse,  21. 
Coccoloba,  360. 
Cocconeidese,  21. 
Cocconeis,  21. 
Cocconema,  20. 
Cocculus,  390. 
Coccus,  356. 
Cochineal,  377. 
Cochineal-insect,  377. 


Cochlearia,  398,  400. 
Cochleariinse,  404. 
Cock's-comb,  369. 
Cock's-foot,  294,  296. 
Cocoa-beans,  423. 
Cocoa-butter,  423. 
Cocoa-plum,  462. 
Cocoa-tree,  422. 
Cocoanut,  298,  300,  302. 
Cocoanut,  Double,  301. 
Cocoanut-palm,  301. 
Cocoineee,  300. 
Cocos,  298,  301,  302. 
Codiacero,  61. 
Codiolum,  59. 
Codium,  62. 
Coelastrum,  52. 
Coalebogyne,  432. 
Coaloglosswm,  332. 
Coelospermese,  493,  497. 
Coelospheeriuin,  24. 
Ccenobia,  47,  51. 
Ctenogoniuin,  142. 
Coffea,  550. 
Coffee®,  550. 
Coffee,  555. 
Coffee-plant,  550,  553. 
Coix,  293. 
Cola,  422,  423. 
Colchicaceffi,  309,  310. 
Colchiceae,  310. 
Colchicin,  311. 
Colchicum,  310,  311. 
Coleochffltaceee,  47,  57. 
Coleochffite,  57,  58. 
Coleonema,  436. 
Coleorhiza,  293. 
Coleosporium,    147,     148, 

152, 154. 
Coleus,  540,  541. 
Collema,  138,  142. 
Colletia,  448. 
Collinsia,  525. 
Collomia,  515. 
Collybia,  171. 
Colocasia,  303,  305,  306. 
Colocynth,  481. 
Colts-foot,  571. 
Columba-root,  390. 
Columbine,  382. 
Columella,  187,  189,  193. 
Columnea,  523. 
Columniferse,  421. 
Colus,  173. 
Colutea,  470,  473. 
Comarum,  457,  458. 
Combretaceee,  487. 
Comfrey,  533. 
Comma-bacillus,  40. 
Commelina,  308. 
Commelinaceee,  308. 


Commersonia,  422. 
Commiphora,  438. 
Compass-plant,  572. 
Composites,  556,  560,  561, 

563,  564. 
Comptonia,  350. 
Condurango-bark,  546. 
Cone,  235. 
Cone-scales,  256. 
Conferva,  54. 
Confervoidese,  47,  53. 
Conidia,  87,  90. 

Liberation    and    distri- 
bution of,  91. 
Conidial-layers,  88. 
Conidiocarp,  89, 147. 
Conidio-fructin'cation,  87. 
Conidiophore,  87,  88. 
Conifers,  3,  237,  238,  252, 

255. 
Female  flower   of,  255, 

257. 

Pollination,  258. 
Coniocybe,  134. 
Conium,  494,  498. 
Conjugate,  1,  12,  14,  41, 

88. 

Conjugation,  11. 
Connaracese,  435. 
Conocarpus,  487. 
Conomitrium,  196. 
Contorts?,  505,  541,  549. 
Convallaria,  314,  316. 
Convallariacese,  309,  314. 
Convallariese,  314. 
Convolvulacese,     5,     515, 

522,  532. 

Convolvuleee,  516. 
Convolvulus,      114,     516, 

517. 

Co-operating  cells,  248. 
Copaifera,  467,  468. 
Copal-balsam,  468. 
Copernicia,  300,  301. 
Copper-beech,  157. 
Coprinarins,  171. 
Coprinei,  172 
Coprinus,  172. 
Coptis,  379,  32. 
Cora,  176. 
Corallina,  79,  84. 
Corallinacese,  84. 
CoralliorrhizH,  332. 
Corallorhiza,  5,  332. 
Coral-root,  332. 
Corchorus,  424,  425. 
Cordaitacese,  271. 
Cordiacese,  47-61, 531, 532. 
Cordyceps,  125,  127,  1£8. 
Cordyline,  316. 
"Core,"  463. 


600 


INDEX. 


Coriander,  497,  498. 
Coriandrum,  493,  497, 498. 
Cork-elm,  352. 
Cork-oak,  348. 
Cormophyta,  1. 
Cormophytes.  234. 
Cornacese^  490,  549. 
Cornel,  490. 
Corn-cockle,  367. 
Coru-flower,  567. 
Corn-poppy,  395. 
Cornus,  490,  491. 
Corona,  317,  476. 
Coronilla,  472,  473. 
Correa,  436. 
Corrigiola,  365,  367. 
Corsinia,  190. 
Corsiniacese,  190. 
Cortex  angosturse,  437. 
Corticium,  144,  161. 
Cortinarius,  171. 
Cortusa,  512. 
Corydalis,  334,  395,   396, 

397. 

Corylacese,  341,  343. 
Corylus,    122,    343,    344, 

348. 

Corypha,  298,  300. 
Coscinodisceee,  21. 
Coscinodiscus,  20. 
Coscinodon,  197. 
Cosmanthus,  515. 
Cosmarium,  42,  43,  44. 
Costus,  326. 
Cotoneaster,  463,  465. 
"  Cotton  "  427,  429,  430. 
Cotton-grass,  286. 
Cotton-thistle,  570. 
Cotyledon,  451. 
Cotyledons,  247. 
Couch,  295. 

Coumarin,  296,  473,  553. 
Cover-scale,  255,  256. 
Cow-bane,  494,  498. 
Cowberry,  509. 
Cow-parsnip,  496. 
Cow-tres,  356. 
Cow-wheat,  526. 
"  Crab's-eyes,"  470. 
Crambe,  400,  403. 
Craniolaria,  529. 
Crassula,  452. 
Crassulaceee,  451. 
Cratsegeae,  465. 
Cratsegus,  152,  465. 
Craterellus,  162,  172. 
Craterocoha,  156. 
Crenothrix,  30,  37. 
Creosote,  438. 
Crepis,  571. 
Crescentia,  529. 


Crinum,  318. 
Crocus,  320,  321. 
Cronartium,  146,  147, 153, 

155,  156. 

Crotalaria,  472,  473. 
Croton,  431,  434. 
Crowberry,  434. 
Crown-imperial,  314. 
Crozophora,  434. 
Crucianella,  552. 
Crucibulum,  176. 
Cruciferee,  398. 
Crucifers,  398. 
Crucigenia,  51. 
Cruoria,  84. 
Cryptogams,  3,  234. 

Vascular,  2, 198. 
Cryptoglena,  15. 
Cryptogramme,  213. 
Cryptomeria,  2fi7. 
Cryptonemia,  84. 
Cryptonemiales,  82,  84. 
Ctenanthe,  327. 
Ctenium,  295. 
Ctenomyces,  119. 
Cubeb,  363. 
Cucubalus,  367. 
Cucullus,  545. 
Cucumber,  481. 
Cucumis,  121,  480,  481. 
Cucurbita,  478,  479,  480, 

481. 
Cucurbitacese,    475,     478, 

561. 

Cucurbitariacese,  130. 
Cud-weed,  573. 
Cuminum,  497,  498. 
Cunninghamia,  263. 
Cunoniaceee,  454. 
Cuphea,  482,  483. 
Cupressaceee,     257,     262, 

267,  272. 
Cupressus,  241,  245,  268, 

269. 

Cupule,  343. 
Cupuliferae,  341,  345. 
Curare,  546. 
Curculigo,  318. 
Curcuma,  326. 
Curly-mint,  541. 
Currants,  447,  454. 
Curvembryae,  H63. 
Cuscuta,  5,  515,  517. 
Cuscutese,  51H. 
"  Cushion,"  206. 
Cusparia,  437. 
Cuspariese,  437. 
Cutleria,  68,  72. 
Cutleriaceee,  11,  72. 
Cyanophyceee,  22. 
Cyanophyll,  22. 


Cyanotis,  308. 
Cyathea,  214,  215. 
Cyatheaceee,  210,  215. 
Cyathium,  432. 
Cyathus,  176. 
Cycadaceag,  252. 
Cycadeee,  3,  236,  252.  254. 
Cycas,  25,  231,  236,  238, 

251,  252,  253,  254. 
Cyclamen,  334,  512,  513. 
Cyclantliacese,  302. 
Cyclanthera,  481. 
Cyclolobeee,  371. 
Cyclosporeae,  68,  73. 
Cydonia,  463,  464. 
Cylindrocapsa,  14,  55. 
Cylindrocapsaceae,  47,  54. 
Cylindrocystis,  44. 
Cylindrospermum,  22,  25. 
Cymbella,  20,  21. 
Cymbelles;,  20,  21. 
Cymodocea,  281. 
Cymopolia,  63. 
Cynanchum,  546. 
Cynara,  570,  574. 
Cynareee,  569. 
Cynips,  355. 
Cynodon,  295. 
Cynodontium,  196. 
Cynoglossum,  533,  535. 
Cynomorium,  503,  504. 
Cynosurus,  294,  296. 
Cypella,  321. 
Cyperacese,  277,  283,  284, 

291. 

Cyperus,  286,  287,  290. 
Cyphella,  162. 
Cyphiaceee,  562. 
Cypress,  267,  2(58. 
Cypripedileee.  329,  330. 
Cypripedilum,  330. 
Cypripedium,  330. 
Cypsela,  564. 
Cyrtandrese,  528. 
Cystocarp,  14,  58,  82. 
Cystoclonium,  83. 
Cystopteris,  214. 
Cystopus,  107. 
Cytinus,  503,  504. 
Cytisus,  472,  473. 

Dacrydium,  255,  200,  261. 
Dacryomitra,  158.  159. 
Dacryomyces,    134,     158, 

159. 

Dacryomycetaceee,  159. 
Dacryomycetes,    96,   145, 

159. 

Dactylis,  287,  294,  296. 
Dactylococcus,  51. 
Dsedalea,  166,  171. 


INDEX. 


601 


Dahlia,  569,  572. 
Daisy,  572. 
Dalbergia,  472. 
Dalbergieas,  472. 
Dalechampia,  434. 
Damasonium,  282. 
Dammara,  263. 
Dansea,  212. 
Dandelion,  571. 
Daphne,  449,  450. 
Darlingtonia,  409. 
Darwinia,  489. 
Dasycladaceae,  63. 
Dasycladus,  63. 
Dasyscypha,  135. 
Date-palm,  298,  299,  301, 

302. 

Date-plum,  511. 
Datisca,  477 
Datiscaceae,  477. 
Datura,  519,  520,  522. 
Dauceae,  496. 
Daucus,  134,  492,  496. 
Davallia,  214. 
Davilla,  413. 
Deadly  nightshade,  521. 
Dead-nettle,  538. 
Delesseria,  79,  80,  83. 
Delesseriacese,  83. 
Delphinieae,  383. 
Delphinium,  379,  383. 
Dendrobium,  332. 
Derbesia,  10,  60. 
Derbesiaceae,  47,  60. 
Dermatea,  116,  134. 
Derrnateaceae,  134. 
Dermateales,  134. 
DermatophytoD,  54. 
Dermocarpa,  25. 
Desmanthus,  475. 
Desmarestia,  71. 
Desrnarestiaceas,  71. 
Desmidiaceae,    10,   18,  21, 

42,  44,  48. 
Desmidium,  44. 
Desmodium,  466,  472. 
Deutzia,  455. 
Devil's-bit,  560. 
Dianthus,  364,  367. 
Diapensiaceae,  509. 
Diatoma,  19. 
Diatomacese,    10,    12,    14, 

19,  20,  21. 
Diatomeae,  1,  18. 
Diatomin,  18. 
Diatoms,  1. 
Diatrypaceaa,  130. 
Diatrype,  130. 
Dicentra,  395,  396,  397. 
Dichaenaceae,  132. 
Dichelyma,  197. 


Dichondreae,  516. 
Dichorisandra,  308. 
Dichospermum,  371. 
Dicksonia,  207,  215. 
Diclinous,  236. 
Dicliptera,  530. 
Dicotyledones,  3,  334. 
Dicranella,  196. 
Dicranum,  196. 
Dictamnus,  436. 
Dictyochaceaa,  15. 
Dictyonema,  176. 
Dictyosiphon,  71. 
Dictyosiphonaceae,  71. 
Dictyosphaerium,  51. 
Dictyostelium,  8. 
Dictyota,  76. 
Dictyotaceae,  76. 
Dictyotales,  1,  14,  76. 
Dicypellium,  392. 
Didiscus,  493. 
Didymium,  8. 
Dieffenbachia,  306. 
Dielytra,  395. 
Diervilla,  554,  556. 
Digitalis,  524,  525,  527. 
Digraphis,  295,  296. 
Dill,  496. 
Dillenia,  413. 
Dilleniaceae,  413. 
Dimorphanthus,  491. 
Dimorphochlamys,  481. 
Dinifera,  17. 
Dinobryinaceae,  15. 
Dinobryon,  15. 
Dinoflagellata,   1,   14,  16, 

17,  18,  21. 
Dinophysis,  17. 
Diodia,  550. 
Dioecious,  236. 
Dionasa,  408. 
Dioon,  254. 
Dioscorea,  322,  323. 
Dioscoreaceas,    276,    309, 

310,  322. 
Diosma,  436. 
Diosmeae,  436. 
Diospyrinae,  505,  510. 
Diospyros,  511. 
Diphtheria,  40. 
Diphjscium,  197. 
Diplarrhena,  321. 
Diplecolobese,  400. 
Diplococcus,  39. 
Diploderma,  78. 
Diplostemonous,  335,  336. 
Diplusodon,  483. 
Dipsacaceae,  549,  556,  558, 

559,  560,  569 

Dipsacales,  505,  556,  564. 
Dipsacus,  559,  560. 


Dipterocarpaceae,  415. 
Dipterocarpus,  415. 
Dipteryx,  472,  473. 
Discelium,  197. 
Discolicbenes,  142. 
Discomycetes,95, 116, 132. 
Discosporangium,  70. 
Disease,  32. 
Disinfection,  32. 
Dispora,  36. 
Distichium,  196. 
Doassansia,  110. 
Docidium,  '44. 
Dock,  359. 
Dodder,  5,  516,  517. 
Dodecatheon,  513. 
Dog's-tail,  294, 296. 
Dogwood,  490. 
Dolichos,  471. 
Dondia.  493. 
Dorema,  496,  498. 
Doronicum,  240,  572. 
Dorstenia,  131,  354. 
Dothideaceaa,  131. 
Double  Cocoa-nut  301. 
Doum-palm,  298,  301. 
Draba,  400. 
Dracaena,  274,  316. 
Dracaeneae,  316. 
Dracocephalum,  539. 
Dracunculus,  303,  305. 
Dragon's  blood,  301,  316. 
Dragon-tree,  316. 
Draparnaldia,  54. 
Drimys,  389. 
Drosera,  408. 
Droseraceae,  407. 
Drosophyllum,  408. 
Dryas,  458. 
Dryobalanops,  415. 
Dry-rot,  165,  166. 
Dry  yeast,  179. 
Duboisia,  522. 
Duck-weed,  307. 
Dudresnaya.  84. 
Dumontia,  84. 
Dumontiaceae,  84. 
Durio,  427. 
Durra,  296. 
Dwarf-elder,  553. 
Dwarf -male,  57. 
Dwarf-palm,  300. 
Dyer's  Weed,  407,  472. 

Earth-nut,  472. 
Earth-star,  174. 
Earth-tongue,  136. 
Eating-chestnut,  346. 
Ebeuacese,  511. 
Ebony,  511. 
Ecballium,  478,  480,  481. 


602 


INDEX. 


Eccremocarpus,  529. 

Echeveria,  451. 

Ecbinocactus,    375,    376, 
377. 

Echinodorus,  281. 

Echinops,  564,  570. 

Echinopsis,  376,  377. 

Echinospermum,  533. 

Echites,  544. 

Echium,    531,   532,    533, 
534,  535. 

Ectocarpacesa,.  70. 

Ectocarpus,  69,  70. 
,   Edelweiss,  573. 
I  Edwardsia,  469. 

Egg-cell,  13. 

Egg-fertilisation,  13. 

Egg-plant,  522. 

Ehretia,  533. 

Eichhornia,  316. 

Elachista,  71. 

Elachistaceee,  71. 

Eleeagnacese,  449. 

Elasaguus,  450. 

Elffiis,  301,  302. 

Elseocarpus,  425. 

Elaphorayces,  124. 

Elaphnum,  438. 

Elatereee,  192. 

Elaters,  189. 

Elatinaceee,  413. 

Elatine,  413. 

Elder,  156,  553,  555,  556. 

Elemi,  438. 

Elettaria,  326. 

Eleusine,  295. 

Elisma,  231,  282. 

Elm,  124, 165,  351. 

Elodea,  232. 

Elymus,  113,  296. 

Elyna,  286,  287. 

Embryo,  246,  247,  248. 

Embryo-sac,  241,  247. 

Emericella,  176. 

Emex,  360. 

Empetracese,  434. 

Empetrum,  434. 

Ernpleuruin,  436. 

Empusa,  101,  102. 

Enantioblastse,  277,  308. 

Encalypta,  197. 

Encephalartos,  254. 

Enchanter's    Nightshade, 
485. 

Encoeliacese,  70. 

Endocarpon,  142. 

Endomyces,  116,  117. 

Endophyllum,  147,  151. 

Endophytic  parasites,  85. 

Endosperm,  233,  246,  24f>, 
249. 


Endospermous,  249. 
Endospheera,  47,  51. 
Endospore,  89,  187. 
Kndosporous,  29. 
Endothecium,  186. 
Endozoic  Fungi,  85. 
Enhalus,  283. 
Entada,  473,  474. 
Enteromorpha,  53. 
Entoderma,  54. 
Entomophthora,  102. 
Entomophthoraceffi,  102. 
Entomophthorales,        95, 

102. 

Entyloma,  109,  111,  113. 
Enzyme,  32. 
Epacridaceee,  509. 
Epacris,  509- 
Ephebe,  139,  142. 
Ephedra,  271. 
Ephemerum,  195. 
Epibasal,  186. 
Epichloe,  125,  127. 
Epiclemmydia,  54. 
Epidendrese,  332. 
Epidendron,  332. 
Epigynum,  544. 
Epilobium,  484,  485. 
Epimedium,  390. 
Epipactis,  331,  333. 
Epipetalous,  336. 
Epiphyllum,  375,  377. 
Epiphytic  parasites,  85. 
Epipogon,  331. 
Epipyxis,  15. 
Episepalous,  335. 
Epithemia,  20,  21. 
Epizoic  Fungi,  85. 
Equisetaceaa,     202,     204, 

221,  234,    235,    236, 
240. 

Equisetinse,    2,    204,  221, 

225. 
Equisetum,  200,  206,  221, 

222,  224. 
Eragrostis,  294. 
Eranthenmm,  530. 
Eranthis,  379,  382. 
Ergot,  125,  127. 
Eria,  332. 

Erica,  507,  508. 
Ericaceae,  238,  505,  507. 
Ericese,  507. 
Erigeron,  573. 
Erinus,  525. 
Eriobotrya,  465. 
EriocaulacejE,  308,  309. 
Kriocaulon,  309. 
Eriodendron,  427. 
Eriophorum,  285,  286. 
Bodium,  419. 


Eruca,  402. 
Ervum,  470,  473. 
Eryngium,  493,  569. 
Erysiminee,  404. 
Erysimum,  402. 
Eryeiphaceae,  119. 
Erysiphe,  119,  120,  121. 
Erytnraaa,  542,  543. 
Erythrina,  471. 
Erythronium,  312. 
Erythrotrichia,  78. 
Erythroxylaceae,  442. 
Erythroxylon,  442. 
Escalloniacese,  454. 
Escalloniaa,  451. 
Eschalot,  312. 
Eschsholtzia,  393,  395. 
Esparto  grass,  296. 
Euactaaa,  379. 
Euaspergillus,  122. 
Euastrum,  44. 
Eucalyptus,  489. 
Eucharidium,  485. 
Eucharis,  317,  318. 
Euchlaana,  293. 
Eucomis,  312. 
Eudorina,  48,  50. 
Eugeissonia,  301. 
Eugenia,  488,  489. 
Euglena,  103. 
Eunotiea3,  21. 
Euonymus,  152,  444. 
Eupatoriess,  571,  572. 
Eupatorium,  569,  571. 
Euphacidiacese,  133. 
Euphorbia,  148,  151,  430, 

432,  433. 

Euphorbiacese,  430. 
Enphorbium,  434. 
Euphoria,  441. 
Euphrasia,  526. 
Eupodisceee,  21. 
Eiirhynchium,  197. 
Eurotium,  121,  122. 
Euryale,  386,  387,  388. 
Eusporangiatee,  202,  210, 

239. 

Euterpe,  301. 
Euthora,  83. 
Eutoca,  515. 
Evening  Prrnrose,  484. 
Everuia,  143. 
Evodia,  436. 
Evolvulus,  516. 
Exalbuminous,  249. 
Exidia,  156. 
Exoasci,  95,  115,  116. 
Exobasidium,  160,  161. 
Exochorda,  457. 
Exospore,  87,  187. 
Exostemma,  553. 


INDEX, 


603 


Exuviella,  16,  17,  18,  21. 

Eye-bright,  526. 
"  Eye-spot,"  10. 

Faba,  468,  470,  473. 
Fabiana,  521. 
Fabroniacese,  197. 
"  Fachel,"  284. 
Facultative  parasites,  84. 
Fagonia,  438. 
Fagus,  122,  347,  348. 
"Fairy-rings,"     86,    136, 

163,  168. 
Falcaria,  494. 
"  Fan,"  284. 
Fan-palm,  198. 
Farinose®,  b08. 
Fatsia,  491. 
Featber-grass,  294. 
Feather  palm,  298. 
Fedia,  557. 
Fegatella,  191. 
Fennel,  492,  495,  498. 
"  Ferment  of  wine,"  178. 
Fermentation,  32. 

Alcoholic,  97. 
Ferns,  2. 

Stem  of,  202,  204,  205. 
True,  204,  205. 
Water,  205. 
Various,  208. 
Ferraria,  321. 
Ferula,  496,  498. 
Fescue,  293,  296. 
Festuca,  293,  296. 
Festucese,  293. 
Fevillea,  478,  481. 
Ficaria,  334,  383. 
Ficus,  351,  354,  355,  356. 
Field-horsetail  '224. 
field-madder,  552. 
Field-thistle,  151. 
Fig-wort,  524. 
imago,  573. 
Filament,  238. 
Filbert,  345. 
Filices,  204,  205. 

Systematic  division   of, 

210. 
Filicinffi,  2,  204,  205,  234, 

236,  239. 
Fiori  di  fico,  355. 
Fiorin,  294. 
Fir,    124,   153,    155,    165, 

170,  263,  264,  265. 
Fir-cones,  163. 
Firneedle-rust,  152. 
Fissidens,  196. 
Fissidentaceee,  196. 
Fission-Algse.    1,   14,    22, 

29. 


Fission-Fungi,  26,  29. 
Fission-plants,  10. 
Fistulina,  166. 
Flag,  320. 
Flagellata,  48. 
Flagellatas,  15. 
Flax,  417. 
Flea-bane,  573. 
"  Fleur  de  vin,"  179. 
Floral-leaves,  235. 
Floridese,  9,  10,  77,  78. 
Flower,  235. 

Monocotyledonous,  276. 
Flowering-plants,  3,  249 
Flowering-rush,  281. 
Flowers-of-tan,  7. 
Fly-mould,  101. 
Fly-mushroom,  167,  171. 
Fly-trap,  408. 
Fceniculum,  495,  498. 
Foliage-leaves,  235. 
Fontinalacese,  197. 
Fontinalis,  197. 
Fool's-parsley,  495,  498. 
Foot,  -186. 
Fore-leaf,  275,  334. 
Forget-me-not,  334. 
Forskohlea,  353. 
Forsythia,  546,  547. 
Fossil  Gymnosperms,  271. 
Fothergilla,  455. 
Fourcroya,  318. 
Fovea,  231. 
Foxglove,  525. 
Fox-tail,  294,  293,  369. 
Fragaria,  468,  461. 
Fragilariese,  21. 
Franciscea,  521. 
Fraucoaceie,  454. 
Frangulinee,  443,  449,  451, 

490. 

Frankeniacese,  411. 
Frankincense,  438. 
Fraxinus,  122,  130,  546, 

547. 

French-bean,  473. 
French  Eose,  460. 
Freycinetia,  302. 
Fritillaria,  312,  313,  314. 
Frog-bit,  282. 
"  Fruit,"  91. 
Fruit,  249. 
Fruit-bearers,  91. 
Fruit-bodies,  91. 
"  Fruit-forms,"  91. 
Frullania,  191,  192. 
Frustule,  18. 
Frustulia,  20. 
Fucacese,  75. 
Fuchsia,  484,  485. 
Fucoideffi,  9. 


Fucus,  13,  73,  74,  75. 
Fuligo,  6,  8. 
Fumago,  124. 
Fumaria,  396,  397. 
Fumariaceae.  395. 
Fumitory,  395. 
Funaria,  182,  188,  197. 
Funariaceee,  197. 
Funckia,  312,  313,  314. 
Fundaments,  90. 
Fungi,  1,  4,  5,  8,  84. 
Fungi-galls,  85. 
Fungi  Imperfect!,  90. 
Fungus  chirurgorum,  164. 

laricis,  164. 
Funicle,  241. 
Furcellaria,  79,  84. 
Furze,  472. 
Fusicladium,  130. 


Gagea,  312. 
Gaillardia,  572. 
Galactodendron,  356. 
Galangal,  326. 
Galanthus,  317,  318. 
Galaxaura,  83. 
Galaxia,  321. 
Galbanum,  498. 
Galega,  470. 
Galeobdolon,  538. 
Galeopsis,  538,  540. 
Galinsoga,  572. 
Galipea,  437. 
Galium,  552,  553. 
Galphimia,  442. 
Gambier,  553. 
Gamboge,  414. 
Gambo-hetup,  430. 
Gametangium,  12. 
Gamete,  11. 
Gametophore,  183. 
Gametophyte,  181. 
Gamopetalte,  336. 
Garcinia,  414. 
Garden-cress,  405. 
Gardenia,  550. 
Gardeniea3,  550. 
Garidella,  383. 
Garlic,  312. 
Garrya,  491. 
Gasteria,  312. 
Gasterolichenes,  176. 
Gasteromycetes,  96,    145,, 

173.  . 

Gastonia,  491. 
Gaultheria,  508. 
Gaura,  485. 
Geaster,  174. 
Gelidiaceas,  83. 
Gelidium,  83,  84. 


G04 


INDEX. 


Genipa,  550. 
Genista,  471,  473. 
Genistese,  471. 
Gentian,  542. 
Gentiana,  542,  543. 
Gentianacese,  542. 
Gentianese,  542. 
Geoglossum,  136. 
Geonoma,  301. 
Georgiaceas,  197. 
Geraniacese,  418. 
Geranium,  419. 
Germ-pores,  93. 
Gesneria,  528. 
Gesnerinceae,     518,     526, 

528 

Gesnerieae,  528. 
Geum,  458,  460. 
Gigartina,  83,  84. 
Gigartinacese,  83. 
Gigartinales,  82,  83. 
Ginger,  326. 
Gilia,  515. 
Gillema,  457. 
Gills,  166. 
Ginkgo,   255,   257,  259, 

260,  272. 
Gipsy-wort,  539. 
Gladiolus,  321. 
Glandules,  329. 
Glasswort,  371. 
Glauciuni,  b94,  395. 
Glaucocystis,  22,  24. 
Glaux,  513. 
Gleba,  172. 
Glechoma,  539,  541. 
Gleditscbia,  468. 
Gleiclienia,  215. 
GleicheniaceaB,  215,  236. 
Glenodinium,  17. 
Globba,  326. 
Globe-thistle,  570. 
Globularia,  541. 
Globulariaceffi,  532,  541. 
Gloeocapsa,  24 
Gloeotrichia,  25. 
Gloiopeltis,  84. 
Gloiosiphoniaceae,  84. 
Gloxinia,  528. 
Glume,  v87. 
Gluimfloree,  277,  283. 
Glyceria,    113,   290,    294, 

296. 

Glyciue,  471. 
Glycyrrhiza,  470,  473. 
Glyptostrobus,  267. 
Gnaphalium,  569,  573. 
Gnetaceae,  3,  251,  271,  272. 
Gnete®,  252,  270. 
Gnetum,  271. 
Gnidia,  449. 


Gnomonia,  130. 
Gnomoniacpse,  130. 
Goafs-beard,  571. 
Godetia,  485. 
Godlewskia,  25. 
Golden- currant,  455. 
Golden-rod,  573. 
Golden  Saxifrage,  452. 
Goldfussia,  530. 
Gold-of-pleasure,  401. 
Gomontia,  58. 
Gomontiacese,  47,  58. 
Gomphonema,  19. 
Gornphonemese,  20,  21. 
Gomphosphseria,  24. 
Gomphrena,  368,  369. 
Gonatozygon,  44. 
Gongrosira,  54. 
Gonidia,  138. 
Gonimoblast,  82. 
Goniotrichaceae,  78. 
Goniotrichum,  78. 
Gonium,  48. 
Gonococcus,  39. 
Gonolobus,  546. 
Goodenia,  564. 
Goodeniaceas,  563. 
Gooseberry,  455. 
Goosefoot,  369. 
Gossypieaa,  427. 
Gossypium,  427,  429,  430. 
Gouania,  448. 
"Gourds,"  481. 
Gout-weed,  494. 
Gracilaria,  83. 
"Grains     of     Paradise,' 

390. 

Gramineas,  277, 283, 287. 
Grape-disease.  121. 
Graphiola,  110. 
Graphis,  140,  142. 
Grasses,  287. 
Grass-flower,  290,  291. 
Grass-fruit,  292 
Grass  of  Parnassus,  453. 
Grass-wrack,  279. 
Grateloupiaceee,  84. 
Gratiola,  525,  527. 
Green  Algae,  1,  14. 
"  Greenheart,"  393. 
Grevillea,  450. 
Griffithsia,  84. 
Grimmia,  197. 
Grimmiaceae,  197. 
Gronovia,  476. 
Ground  Ivy,  539. 
Groundsel,  153,  572. 
Gruinales,  416. 
Guaiacum,  438. 
Guano,  20. 
Guava,  489. 


Guava-rum.  490. 
Guazuma,  422. 
Guelder-rose,  455,  555. 
Guepinia,  159. 
Guinea-corn,  296. 
Guinea  Pepper-plant,  521 
Guizotia,  574. 
Gulf-weed,  75. 
Gum-arabic,  475. 
Gum-benzoin,  511. 
Gum-tragacanth,  473. 
Gum-trees,  490. 
Gunnera,    25,    482,    485 

486. 

Guttapercha,  511. 
Guttiferae,  414. 
Gynmadenia,  332. 
Gymnoascacese,  119. 
Gymnoascales,    95,    116. 

118. 

Gymnoascus,  119. 
Gymnodinium,  17. 
Gymnogramme,  214. 
Gymnospermae,     2,    234, 

239,  250,  251. 
Gymnosperms,  244,  246. 

Fossil,  271. 
Gymnosporanginm,     146, 

147,  151,  154. 
Gymnosporese,  82. 
Gymnostomum,  196. 
Gymnozyga,  42,  44. 
Gynandraa,  278,  328. 
Gynandropsis,  405,  406. 
Gynerium,  294.  296. 
Gynceceum,  237. 
Gynophore,  367. 
Gynostemium,  329. 
Gysophila,  368. 


Habenaria,  332. 
Hablitzia,  370. 
Babrothamnus,  522. 
Hacquetia,  493. 
Heemanthus,  317,  318. 
Hagmatoxylori,  467,  4<'i8. 
Haamodoraceas,  320. 
HaBmodorum,  320. 
Hagenia,  460. 
Hair-grass,  294. 
Hakea,  450. 
tfalesia,  511. 
Flalianthus,  366. 
Halidrys,  73,  75. 
Halimeda,  62,  63. 
Halimus,  371. 
Halophila,  283. 
Haloragidaceaa,  482,  485, 

486. 
Haloragis,  486. 


INDEX. 


605 


Halyrnenia,  84. 
Hamamelidacese,  455. 
Hamamelis,  455. 
Hamelia,  550. 
Hankornia,  544. 
Hapalosiphon,  26. 
Haplomitriuni,  192. 
Haplospora,  72. 
Haptera,  4,  10. 
Hard-fern,  214. 
Hare's-ear,  494. 
Hart's-tongue,  214. 
"  Harzsticken,"  169. 
Haschisch,  358. 
Hassalia,  26. 
Haustoria,  86. 
Hawkbit,  571. 
Hawksbeard,  571 . 
Hawthorn,  465. 
Hay-bacillus,  37,  38.  39. 
Hazel,  526. 
Hazel-nut,  343. 
Heal-all,  539. 
Heath,  507. 
Hebenstretia,  54l. 
Hechtia,  319. 
Hedera,  491. 
Hedge-mustard,  402. 
Hedge-parsley,  497. 
Hedwigia,  197. 
Hedycarya,  389. 
Hedychium,  326. 
Hedyosmum,  363. 
Hedysareae,  472. 
Hedysarum,  472,  473. 
Helenium,  572. 
Heleocharis,  285. 
Helianthese,  572,  573. 
Helianthemum,  412. 
Helianthus,  569,  572,  574. 
Helichrysum,  573. 
Heliconia,  325. 
Helicon!®,  325. 
Helicophyllum,  303. 
Helicteres,  422. 
Heliophilinae,  404. 
Heliotropieae,  533. 
Heliotropium,  533,  535. 
Hellebore,  382. 
Helleboreae,  381. 
Helleborus,  379, 380, 382. 
Helminthocladiacese,  83. 
Helobieae,  277,  278. 
Helosciadium,  494. 
Helosis,  504. 
Helotiacese,  135. 
Helotium,  135. 
Helvella,  136. 
Helvellacese,  136. 
Helvellales,  95, 116, 136. 
Helwingia,  491. 


Hemerocallidece,  312. 
Hemerocallis,    312,     313, 

314. 

Hemiasci,  95,  108. 
Hemibasidii,  95, 108,  109. 
Hemicblamydeous,  257. 
Hemileia,  155. 
Hemlock,  494,  498. 
Hemp,  356,  529. 
Hemp-agrimony,  571. 
Hemp- nettle,  538. 
Henbane,  521. 
Henriquezia,  549. 
Hepaticse,  2,  188. 
Hepialus,  128. 
Heracleum,  492,  496. 
Herb-Paris,  314. 
Heritiera,  422. 
Herman  nia,  422. 
Hermaphrodite,  236. 
Herminium,  332. 
Hermodactylus,  321. 
Hernandia,  392. 
Herniaria,  365,  367. 
Herpesti?,  525. 
Herposteiron,  54. 
Herpotrichia,  129. 
Hesperideas,  404. 
Hesperidinas,  404. 
Hesperis,  400,  402. 
Heteranthera,  316. 
Heterobasidion,  145, 165. 
Heterocystese,  24. 
Heterocysts,  22. 
Hetercecious,  148. 
Heteromerous,  138. 
Heteropteris,  442. 
Heterosphaeria,  116, 133. 
Heterosphaariaceee,  133. 
Heterosporous     Vascular 

Cryptogams,  200. 
Heterotoma,  563. 
Heuchera,  452. 
Hibiscus,  427,  430. 
Hickory,  350. 
Hieracium,  571. 
Hierochloa,  295,  296. 
Higher  Fungi,  95,  114. 
Hiluin,  243. 
Himanthalia,  75. 
Hiniantidiurn,  20. 
Hip,  459,  460. 
Hippocrateaceas,  444. 
Hippocrepis,  472. 
Hippomane,  434. 
Hippophae,  450. 
Hippuris,  486. 
"  Hochblatt,"  235. 
Hog's-fennel,  496. 
HolboeUia,  390. 
Holcus,  294,  296. 


Holly,  444. 
Hollyhock,  151,  430. 
Holochlamydeous,  256. 
Holosteum,  366. 
Homalia,  197. 
Homalotheciuro,  197. 
Homocysteffi,  24. 
Homoiomerous,  138. 
Honckenya,  366. 
Honesty,  400. 
Honey- dew,  126. 
Honey-leaves,  379. 
Honeysuckle,  553,  554 
Hookeriaceaa,  197. 
Hop,  124,  356,  357. 
Hopea,  415. 
Hordeee,  295. 
Hordeum,  291,  296. 
Horehound,  538. 
Hormidium,  54. 
Hormogonia,  10,  24. 
Hornbeam,  157,  343,  344. 
Horned  Pond-weed,  279. 
Horn-nut,  485. 
Horn-poppy,  395. 
Horn-wort,  388. 
Horse-bean,  470,  473. 
Horse-chestnut,  440. 
Horse-radish,  400,  405. 
Horsetails,  2,  204,  221. 
Hosta,  312. 
Hoteia,  452. 
Hottonia,  512. 
Hound's-tongue,  5D3. 
House-leek,  452. 
Houttuynia,  359,  362. 
Hoya,  546. 
Humiriacese,  421. 
Humulus,  121,  357,  358. 
Hura,  432. 
Hyaciutheaa,  312. 
Hyacinthus,  31'2,  313,  314. 
Hyalotheca,  42,  44. 
Hydnaceae,  162. 
Hydnophytum,  550,  553. 
Hydnora,  504. 
Hydnum,  162. 
Hydra,  9. 
Hydrangea,  455. 
Hydrangeaceas,  455. 
Hydrastin,  385. 
Hydrastis,  381. 
Bydrilla,  283. 
Bydrocharis,  282. 
Hydrocharitaceae,        278, 

282. 

Hydrocleis,  281. 
Hydrocotyle,  491,  493. 
Hydrocotyleaa,  493. 
Hydrodictyaceaa,  47,  51. 
Hydrodictyon,  9,  52. 


606 


INDEX. 


Hyclrolea,  515. 

Ilex,  444. 

Jateorhiza,  390. 

Byolrophyllaceee,  515. 

Illecebrum,  3G7. 

Jatropha,  431. 

Hydropteridess,  205,  215, 

Illiciete,  389. 

Jequirty,  470. 

239. 

Illicium,  389. 

Jerusalem  -  Articholio, 

Hydruraceas,  16. 

Impatiens,  421. 

572,  574. 

Hydrurus,  16. 

Irnperatoria,  496,  498. 

Jonquil,  318. 

Hygrophorei,  172. 

Incense,  438. 

Judas'-ear,  156. 

Hygrophorus,  172. 

Indian-corn,  293. 

Judas-tree,  468. 

Hylocomium,  197. 

Indian-cress,  4'20. 

Juglandaceee,  337,  349. 

Hymensea,  468. 

Indigo,  470,  473. 

Juglandiflorje,  349. 

Hymenium,  88. 

Indigofera,  470,  473. 

Juglans,  349,  350.  • 

Hymeiiogaster,   174,  175, 

Indusium,  210. 

Juncaceaa,  277,  283,  284, 

176. 

Inflorescence     of     Palm, 

291. 

Hymenogastracese,  176. 

299. 

Juncaginaoeffi,  278. 

Hymenolichenes,  176. 

Infusoria,  9. 

Juncus,  283,  284. 

Hymenomycetes,  96,  145, 

Inga,  473,  475. 

Jungermannia,  191,  192. 

159. 

Integuments,  242. 

Jungermannieee,  191. 

Hymenophore,  159. 

Inula,  569,  573,  574. 

Juniper,  259,  268,  269. 

Hymenophyllacese,      206, 

Inulin,  574. 

Juniperus,  151,   152,  241, 

210,  215. 

Involucre,  189. 

268,  269. 

Hymenophyllum,  215. 

Involution-forms,  36. 

Jurinea,  570. 

Hyoscyainine,  522. 

lonidiurn,  4  10. 

Jussia3a,  485. 

Hyoscyamus,    518,     519, 

Ipecacuanha,  553. 

Justicia,  530. 

520,  521,  522,  523. 

Ipomaea,  515,  517. 

Jute,  425. 

Hypecoum,  395,  396. 

Iridaceaa,  277,  310,  320. 

Hypericaceee,  413. 

Iris,  276,  291,  320,  321. 

Kalancboe,  451. 

Hypericum,  413,  414. 

Irish-moss,  84. 

Kale,  403,  405. 

Hypha,  85. 

Iron-bacteria,  33. 

Kalmia,  509. 

Hyphse-like  threads,  9. 

Iron-wood,  339,  511. 

Ksempferia,  325,  326. 

Hypbame,  298,  301. 

Irpex,  163. 

Kaniala,  434. 

Hypholoma,  171. 

Isactis,  25. 

Kaulfussia,  212. 

Hypnacese,  197. 

Isaria,  127,  128. 

Kefir-  grains,  36. 

Hypnum,  47,  196,  197. 

I  satis,  403,  404. 

Kelp,  76. 

Hypobasal,  186. 

Isnardia,  485. 

Kerria,  457,  460. 

Hypochseris,  571. 

Isoetaceee,  230. 

Kidney-bean,  471,  473. 

H^pochnus,  161. 

Isoetes,    200,    202,    204, 

Kielmeyera,  415. 

Hypocreacea3,  125. 

228,  230,  245. 

"  King   Charles    and   the 

Hypocreales,  125. 

Isogamous      fertilisation, 

Oak,"  207. 

Hypoderma,  132. 

11. 

Kingia,  312. 

Hypodermaceas,  132. 

Isolepis,  287. 

Kino,  473. 

Hypomyces,  l'J5. 

Isonandra,  511. 

Kitaibelia,  429. 

Hyporhodius,  171. 

Isopyrum,  382. 

Knap-weed,  570. 

Hypothecium,  132. 

Isosporotis    Vascular 

Knapwell,  367. 

Hypoxidese,  317. 

Cryptogams,  200. 

Knautia,  560. 

Hypoxis,  318. 

Isothecium,  197. 

Knot-grass,  359. 

Hypoxylon,  131. 

Isotorna,  563. 

Knowltonia,  379. 

Hypsophyllarv  If  aves,  235. 

Iw,  491. 

Kobresia,  287. 

Hyssop,  540,  541. 

Ixia,  321. 

Kochia,  371. 

Hyssopus,  540,  541. 

Ixora,  550. 

Koeleria,  294. 

Hysteriaceaa,  132. 

Koelreuteria.  441. 

Hysteriales,  95,  116,  132. 

Jacaranda,  529. 

Koenigia,  361. 

Hysterium,  132. 

Jack,  356. 

Kohlrabi,  405. 

Hysterophyta,  498. 

Jacquinia,  513. 

Krameria,  468. 

Jalap,  517. 

Kramerieffi,  468. 

Iberis,  398,  400,  401. 

Jambosa,  488. 

Koso-tree,  460. 

Icacinaceee,  439. 

Japanese  wax,  439. 

Iceland-lichen,  142. 

Jasione,  541,  561,  562. 

Labellum,  277,  323,  325. 

Iceland-moss,  143. 

Jasminaceee,      541,     542, 

Labiate,    515,    532,   535, 

Ice-plant,  375. 

547. 

536. 

Icica,  438. 

Jasmine,  547. 

Labiate-flowered,  567,  570. 

Ignatius-beans,  546. 

Jasmin  urn,  547. 

Laburnum,  472,  473. 

INDEX. 


C07 


Labyrinth  Fungus,  166. 
Lace-tree,  449. 
Lacmus,  142. 
Lactarius.  171. 
Lactoridaceae,  362. 
Lactoris,  3^2. 
Lactuca,  571,  574. 
Ladanum,  412. 
Ladenbergia,  550,  553. 
Ladies-mantle,  460. 
Lady-fern,  213. 
Lady's-finger,  471. 
Lealia,  332. 
Lageuandra,  306. 
Lagenaria,  479,  481. 
Lagenedium,  104. 
Lagerstrcemia,  483. 
Lagetta,  449. 
Lagoecia,  494. 
Laguncularia,  487. 
Lagurus,  296. 
Lamella,  166. 
Laminaria,  71. 
Liminariaceaa,  71. 
Lamium,   536,    538,   540, 

541. 

Lamprothamnus,  67. 
Landolphia,  544. 
Langsdorffia,  504. 
Lantana,  535. 
Lappa,  570,  574. 
Lapsana,  570. 
Larch,  266,  267. 
Larch-canker,  135. 
Larch-fungus,  164. 
Lardizabalaceae,  390. 
Larix,  266. 
Larkspur,  383. 
Larrea,  438. 
Laserpitium,  497. 
Lasiandra,  484. 
Latania,  301. 
Lathreea,  525,  526,  528. 
Lathyrus,  470,  473. 
Lattice-rust,  147. 
Laudatea,  176. 
Lauraceae,  238,  391.  449. 
Laurus,  161,  391,  392,  39  *. 
Lavandula,  536,  540,  541. 
Lavatera,  428,  430. 
Lavender,  540. 

Oil  of,  541. 
Lawsonia,  483. 
Leafy-mosses,  188. 
Leathesia,  71. 
Leaven,  179. 
Lecanora,  140,  142. 
Lechenaultia,  564. 
Lecidea,  142. 
Lecythideae,  489. 
Lecythis,  489. 

W.  B. 


Ledum,  153,  509. 
Leea,  445. 
Leek,  312. 
Leersia,  291,  293. 
Legumiuosas,  466. 
Legume,  466. 
Lejolisia,  81. 
Lemanea,  80,  82. 
Lemaneacese  82. 
Lemna,  25,  47,  307. 
Lemnaceae,  307. 
Lemon,  438. 
Lentil,  470,  473. 
Lentinus,  171. 
Leontice,  390. . 
Leontodou,  568,  571. 
Leontopocliuni,  593. 
Leonurus,  538. 
Lepidiinae,  404. 
Lepidium,  400,  401. 
Lepidocaryinas,  301. 
Lepidodendracese,  233. 
Lepidozia,  192. 
Lepiota,  171. 
Leptobryum,  197. 
Leptogium,  140,  142. 
Leptomitus,  108. 
Leptopleura,  387. 
Leptopuccinia,  151. 
Leptosiphon,  515. 
Leptospermeas,  489. 
Leptospermum,  489. 
Leptosporangiataa.       202, 

210,  212,  2b9. 
Leptotbrix,  26,  H3,  35,  38. 
Leptotriclium.  196. 
Lepturus,  295. 
Lescuraea,  197. 
Leskea,  197. 
Leskeaceae,  197. 
Lessonia,  72. 
Lettuce,  571   574. 
Lencobryaceae,  196. 
Leucobryum,  192,  196. 
Leucodon,  197. 
Leucojum,  317,  318. 
Leuconostoc,  28,  29,  35. 
Levisticum,  496,  498. 
Liagora,  83. 
Libanotis,  495. 
Libocedrus,  269. 
Lichen,  4,  8. 
Lichen-forming    Ascomy- 

cetes,  116,  136. 
Basidiomycetes,  176. 
Lichenin,  142. 
Lichina,  142. 
Licmophoreae,  21. 
Lignum  Vitas,  438. 
Ligularia,  572. 
Ligulate-flowered,  567. 


Ligule,  283. 
Liguliflorae,  570. 
Ligustrum,  547. 
Lilac,  547. 
Lilasa,  279. 

Liliaceae,  274,  309,  311. 
Lilies,  311,  314. 
Liliifiorae,  278,  309. 
Lilium,    245,     312,    313,. 

314. 

Lily  of  the  Valley,  314. 
Lime,  165. 
Limnanthaceae,  421. 
Limnanthemum,  543. 
Limnanthes,  421. 
Limnocharis,  281. 
Limodorum,  331. 
Limonia,  437. 
Limosella,  525. 
Linaceae,  417. 
Linaria,  523,  525,  527. 
Lindera,  393. 
Ling,  507. 
Linnaea,  555. 
Linnaeeae,  555. 
Linociera,  547. 
Linseed,  418. 
Linum,  417,  418. 
Liparis,  332. 
Lippia,  535. 
Liquidambar,  455. 
Liquorice,  470,  473. 
Liriodendron,  388. 
Listera,  331. 
Litchi,  441. 
Lithoderma,  71. 
Lithodermataceae,  71. 
Lithophyllum.  84. 
Lithospermum,  533 
Lithothamnion,  80,  84. 
Littorella,  530,  531. 
Liverworts,  2,  181,  188. 
Livistona,  298,  299,  300r 

302. 

Lloydia,  312. 
Loasaceae,  476. 
Lobelia,  562,  563. 
Lobeliaceae,  335,  562. 
Lobeline,  503. 
Loclmera,  544. 
Locusts,  468. 
Lodicules,  288,  291. 
Lodoicea,  30 1 . 
Loganiaceae,  542, 546,  549- 
Logwood,  468. 
Loiseleuria,  509. 
Lolium,  295.  296. 
Lomandra,  312. 
Lomaria,  214. 
Lornentaceae,  403. 
Lomentaria,  83. 
RR 


608 


INDEX. 


Lonicera,  553,  554,  556. 

Macrosporangium,       241, 

Maranta,  327. 

Lonicereffi,  549,  553. 

243. 

Marantaceas.  277,  327. 

Loug  -pepper,  363. 

Macrospore,  200,  242,  243, 

Marasmiei,  J71. 

Loose-strife,  482. 

245,  246. 

Marasmius,  168.  171. 

Lopezia,  484,  485. 

Macrocystis,  72. 

Marattta,  212. 

Lophiostomacese,  130. 

Macrozamia,  254. 

Marnttiacese,  209,  210,  212, 

Lophocolea,  192. 

Madder,  552,  553. 

236. 

Lophodermium,  132. 

Madia,  574. 

Marcgraviace*.  415. 

Lophospermum,  525. 

Madotheca,  192. 

Marcbanlia,  181,  183,  184, 

Loquat,  465. 

Mad-wort,  534. 

190. 

Lorauthaceee,  501. 

Maesa,  513. 

Marchantiaceae,  190. 

Loranthoidese,  501. 

Magnolia,  389. 

Marchantiete.  190. 

Lorantlius,  504. 

Magnoliaceee,  388. 

Mare's-tail.  486. 

Loteae,  471. 

Magnoliese,  388. 

Marigold,  572. 

Lotus,  471. 

Mahernia,  422. 

Marjoram,  539,  541. 

Louse-wort,  526. 

Mahogany,  436. 

Marrow,  480. 

Love-in-the-mist,  382. 

Mahonia,  149,  390. 

Marrnbium,  538. 

Lucerne,  473,  529. 

Maiden-hair,  206,  213. 

Marsilia,   216,    217,   219, 

Lucuma,  511. 

Maize,  289,  293,  296. 

220,  245. 

Luehea,  424,  425. 

Maize-blight,  113. 

Marsiliaceaj,  210,  218,  239. 

Luffa,  481. 

Majanthemutu,  309,  314. 

Marsh  Cinquefoil,  458. 

Lunaria,  400,  401. 

Malachium,  366. 

Marsh-marigold,  382. 

Lung-Lichen,  143. 

Malachra,  428. 

Martynia,  529. 

Lung-wort,  533. 

Malaxis,  332. 

Masdevallia,  332. 

Lunularia,  191. 

Malcolmiinae,  404. 

Massariaceee,  130. 

Lupin,  472. 

Male-Fern,  214. 

Massulse,  331. 

Lupinus,  472. 

Mallow,  425. 

"  Mast,"  347. 

Lnzu'a.  283,  284. 

Malope,  429,  430. 

Mastic.  439. 

Lychnis,  365,  367. 

Malopese,  428. 

Mastigobryum,  192. 

Lycbnothamnus,  67. 

Malpighiacese,  442. 

Mastigocoleus,  24. 

Lycium,  521. 

Malpighia,  422. 

Mate,  445. 

Lycogala,  6,  8. 

Malt,  296. 

Matico.  363. 

Lycoperdacese,  174. 

Malus,  152,  463,  464,  465. 

Matricaria,  572,  574. 

Lycoperdon,  174. 

Malva,  426,  428,  429,  430. 

Matthiola,  400,  402. 

Lycopersicum,  521,  522. 

vTalvaceee,  425. 

Maurandia,  525. 

Lycopodiaceae,  202,  226. 

Malveee,  428. 

Mauritia,  301. 

Lycopodiese,  205,  226. 

Malvaviscus,  428. 

Maxillaria,  332. 

Lycopodinse,   2,  205,  22«, 

Mamme,  355. 

May,  465. 

228,    234,    235,    236, 

Mammea,  414. 

Mayacaceae,  308. 

240. 

Mammilles,  377. 

Maydeaj,  293. 

Lycopodium,     200,      226, 

Mammillaria,  375,  377. 

Meadow-grass,   151,    294, 

227,  228,  233. 

Mammoni,  355. 

296. 

Lycopsis,  534. 

Mancinil-tree,  432. 

Meadow  Eue,  385. 

Lycopus,  536,  539. 

Mandragora,  522. 

Meadow-sweet,  457. 

L.vgeum,  293. 

Mandrake,  522. 

Mecca-balsam,  438. 

Lygodium,  215. 

Manettia,  550. 

Mecouopsis,  395. 

Lynie-grass,  296. 

Mangifera,  439. 

Medicago,  471,  473. 

Lyngbya,  24. 

Manglesia,  450. 

Medick,  471,  473. 

Lyngbyaceas,  22,  24. 

Mango,  439. 

Medinilla,  484. 

Lyonia,  508. 

Mangold,  369,  372. 

Medlar,  465. 

Lysimacbia.  47,  151,  513. 

Mangosteen,  414. 

Meesea,  197. 

Lysipoma,  563. 

Mangrove,  486. 

Megaearpsea,  400,  401. 

Lytbraceae,  482. 

Manibot,  431,  434. 

Melaleuca,  489. 

Ly  thrum,  482,  483. 

Manilla  Hemp,  325. 

Melampodium,  572. 

Maniok,  434. 

Melampsora,  147,  152,153. 

'  Manna,"  547. 

Melampsorella.  147. 

Maba,  511. 

Manna  Ash,  546,  547. 

Melampyrum,  526. 

Macbserium,  472. 

Manna-grass,  296. 

Meianconidacese,  130. 

"  Mace,"  393. 

Manna-lichen,  142. 

Melandriurn,  367. 

Macleya,  395. 

Mannit,  72. 

Melanogaster,  176. 

Madura,  354,  356. 

Maple,  442. 

Melanommacese,  30» 

INDEX. 


609 


Melanoselinura,  497. 
Melanosinapis,  402. 
Melanospora,  125. 
Melanoxylon,  468. 
Melanthium,  310. 
Melastomacese,  483. 
Meliaceee,  435. 
Melianthaceae,  440. 
Melianthus,  440. 
Melica,  287,  290,  294. 
Melilotus,  466,  470,  471. 
Melinophyl,  18. 
Melissa,  540,  541. 
Melobesia,  80,  84. 
Melocactus,  375,  377. 
Melocbia,  422. 
Melogrammatacese,  130. 
Melon,  481. 
Melosira,  19. 
Melosirese,  21. 
Menispermaceee,  390. 
Menispermum,  390. 
Mentha,  47,  536,  539,  541. 
Menthol,  541. 
Mentzelia,  476. 
Menyantbeae,  542,  543. 
Menyanthes,  240,  543,  550. 
Menziesia,  509. 
Mercurialis.  431,  434. 
Mercury,  431. 
Merendera,  310. 
Mericarp,  492. 
Meridieffi,  21. 
Merismopedium,  10,  24. 
Merismopedium  form,  27. 
Mertensia,  533. 
Merulius,  166. 
Mesembriantheinese,  375. 
Mesembrianthemum,  375. 
Mesocarpaceas,  46. 
Mesomycetes,  1,  95,  108. 
Mesotamium,  43,  44. 
Mespilus,  463,  465. 
Metaxenous,  148. 
Metrosideros,  489. 
Metroxylon,  298.  301. 
Metzgeria,  191,  192. 
Metzleria,  563. 
Meum,  495. 
Michauxia,  562. 
Miconia,  484. 
Micrasterias,  44. 
Microcachrys,     255,    260, 

261.     ' 

Microchffite,  26. 
Microchloa,  295. 
Micrococcus,  26,  35,  38. 
Microcoleus,  22,  24. 
Microconidia,  89. 
Microcycas,  254. 
Microdictyon,  62. 


Microglena,  15. 
Micropyle,  242. 
Microsphsera,  121. 
Microspira-comma,  40. 
Microspora,  54. 
MicrosporangiH,  2i7,  240. 
Microspore,  200,  244. 
Microtea.  372. 
Mignonette.  406. 
Mikania,  571. 
Mildews,  119, 122. 
Milfoil,  572. 
M  ilium,  294. 
Milk-thistle,  570. 
Milk-vetch,  470. 
Milk-wort,  443. 
Millet,  296. 
Mimosa,  473. 
Mimosaceffi,  466,  473. 
Mimulus,  525,  526,  527. 
Mimusops,  511. 
Mint,  539. 
Mirabilis,  374. 
Mistletoe  501. 
Mitella.  452. 
Mitromyces,  173. 
Mitrula,  136,  159. 
Mnium,  197. 

Mock  Orange-blossom, 455. 
Modiola,  427. 
Moehringia,  366. 
Mohria,  215. 
Molinia,  151,  294. 
Mollinedia,  389. 
Mollisia,  135. 
Mollisiaceee,  135. 
Mollugo,  375. 
Momordica,  481. 
Monacanthus,  333. 
Monangic,  243. 
Monarda,  540,  541. 
Monardeae,  540. 
Money-wort,  513. 
Monimia,  389. 
Monimiaceee,  389. 
Monkshood,  383. 
Monoblepharis,  102,  108. 
Monocotyledones,  3,  273, 

274,  276. 
Monocotyledonous  flower, 

276. 

Monoecious,  236. 
Monostroma,  53. 
Monotropa,  334,  506,  507. 
Monstera,  303,  305,  307. 
Montia,  373. 
Moonwort,  211. 
Moraceee,  351,  353. 
Morsea,  321. 
Morchella,  136. 
More*,  354. 


Morell,  136 
Moricandiinte,  404. 
Morina,  560. 
Murinda,  549. 
Mortierellaceee,  100. 
Mortierella,  100. 
Morus,  361,  354. 
Moschatel,  453. 
Moss,  182.  •          '  .. 
"  Moss-tiower,"  183. 
Moss-fruit,  186. 
Moss-rose,  460. 
Mosses,    1,    2,   181,    183, 

192,  234. 
Mougeotia,  46. 
Moulds,  31,  94.  122. 
Mountain-ash,  465. 
Mountain-meal,  20. 
Mountain-pine,  2G6. 
Mouse-tail,  383. 
Mucor,  97,  98,  99. 
Mucoracese,  96. 
"  Mucor-yeast,"  97. 
Mucro,  257. 
Mucuna,  471. 
Mud-wort,  525. 
Muehleiibeckia,  360. 
Mulberry.  353,  356. 
Mullein,  523. 
Murracytaceee,  15. 
Musa,  324,  325. 
Musaceee,  277,  323. 
"  Muscardine,"  128. 
Muscari,  312,  314. 
Musci,  2. 

frondosi,  188,  192. 
Muscineffi,  1,  181. 
Musese,  325. 

Mushroom,  159,  166,  168. 
Musk-rose,  460. 
Mutisieee,  570. 
Myauthus,  333. 
Mycelium,  85. 
Mycena,  171. 
Mycoidea,  8,  54. 
Mycoideacea3,  47,  54. 
Mycomycetes,  1,  95,  114. 
Mycorhiza,  124,  175,  180, 

506. 

Mycosiphonales,  95,  104. 
Myosotis,  533,  534.  535. 
Myosurus,  379,  380,  383, 

384. 

Myrcia,  488. 
Myrica,  350. 
Myricaceee,  337,  350. 
Myricaria,  411,  412. 
Myriophyllum,  486. 
Myriotrichia,  71. 
Myriotrichiaceee,  71. 
Myristica,  392,  393. 


610 


INDEX. 


Myristicaceee,  393. 
Myrmecodia,  550,  553. 
Myroxylon,  473. 
Myrrh,  438. 
Myrrha,  438. 
Myrrhis,  495,  498. 
Myrsinaceae,  513. 
Myrsine,  513. 
Myrtaceae,  487. 
Myrtese,  488. 
Myrtiflorae,  451,  482. 
Myrtle,  487,  488. 
Myrtus,  488,  489. 
Myxamceba,  6. 
Myxogasteres,  5. 
Myxomycetes,  1,  4,  5. 
Myxophycese,  22. 
Myzodendroo,  500,  501. 

Naccaria,  83. 
Naegelia,  528. 
Najadacea3,  278,  281. 
Najas,  281. 
Nandina,  390. 
Narcissus,  316,  317,  318. 
Nardostachys,  557,  558. 
Nardus,  291,  295,  558. 
Narthecium,  310. 
Narthex,  496. 
Nasturtium,  400,  402,  420. 
Navicula,  19. 
Naviculeaa,  20,  21. 
Neck-canal-cells,  184. 
Neckera,  197. 
Neckeraceas,  197. 
Nectandra,  392,  393. 
Nectria,  116,  125,  127. 
Neea,  374. 
Negundo,  441,  412. 
Nelumbo,  386. 
Nelumbonese,  386. 
Nunaliou,  81. 
Netnalionales,  82. 
Nemastomacese,  84. 
Nemesia,  525 
Netnopbila,  515. 
Neomeris,  63. 
Neottia,  5,  331. 
Neottieae,  331. 
Neovossia,  111. 
Nepenthaceae,  408,  409. 
Nepenthes,  409. 
Nepeta,  536,  539. 
Nepeteae,  539. 
Nephelium,  441. 
Nephrolepis,  214. 
Nephroselrnls,  15. 
Nerium,  544. 
Nfissea,  483. 
Neslia,  403. 
Nest-fungi,  176. 


Nettle,  351,  352,  353. 
Neuradeae,  457. 
Neuwiedia,  32:^. 
Nicandra,  519,  5 '22. 
Nicotiana,  520,  522. 
Nicotine,  522. 
Nidularia,  176. 
Nidulariaceae,  176. 
Nierembergia,  521. 
Nigella,  379,  380,  382. 
Nightsbade,  521. 
Nigritella,  332. 
Nile-lily,  305. 
Nipa,  301. 
Nipplewort,  570. 
Nitella,  65. 
Nitelleae,  67. 
Nitraria,  438. 
Nitrifying  Bacteria,  5. 
Nitzcbieae,  21. 
Noble  Pine,  264. 
Noctiluca,  17. 
Nodularia,  25. 
Nolana,  522. 
Nolanaceae,  518,  522. 
Noli-me-tangere,  4.21. 
Nonnea,  533. 
Nonsexual     reproduction, 

10. 
Nostoc,  22,  23,  25,  27,  29, 

138,  486. 

Nostocaceae,  22,  24,  25. 
Nostocopsis,  26. 
NothofftKiis,  347,  348,  501. 
Notorhizae,  400. 
Nucellus,   235,   241,   243, 

247. 

Nuculifersa.  505,  515,  531. 
Nucumentaceae,  403. 
Nullipora,  84. 
Nuphar,  387. 
Nutmegs,  393. 
Nutritive-tissue,  248. 
Nux  vomica,  546. 
Nyctagiuiaceae,  373. 
Nyctalis,  172. 
Nyctantb.es,  547. 
Nycterinia,  525,  526. 
Nymphaaa,  387,  388. 
Nympbaeaceae,  385. 
Nymphaeeae,  386. 

Oak,  117,  130,  134,  135, 
161,  164,  166,  346, 
347,  348. 

Oat,  113,  151,  292,  294, 
296. 

Oat-grain,  290. 

Oat-grass,  296. 

Obdiplostemonous,  336. 

Obelidium,  103. 


Obligate  parasite e,  85. 
Ocbna,  439. 
Ocbnaceae,  439. 
Ochroma,  427. 
Ocimum.  541. 
Ocrea,  359. 
Odonthalia,  83. 
Odontites,  526. 
(Edogoniaceee,  47.  55. 
lEdogonium,  10,  ll,55,-r>(>. 
(Enantbe,  495,  498. 
CEnothera,  434.  485,  486. 
(Enotheracese,  484. 
Oidia,  90. 
Oidium,  121,  179. 
Oidium  forms,  179. 
Oil-mould,  99. 
Oil-palm,  301. 
Olea,  547. 

Oleacese,  541,  542,  546. 
Oleander,  544. 
Oligorus,  166. 
Olive,  547. 

Olive-brown  Seaweeds,  68. 
Olive  Oil,  547. 
Olpidiaceee,  103. 
Olpidieae,  103. 
Olpidium,  103. 
Olyreaa,  296. 
Orapbalodes,  533,  534, 
Onagracese,  484. 
Oncidium,  332. 
Oncobyrsa,  24. 
Onion,  312. 
Onobrycbis,  472,  473. 
Ononis,  471. 
Onopordon,  570. 
Ooblasteraa-nlaments,  82. 
Oocystis,  51. 

Oogamous  fertilisation,  13, 
Oogonium,  13. 
Oomycetes,  95,  96,  100. 
Oopbyte,  181. 
Oosphere,  13,  248. 
Oospore,  14. 
Operculum,  193. 
Opbiocytinm,  51. 
Ophioglossacere,  209,  210. 
Ophioglossurn,    210,    211r 

238. 

Ophiopogon.  320. 
Opbrydeee,  331. 
Ophrys,  332,  333. 
Opium-poppy,  395. 
Oplismenus,  295. 
Opuntia,  375,  377. 
Orange,  438. 
Orcbid,  diagram  of  flower, 

329. 

Orchidaceae,  5,  23S,  328. 
Orchideae,  277. 


INDEX. 


611 


Orchids,  151. 

Palm-wine,  301. 

Orchis,  276,  331,  332,  333. 

Palm®,  297. 

Oreobolus,  285. 

Palmella-stage,  15,  16. 

Oreodoxa,  301. 

Palmyra-palm,  301. 

Organs  of  attachment,  4. 

Paludella,  197. 

Origanum,  536,  539,  541. 

Pampas-grass,  296. 

Ornitbogalum,  312,  314. 

Panama  hats,  302. 

Ornithopus,  466,  472. 

Panax,  491. 

Orobauche,  334,  528,  529. 

Pancratium,  317. 

Orontieae,  303. 

Pandanaceaa,  302. 

Oroutium,  304. 

Pandanus,  302. 

Orris-root,  321. 

Pandorina,  45,  48. 

Orseille,  142. 

Panicete,  295. 

Orthoploceee.  400. 

Panicum,  295,  296. 

Orthospermeee,  493. 

Pansy,  411. 

Orthothecium,  197. 

Panus,  171. 

Orthotrichurn,  197. 

Papaveraceee,  394. 

Orthotropous,  242,  213. 

Papaver,  394,  395. 

Oryza,  293. 

Papaw,  476. 

Oryzeee,  293. 

Papayacese,  476. 

Osciliaria,  10,  23,  24,   26, 

Paper-mulberry  tree,  354, 

37. 

356. 

OscillariacesB,  24. 

Papilionacea?,  335,  468. 

Osiers,  152. 

Pappus,  564,  566. 

Osmunda,  209,  215. 

Papyrus,  287. 

Osmundaceee,     202,    210, 
215. 

Paradise  apple,  465. 
Paraglobulin,  473. 

Ostioles,  73. 

Paraphyses,  88. 

Ostropa,  133. 

Paraguay  tea,  445. 

Ostropaceee,  133. 

Parasites,  5. 

Ostrya,  345. 

Parasites,  endophytic,  85. 

Osyris,  500. 

endozoic,  85. 

Oura'tea,  439. 

epiphytic,  85. 

Ouvirandra,  281. 

epizoic,  85. 

Ovary,  3,  239,  250. 

facultative,  84. 

Ovule,  241,  242,  248. 

obligate,  85. 

Ovuliferous  scale,  256,  257. 

pathogenic,  85. 

Oxalidaceee,  416. 

Parasitic  Bacteria,  38. 

Oxalis,  416. 

Parasol-fungus,  171. 

Ox-eye,  572. 

Pariana,  291. 

Oxslip,  513. 

Parietaria,  353. 

Oxybaphus,  374. 

Paris,  309,  314,  316. 

Oxycoccus,  509,  510. 

Paritium.  430. 

Oxyria,  360. 

Parkia,  475. 

Oyster  Mushroom,  171. 

Parmelia,    140,    141,    142, 

143. 

Padina,  76. 

Parnassia,  453. 

Paaonia,  379,  381. 

Paronychia,  365,  367. 

Pseonieas,  381. 

Paronychieffi,  366. 

Paepaianthus,  309. 

Parrotia,  455. 

Palaquium,  511. 

Parsley,  494,  498. 

Palava,  429. 

Parsnip,  492,  49d,  493. 

Palere,  209. 

Parthenogenesis.  14. 

Pales,  288. 

Pasanea,  346  348. 

Palisander-wood.  529. 

Paspalum,  295. 

Paliurus,  448. 

Pasta  guarana,  441. 

Palm,  275,  276,  297. 

Pastinaca,  493,  496. 

Branching  of,  298. 

Passerina,  449. 

Inflorescence  of,  299. 

Passiflora,  475,  476. 

Palm-oil,  301. 

Piiss-ifloracese,  476. 

Palm-wax,  301. 

Passiflorinas,  475. 

Passion-flower,  476. 
Patellaria,  134. 
Patellariaceee,  134. 
Patellea,  134. 
Paternoster  peas,  470. 
Pathogenic   Rod-Bacteria, 

39. 

Patrinia,  557. 
Paullinia,  441. 
Paulownia,  527. 
Pavonia,  428. 
Paxillei,  172. 
Payena,  511. 
Paypayroleae,  411. 
Pea,  470. 

Peach,  117,  121,  461. 
Pear,  130,  464,  465. 
Pedagnuoli,  355. 
Pedaliacese,  518,  529. 
Pediastrum,  52. 
Pedicularis,  151,  526. 
Peganum,  438. 
Pelargonium,  418,  419. 
Peliosanthes,  320. 
Pellia,  191,  192. 
Pellitory,  353. 
Peltigera,  143. 
Pelvetia,  73. 
Penicillium,  122,  123. 
Penium,  43,  44. 
Pennisetum,  295. 
Penny-cress,  401. 
Penny-wort,  493. 
Pentacyclicee,  505,.  506. 
Pentadesma   414. 
Pentapera,  505. 
Pentstemon,  524,  527. 
Peplis,  47,  483. 
Pepper,  361. 
Peppermint,  541. 
Peperomia,  361,  362. 
Pepperwort,  401. 
Pere«kia,  375,  376. 
Perianth,  235. 
Pericha;tium,  192. 
Pericarp,  2i9. 
Pericallis,  574. 
Peridermiurn,     147,     118, 

153,  154,  1C5,  150. 
Peridinta,  1.  14,  17. 
Peridinin,  16. 
Peridinium,  17. 
Peridiola,  176. 
Peridinm,  88,  89.  147. 
Perigynium,  189. 
Perilla,  541. 
Periphyses,  88. 
Periplasm,  104. 
Periploca,  546. 
Perisperm,  249. 
Perisporiaceae,  122. 


612 


INDEX. 


Perisporiales,     95,      116, 

118,  119. 
Peristome,  195. 
Perithecia,  125. 
Periwinkle,  513,  544. 
Perizonium,  20. 
Pernambuco-tree,  468. 
Peronocarpic      ascocarps, 

125. 
Peronospora,     101,      104, 

105   107. 

PeronosporaceaB,  104. 
Persea,  393 
Persica,  461. 

Personals,  505,  515,  517. 
Pertusaria,  140,  142. 
Petals,  235. 

Petasites,  153,  569,  571. 
Petiveria,  372. 
Petrocelis,  84. 
Petunia,  518,  521. 
Peucedanese,  496. 
Peucedarmm.  496. 
Peyssonellia,'84. 
Peziza,  115,  135,  159. 
Pezizaceas,  135. 
Pezizales,  134. 
Phacelia,  515. 
Phacidiales,  133. 
Phacidium.  133. 
Phacotus,  48. 
Phseophyceie,  1,  14,  68. 
Phffiophyl,  68. 
Pbseosporese,  68. 
Phseothamnion,  54. 
Phagocytes,  41. 
Phajus,'  332. 
Phalaridea3,  295. 
Phalaris,  2!)o. 
Pha>]acese,  172. 
Phalloideffi,  96,  145,  172. 
Phallus,  172,  173. 
Phanerogams,      3,      234, 

236,  249. 
Pharbitis,  516. 
Pharus,  291,  293. 
Phascum,  195. 
Phaseoleae,  470. 
Phaseolus,   134,  469,  471, 

473. 

Phegopteris,  213,  214. 
Phellodendron,  437. 
Philadephu*,  451,  455. 
Phillyrea.  547. 
Philodendron,  303,  305. 
Philonotis,  197. 
Phlebia,  163. 
Phleum,  290,  294,  296. 
Phloem,  251. 
Phloeospora,  70. 
Phlomis,  538,  541. 


Phlox,  515, 

Phoeniceaa,  299. 

Phoenix,    298,    299,    301, 

302. 

Pholiota,  171. 
Phormium,  312;  313,  314. 
Phragmidium,    146,    147, 

148,  151,  152. 
Phragmites,  113, 131,  291, 

294. 

Phragmonema,  22,  25. 
Phrynium,  327. 
Phycocyan,  22,  77. 
Phycoerythrin,  22,  77. 
Phycomyces,  99. 
Phycomycetes,    1,    5,    95, 

96. 

Phycophffiin,  69. 
Phycopyrrin,  16. 
Phycoxanthin,  69. 
Phylica,  448. 
Phyllachora,  131. 
Pbyllactinia,  122. 
Phyllactis,  560. 
Phyllanthus,  431,  432. 
Phyllitis,  70. 
Phyllobium,  47,  51. 
Phyllocactus,  377. 
Phyllocladus.  260. 
Phyllodia,  474. 
Phyllodoce,  509. 
Phylloglossum,  228. 
Phyllophora.  83. 
Phyllosiphou,  8. 
PhyllosiphonaceaB,  47,  61. 
Physalis.  521. 
Pbysarutn,  6,  8. 
Physcia.  139,  143. 
Physcomitrium,  188,  197- 
Physiological  varieties,  41. 
Physoderma,  103. 
Physostigma,  471,  473. 
Phytelepbantinae,  301. 
Phytelephas,     299,     301, 

302. 

Phyteurna,  562. 
PhytoamcebaB,  10,  61. 
Phytolacca,  372. 
Phytolaccacese,  372. 
Phytomyxa,  b. 
Phytophtbora,     101,    104, 

105,  106. 
Piassava,  297. 
Picea.  124,  129,  132,  155, 

165,  265. 
Pichuriin,  392. 
Picraena,  438. 
Picris.  571. 
Picrotoxine,  390. 
Pilacracese,  157. 
Pilacre,  157. 


Pilea,  353. 
Pilobolus,  99,  100. 
Pilostyles.  504. 
Pilularia,  216,  220. 
Pimelea,  449 
Pimenta,  489. 
Pimento,  489. 
Pimpernel,  513. 
Pimpinell,  498. 
Pimpinella,  494,  498. 
Pine,    127,  153,  161,  165, 

255,  263,  266. 
Pine-apple,  320. 
Pine-shoot  Fuugus,  152. 
Pinellia,  305. 
Pinguicula,  334,  528. 
Pink,  367. 
Pin-mould,  99. 
Pinnularia,  19. 
Pinus,  129,   132,  153,  155, 

165,    264,    265,    266, 

267,  272. 
Pinoideffi.   256,  258,   259, 

262. 

Pipe-flower,  500. 
Piper,  361,  363. 
Piperaceae,  361. 
Pipere®,  361. 
Piptocephalidacere,  100. 
Piptocephalis,  100. 
Pircunia,  372. 
Pisonia,  374. 
Pistia,  306. 
Pistacia,  439. 
Pistil,  239. 
Pistillaria,  161. 
Pistillate,  236. 
Pisum,  469,  470,  473. 
Pitcairnia,  320. 
Pitcher-plant,  409. 
Pittosporacese.  451,  455. 
I  ittosporum,  455. 
Placenta,  237,  241. 
Placochromaticaa.  21. 
Plagiochila,  189,  192. 
Plagiothecium,  197. 
Plagiotropidea?,  21. 
Planera,  351. 
"  Plankton,"  15,  17,  20. 
Planogametes,  12. 
Plantago.    335,    530,    531, 

536.  559. 

Plantaginacese,  518,  530. 
Plantain,  530. 
Plasmodia,  4,  5,  7. 
Plasmodiophora,  8. 
Plasmodiophorales,  6. 
Platanacese,  455. 
Platanus,  456. 
Platanthera,  332,  333. 
Plate-cultures,  33. 


INDEX. 


613 


Platonia,  414. 
Platyceriuin,  213. 
Platycodoii,  562. 
Platystemon,  395. 
Plectonema,  24. 
Plectranthus,  541. 
Pleospora,  130. 
Pleosporaceae,  130. 
PJeurandra,  413. 
Pleuridium,  195. 
Pleurocarpi,  197. 
Pleurococcaceae,  47,  51. 
Pleurococcus,  10,  51,  138. 
Pleurorhi/ffi  400. 
Plenroloenium,  44. 
Pleurothallis,  332. 
Pleurotus,  171. 
Plocamium,  83. 
Plum,  117,  164,  461,  462. 
Plunibaginacese.  514. 
Plumbago,  514. 
Plumeria,  544. 
Plumule,  247. 
PneumatlxHlia,  267. 
Poa,  2S7,  290.  294,  296. 
"Pocket-plum,"  85. 
"  Pockets,"  117. 
Pod,  466. 
Pod-pepper,  522. 
Podalyrieae,  469. 
Podocarpeae,  260. 
Podocarpus,  2-31,  255,  261, 

272. 

Podopbylium,  390. 
Podosphaera,  120. 
Podospora,  129. 
Podostemaceas,  451,  456. 
Pogostemon,  541. 
Poinciana,  468. 
Point  Caraway,  498. 
Polanisia,  406\ 
Polemoniaceae,  509,  515. 
Polemonium,  515. 
Polianthes,  318 
Pollinarium,  332. 
Polliuia  329. 
Pollinodium,  100.  120. 
Pollen-chamber,  251. 
Pollen-grain,     240,     244, 

245. 

Pollen-sac,  235,  237,  240. 
Pollen-tube,  2i4,  249. 
Polyearpicaa,  377. 
Polycystis,  24. 
Polydinida,  16,  17,  18. 
Polyembryony,  247. 
Polygala,  442.  443. 
Polygalaceae,  442. 
Polygamous,  236. 
Polygouaceae,  239,  359. 
Polygonatum,  314,  316. 


Polygoniflorse,  358. 
Polygonum,  359,  360,  361. 
Polyides,  84. 
Polykrikos,  17. 
Polypetalee,  336. 
Polyphagus.  103,  104. 
Polypodiacese,     202.    205, 

206,  209,  210,  212. 
Polypodium,  207,  213. 
Poiyporaceaa,  163. 
Polyporus,  163,  164,  165. 
Polysipbonia,  79,  83. 
Polystachya,  332. 
Polystipma,  125,  127. 
Polytrichaceae,  197. 
Polytrichum,  197. 
Pomace®.  456,  462. 
Pomaderris,  448. 
Pomalo,  438. 
Pomegranate,  488,  489. 
Pomona-fungus,  171. 
Pond-weed,  279. 
Pontederia,  316. 
Pontederiaceffi,  308,  316. 
Poplar,  124,  164,  338. 
Poppies,  394. 
Populus,  152,  338. 
Pore-fungus,  163. 
Porogames,  273. 
Porouia,  131. 
Porphyra,  10,  78. 
Porphyraceae,  78. 
Portulaca,  373. 
Portulacaceae,  373. 
Posidonia,  281. 
Potamogeton,  47,  278,  279. 
Potamogetonaceaa,       278, 

279. 

Potato-fungus   104,  107. 
Potato-plant,  521,  522. 
Potentilla,  458,  460. 
Potentilleae,  458. 
Poterium,  460. 
Pothos,  304. 
Pottia,  196. 
Pottiacese,  196. 
Pouzolzia,  353. 
Prasiola,  53 
Preissia,  191. 
Preslia,  539. 
Primrose,  512. 
Primula,  511,  512,  513. 
Primulacea3,239,512,  514 
Primulinaa,  505,  511. 
Pringsheimia,  54. 
Prionium,  284. 
Pritcbardia,  ^98. 
Priva,  535. 
Privet,  547. 
Procarpiuin,  81. 
Proembryo,  64. 


I  Profichi,  355. 
Promyceliuui ,  94,  146. 
Pronucleus,  245. 
Prorocentrum,  17,  18. 
Protea,  450. 
Proteaceee,  450. 
Prothallium,  193.244,248. 

Secondary,  233. 
Protistae,  5. 
Protium,  438. 
Protobasidia,  144. 
Protobasidiomycetes,     96, 

145. 

Protococcaceae,  47,  48,  51. 
Protococcoideae,  8,  47. 
Protomyces,  108. 
Protomycetacese,  108. 
Protonema,  181. 
Provence  oil,  547. 
Prunella,  539. 
Prunus,  117, 118, 127, 130, 

152,  461,  462. 
Psalliota,    167,  168,    169, 

171. 

Psamma,  295,  296. 
Pseudopbacidiacese,  133. 
Pseudopodium,  10,  193. 
Pseudotsuga,  264, 265, 266. 
Psidium,  488,  489. 
Psilotaee83,  228. 
Psilotum,  201,  228. 
Psycbotria,  550. 
Ptelea,  437. 
Pteridium,  131,  207,  213, 

214. 

Pteridophyta,  2,  198,  234. 
Pterigynandrum,  197. 
Pteris,  199,  203,  213,214. 
Pterisanthes,  445. 
Pterocarpus,  473. 
Pterocarya,  350. 
Pterocepbalus,  560. 
Pterogoniaceaa,  197. 
Pterogyne,  468. 
Pterostegia,  360. 
Pterygophyllum,  197. 
Ptilidium,  192. 
Ptilota,  84. 
Ptychogaster,  166. 
Puccinia,  147,    148,    149, 

150. 

Puff-ball,  174. 
Pulmonaria,  533,  534. 
Pulque,  318. 
Pulsatilla,  384. 
Pumpkin,  480,  481. 
Punctaria,  70. 
Punica,483,  488,489,490. 
Puniceae,  488. 
Puschkinia,  312. 
Putrefaction,  32. 


€14 


INDEX. 


Puya,  319. 
Pycnidia,  89. 
P.ylaiella,  70. 
Pyrenoid,  46. 
Pyrenolichenes,  142A. 
Pyrenomycetes,    95,   116, 

118,  125. 
Pyrenula,  142. 
Pyrethrum,  572,  574. 
Pyrola,  334,  506,  507. 
Pyrolacese,  506. 
Pyrrophv],  16. 
Pyrus,  152,  463. 
Pythium,  101,  106. 


Quaking-grass,  294,  296. 
Quassia,  438,  439. 
Quassine,  438. 
Quercitiorffi,  337,  340. 
Quercitron- wood,  348. 
Quercus,    341,    346,   347, 

348,  504. 

Quillaja,  457,  460. 
Quillajese,  457. 
Quill-wort,  230. 
Quince,  464,  465. 
Quinchamalium,  500. 
Quinine,  550,  553. 


Racomitrium,  197. 
Radiatas,  571. 
Radicle,  247. 
Radiola,  418. 
Radiolarias,  9. 
Radish,  408.  404,  405. 
Radula,  192. 
Rafflesia,  504. 
Rafflesiaceae,  499,  504. 
Raisins,  447. 
Rajauia,  323. 
Ralfsia,  71. 
Ralfsiacese,  71. 
Ramalina,  143. 
Ramenta,  209. 
Ramie,  353. 
Rampion,  562. 
Randia,  550. 
Ranunculaceee,  278,  378. 
Ranunculese,  383. 
Ranunculus.  151, 378,  379, 

380,  382,  383,  384. 
Rapateaceae,  308. 
Rape,  404. 
Raphanus,  400,  403. 
Raphia,  301. 
Raphidium,  51. 
Raphiolepis,  463,  465. 
Raspberry,  45'J,  460,  461. 
Ravenala,  325. 


Ray-flowers,  567. 
Reboulia,  191. 
Receptacle,  210. 
Red  Algte,  1. 
Red-beet,  372. 
Red-cabbage,  405. 
Red-clover,  46^,  517. 
Red-currant,  455. 
Red-pine,  264,  266. 
"Red-rot,"  164,  166. 
Red  Sandalwood,  473. 
Red  Seaweeds,  4,  77. 
Red  Snow,  48. 
Red-strip,  165. 
Red-tree,  468. 
Reed,  151,  294. 
Reed-mace,  303. 
Reindeer  Moss,  141. 
Reineckea.  314. 
Remijia,  550,  553. 
Renealmia,326. 
Replum,  398. 
Reseda,  407. 
Resedaceee,  406. 
Resin,  266. 
Rest-harrow,  471. 
Restiacese,  309. 
Restio,  309. 
Restrepia,  332. 
Retama,  472. 
Reticularia,  8. 
Retinospora,  268. 
Rhamnaceae,  447,  449. 
hhamnus,  151,  448. 
Rhaphidophora,  305. 
Rhatany,  468. 
Rheum,  151,  359,  360. 
Rhinauthacese,  153. 
Rhinanthere,  525,  526. 
Rhinanthus,  526. 
Rhingia,  320. 
Rhipidium,  320. 
Rhipsalis,  375,  376,  377. 
Rbizidiaceae,  103. 
Rhizoboleso,  415. 
Rhizocarpfffi,  205,  215. 
Rhizoclonium,  58. 
Rhizoids,  4,  10. 
Rhizomorpha,  169.  170. 
Rhizopacete,  99. 
Rhizophora  482,  486,  487, 

513. 

Rhizophoraceee,  482,  486. 
Rhizophyllidacese,  84. 
Rhizophyllis,  84. 
Rhizopods,  5. 
Rhizopogon,  175,  176. 
Rhizopus,  99. 
Rhizosolenia,  20. 
Rhodanthe,  573. 
Rhodiola,  451,  452. 


Rhodochiton,  525. 
Rhododendron,  161,  508. 
Rhodomela,  83. 
Rhodoraelacese,  83 
Rhodophycese,  1,  14,  77. 
Rhodophyll,  77. 
Rhodophyllidaceee,  83. 
Rhodophyllis,  83. 
Rhodoraceee,  335,  508. 
Rhodotypus,  457. 
Rhodymenia,  83,  84. 
Rhodymeniacese,  83. 
Rhodymeniales,  82,  84. 
Rhoeadina3,  393. 
Rhopographus,  131. 
Rhubarb,  359. 
Rhus,  439. 
Rhynchosia,  471. 
Rhynchospora,  285,  286. 
Rhytisma,  132. 
Ribbon-grass,  296. 
Ribes,  121,   152,  153,  241, 

454,  455. 
Ribesiacese,  454. 
Rib-grass,  530. 
Riccia,  186,  189,  190. 
Ricciacese,  190. 
Rice,  291,  293,  296. 
Richardia,  305. 
Richardsonia,  550. 
Ricinus,  431. 
Riella,  192,  231. 
Ringworm,  180. 
Rivina,  372. 
Rivularia,  10,  25. 
Rivn!ariacea3,  22,  24,  25. 
Robinia,  470,  473. 
Roccella,  142. 
Rock-cress,  402. 
Rock-rose,  412. 
Rod-bacteria,  39. 
Roestelia,   147,   148,  151, 

152, 153. 

Roman  spinach,  372. 
Roots,  4. 
"  Ropiness,"  35. 
Rosa,  148,  459,  460. 
Rosaceaa  451,  457. 
Rose,  121. 
Rose-mallow,  428. 
Rose  of  Jericho,  401,  574. 
Rosese,  459. 
Rosellinia,  130. 
Rosemary,  540. 

Oil  of,  541. 
Rosiflorse,  456,  466. 
Rosmarinus,  536,  540,541. 
Rostellum,  329,  332. 
Rotang.  298. 
Royal-fern,  209,  215. 
Rubeffi,  458. 


INDEX. 


615 


Eubia,  551,  552,  553. 
Rubiacea^.  542,  546,   548, 

549,  553. 
Rubiales,    490,   505,  548, 

556,  564. 

Rubus,  458,  460,  461. 
Rndbeckia,  572. 
Ruellia,  530. 
Rulingia,  422. 
Rumex,  151,  359,  360. 
Ruppia,  278,  279. 
Ruscus,  316. 
Rush,  283,  284. 
Russula,  171. 
Russulei,  171. 
Rust  of  Wheat,  148. 
Rusts,  146. 
"Rust  spots,"  130. 
Ruta,  436. 
Rutacese,  436. 
Ruteffi,  436. 
Rye,  125,  151. 
Rye-grass,  295,  296. 
Rye-stem  blight,  113. 

Sabal,  300. 
Sabaleae,  299. 
Saccharomyces,  177,  178. 
Saccharomyces-forms,176. 
Saccharum,  293. 
Safflower,  574. 
Saffron,  321. 
Sagina,  364,  365,  366, 
Sagittaria,  281,  282. 
Sago,  254. 
Sago-palm,  298. 
Sainfoin,  472,  473. 
"  Salep,"  333. 
Salicaceae.  338. 
Salicin,  339. 
Saliciflorae,  337. 
Salicornia,  369,  371,  503. 
Salicoruieae,  371. 
Salisburia,  259. 
Salix,  122.  337,  333. 
Salpiglossis,  521. 
Salsafy,  574. 
Salsola,  370.  371,  372. 
Salsolea?,  370. 
Saltpetre  formation,  35. 
Saltwort,  370. 
Salvadora,  547. 
Salvadoruceae,  542,  547. 
Salvia,  536.  540,  541. 
Salvinia,  201,    216,    217, 

218,  245. 

Salviniaceae,  210,  218. 
Sambuceae,  555,  557. 
Sambucus,  156,  553,  555. 
Samolus,  513. 
Samydacese,  476. 


Sandal-wood,  473,  500. 
Sandarack  resin,  269. 
Sand-box  tree,  432. 
Sand-star,  287. 
Sanguinaria,  395. 
Sanguisorba,  460. 
Sanicula,  493. 
Sannicle,  493. 
Sanseviera,  320. 
Santalaceae,  500. 
Santalum,  500. 
SantoHna.  572. 
Sapindaceae,  440. 
Sapindus,  441. 
Saponaria,  368. 
Saponin,  460. 
Sapotaceas,  510. 
Saproleguia.  102, 107, 103. 
Saprolegniaceae,  107. 
Saprophytes,  5. 
Sapucaia-nuts,  489. 
Saranthe,  327. 
Sarcina,  27,  28,  38. 
Sarcophyte,  504. 
Sargassum,  4,  10.  73,  75. 
Sarothamnus,  472. 
Sarracenia,  409. 
Sarraceniaceae,  408. 
Sarsaparilla,  316. 
Sassafras,  392. 
Satureia,  540,  541. 
Satureieaa,  539. 
Saurureae,  362. 
Saururus,  362. 
Saussurea,  570. 
Sauvagesieae,  411. 
Saw-wort,  570. 
Saxifraga,  161,  452. 
Saxifragaceae,  451,  452. 
Saxifrage,  452. 
Saxifraginae.  451,  555. 
Scabiosa,  55S,  559,  560. 
Scaevola,  564. 
Scale-leaves,  235. 
Scammony,  517. 
Scandicese,  495. 
Scandix,  495. 
Scapania,  192. 
Scarlet-runner,  473. 
Scenedesmus,  51. 
Scheuchzeria,  278. 
Schistostega,  196,  197. 
Schistostegacese,  197. 
Schizaea,  215. 
Schizaeacea?,  210,  215 
Scliizanthus,  521. 
Schizocarp,  492. 
Schizocarpeae,  195. 
Schizochlamys,  51. 
Schizomeris,  53. 
Schizomycetes,  26,  33. 


Schizopetaleae,  404. 
Schizopetalum,  4n2. 
Schizopliyceae,8, 10,22,26 
Schizophylluin,  171. 
Schizophyta,  1,  14, 19,  22, 

24. 

Schcenocaulon,  310. 
Schoenoxiphium,  287. 
Schcenus,  286. 
Schwendenerian    Theory, 

139. 

Sciadium,  51. 
Sciadopitys,  267. 
Scilla,  312,  314. 
Scirpeae,  2*5. 
Scirpus,  285,  286,  287. 
Scirrhia,  131. 
Scitamineae,  276,  278,  323, 

328. 
Scitonemaceae,  22,  24,  25, 

27. 

Scleranthus,  365,  367. 
Scleria,  286. 
Scleroderma,  175. 
Sclerodermataceae,  175. 
Sclerotinia,  116,  134,  135. 
Sclerotium,  87,  127. 
Scolopendrium,  213,  214. 
Scoparia,  525. 
Scopolia,  519,  521. 
Scorodosma,  496. 
Scorzonera,  571,  574. 
Scotch  Fir,  259,  2(iti. 
Scotinosphaera,  47,  51. 
Screw  Pine,  302 
Scrophularia,  524,  526. 
Scrophulariaceae,  518,  521, 

522,  527. 

Scutellaria,  536,  539. 
Scutellum,  293. 
Scyballium,  504. 
Scytonema,  22,  26,  176. 
Scytonemac^ae,  25. 
Sea-holly,  493. 
Sea-kale,  403,  405. 
Sea-lavender,  514. 
Sea-milkwort,  513. 
Seaweed,  4. 
Sea-wormwood,  574. 
Sebacina,  156. 
Secale,  127,  295,  236. 

cornutum,  127. 
Sechium,  481. 
"  Sedimentary-yeast, "178. 
Sedum,  451,  452. 
Seed,  247,  248,  249. 
Seguieria,  372. 
Selaginaceae,  532,  541 
Selaginella,  200,  203,  223, 

229,    230,    2J2,    233, 

245,  254. 


G16 


INDEX. 


Selaginellacese,  231. 
SeJaginellese,  205,  228. 
Selago,  541. 
Selenastrum,  51. 
Selenipedilum,  329,  330. 
Seligeria,  196. 
Seligeriaceee,  196. 
Semele,  316. 
Sempervivum,    151,    451, 

452. 

Senebiera,  400,  401. 
Senecio,    566,     569,    572, 

573. 

Senecionese,  572. 
Senna,  468. 
Sepals,  235. 
Sequoia,  267,  272. 
Serapias,  332. 
Serjania,  441. 
Serratula,  570,  574. 
Serum,  33. 
Service-tree,  465. 
Sesamum,  529.  ' 
Seseli,  495. 
Seselineae,  495. 
Sesleria,  294. 
Sesuvium,  375. 
Seta,  186 
Setaria,  295. 
Sexual  reproduction,  11. 
Sheep-seaweed,  84. 
Shellac,  356,  434. 
Shepherdia,  450. 
Shepherd's-needle,  495. 
Shepherd's-purse,  401. 
Sherardia,  552. 
"Sichel,"  284. 
"  Sickle,"  284. 
Sicyos,  481. 
Sida,  428,  430. 
Sideritis,  538. 
Sideroxylou,  511. 
Sigillari'aceae,  233. 
Silaus,  495. 
Sileneaa,  367. 
Silene,  367. 
Siler,  495. 
Siliceous  earth,  20. 
Siliculosse     angustisept®, 

401. 

latiseptaj,  400. 
Siliqua,  398. 
Siliquosffi,  402 
Silk-cotton,  427. 
Silphium,  498,  572. 
Silver-leaf,  450 
Silybum,  567  570. 
Simaba,  439. 
Simaruba,  439. 
Simarubaceaa,  438. 
Sinapese,  404. 


Sinapis,  400,  40^. 
Siphocampylos,  563. 
Siphoneae,  4,  9,  47,  59. 
Siphonia,  434. 
Siphonocladus,  62. 
Sirosiphoniacese,    22,    24, 

26. 

Sisal  hemp,  318. 
Sisymbriinse,  404. 
Sisymbrium,     399,     402, 

410. 

Sisyrinchium,  321. 
Sium,  494,  498. 
Skimmia,  437. 
Skull-cap,  539. 
Slime-fungi,  1,  4,  5. 
Sloe,  461,  462. 
Sloth,  8,  356. 
Smilaceee,  316. 
Smilacina,  314. 
Smilax,  315,  316. 
"  Smut,"  113,  130. 
Smut-fungi,  114. 
Smike  cucumber,  481. 
Snapdragon,  523,  524. 
Snowberry,  554. 
Snowdrop,  317. 
Soapwort,  368. 
Soft-grass,  296. 
Soja,  471. 
Solanaceae,  514,  518,  520, 

522. 

Solaniue,  522. 
Solanum,  238,  521,  522. 
Soldanella.  513. 
Solenia,  162. 
Solidago,  573. 
Sollya,  455. 
Solomon's  seal,  314. 
Sonchus,  153,  571. 
.Sophora,  469. 
Sophoreas,  469. 
Sorbus,  152,  465. 
Sordaria,  129. 
SordariaceaB,  12d. 
Soredia,  141. 
Sorghum,  296. 
Sori,  205. 
Sorocea.  356. 
Sorrel,  361. 
Southernwood.  574. 
Sow-thistlp,  571. 
Spadiciflone,  277,  297. 
Spadix,  297. 
Sparassis,K)l. 
Sparaxis,  321. 
Spargauium,  302,  303. 
Sparmannia,  425. 
Spartium,  473. 
Spathe,  297. 
Spathicarpa,  306. 


Spathulea,  137. 
Spearmint,  541. 
Specularia,  562. 
Speedwell,  525. 
Spergula,  366. 
Spergularia,  366. 
Spermacoce,  5-50. 
Spermacoceffi,  550. 
Spermagonia,  116. 
Spermaphyta,  3. 
Spermatahgia,  81. 
Spermatia,     13,     76, 

111,146. 

Spermatochnaceee,  71. 
Sperrnatochnus.  71. 
Spermatozoid,  13,  183. 
Sperm-nucleus,  245. 
Spermocarp,  58. 
Spermogonia,      89,      141, 

146,  147,  149. 
SpermothamnioD,  84. 
Sphacelaria,  70. 
Sphacelariaceee,  70. 
iSpbacelia,  125,  126. 
Sphacelotheca.  110. 
Sphseralcea,  430. 
Sphau-ella,  48,  130, 
Spbserellacese,  130. 
Sphasriales,  129. 
Sphaerobolacese,  173. 
Sphasrobolus,  173. 
Spheerocarpus,  191,  192. 
Sphaerococcaceas,  83. 
Spbserophorus,  142. 
Sphseroplea,  13,  14,  58. 
Spheeropleaceas,  47,  58. 
Spheerotheca,  120,  121. 
Spheerozosma,  44. 
Sphagneee,  193 
Sphagnum,  186,  188,  192, 

194,  195,  197. 
Sphenogyne,  5b6. 
Sphenophyllacese,  233. 
Sphinctrina,  140. 
Spigelia,  546. 
S pikelet,  285,  287,  289. 
Spikes,  285. 
Spilanthes,  572. 
Spinach,  371. 
Spinacia,  371,  372. 
Spindle-tree,  444. 
Spirffia,    151,     451,     456. 

457,  4^0. 
Spiraeeje,  457- 
Spiranthes,  331. 
Spirilla,  27. 
Spirillum,  30. 
Spirochffitse,  27. 
Spirochajte,  28,  38,  40. 
Spirodela,  307. 
Spirogyra,  44,  45. 


INDEX. 


617 


Spirolobeaa,  371,  400. 
Spirotseuia,  44, 
Spiruliua,  24. 
Splacbnacese,  197. 
Splachuum,  196,  197. 
Spondias,  439. 
Sponges,  8. 
Sporangia,  239. 
Sporangial-layers,  88. 
Sporangiocarp,  88. 
Sporangio  -  fructification, 

87. 

Sporangiophore,  88. 
Spores,  10. 

Liberation   and    Distri- 
bution of,  91. 

Germination  of,  93. 
Sporidia,  112. 
Sporobolus,  295. 
Sporocarp,  205,  219. 
Sporochnaceae,  71. 
Hporochnus,  71. 
Sporogonium,  186. 
Sporophylls,  223,  235,  236. 
Sporophyte,  181,  186. 
Spring-spores,  147. 
Spumaria,  8. 
Spurge,  431,  432. 
Spurge-laurel,  449. 
Spnrry,  366. 
Squatii ariacee,  84. 
"Squills,"  314. 
Squirting  cucumber,  480. 
Stachydeaa,  538. 
Stacbys,  538,  541. 
Stachytarpbeta,  535. 
Staehelma,  570. 
Stag-truffle,  124. 
Stalk,  186. 
Stamen,  235,  236. 
Stamiuate,  286. 
Stangeria,  253,  254. 
Stanhopea,  332. 
Stanleyinae,  404. 
Stapelia,  546. 
Stapbylea,  440. 
Stapbyleacea3,  440. 
Stapbylococcus,  39. 
Star-aniseed,  389. 
Statice,  514. 
Staurastrum,  42,  43,  44. 
Steenbammera,  533. 
Stegocarpeee,  195. 
Stellaria,  364,  365,  366. 
Stella  ta3,  550,  552,  553. 
Stemless  Plants,  1. 
Stemonitis,  7,  8. 
Stepbanospermum,  272. 
Stephanosphaara,  48. 
Sterculia,  422. 
Sterculiacese,  422. 


Stereocaulon,  143. 
Stereum,  162. 
Sterigmata,  144,  146. 
Sterilization,  32. 
Sticta,  134,  137,  143. 
Stictidacese,  133. 
Stictidales,  133. 
Stictis,  133. 
Stigeoclonium,  54. 
Stigma,  3,  250. 
Stigmaria,  233. 
Stigonem*,  26,  142. 
Stilbaceas,  532,  541. 
Stilbe,  541. 
Stillingia,  434. 
Stilopbora,  71. 
Stilophoraceee,  71. 
Stinkbrand,  113. 
Stink-born,  172,  173, 
Stipa,  291,  294,  296. 
Stitcbwort,  366. 
St.  John's-wort,  413. 
Stock,  402,  405. 

Stonebrand,  113. 

Stonecrop,  451. 

Stone-wort,  1,  14. 

Stork's-bill,  419. 

Stratiotes,  282. 

Strawberry,  458. 

Strawberry-tree,  508. 

Strelitzia,  325. 

Streptocarpus,  528. 

Streptocbaeta,  290. 

Streptococcus,  39. 

Streptopus,  314. 

Striaria,  70. 

Striariacese,  70. 

Strickeria,  129,  130. 

Stroma,  88. 

Stromantbe,  327. 

Strophanthus,  544. 

Struthiopteris,   209,    214, 
254. 

Struvea,  9,  62. 

Strychnine,  546. 

Strychnos,  546. 

Sturmia,  332. 

Stylar-column,  328 
-brusb,  567. 

Style,  250. 

Stylidiacea?,  564. 

Stylidfum,  564. 

Stylochrysalis,  15. 

Stylopod,  492. 

Styphelia,  509. 

Styracaceaa,  511. 

Styrax,  511. 

Styrax-balsam,  455. 

Subhymenial  layer,  167. 

Subularia,  393,   39iJ,   400, 
401. 


Succisa,  517. 
Sugar-beet,  372. 
Sugar-cane,  239,  293,  296. 
Sugar-root,  498. 
Sulpbur-bacteria,  37,  38. 
Sumach,  439. 
Summer-spores,  147. 
Sundew,  407. 
Sun  -flower,  572. 
Sunn  hemp,  473. 
"  Surface  yeast,"  178. 
Surirayeae,  21. 
Suspensor,  233,  240,  247. 
Swamp  cypress,  267. 
Swarmspores,  10,  87. 
Swede,  405. 
Sweet  Cicely.  498. 
Sweet-flag,  303. 
Sweet-gale,  351. 
Sweet  oil,  547- 
Sweet-pea,  470. 
Sweet-potato,  517. 
Sweet-vernal,  295,  296. 
Swertia,  542. 
Swietenia,  436. 
Swine's-succory,  571. 
Sycamore,  133,  442. 
Symbiosis,  85. 
Sympetalas,  336.  50.4. 
Symphoricarpus,  554,  556. 
Sympbyandra,  562. 
Symphyllodium,  257. 
Syrnphytopleura,  387. 
Symphytum,  533,  535. 
Symploca,  24. 
Synalissa,  139. 
Syuandrium,  306. 
Synangium,  212. 
Syncarp,  278. 
Syncepbalis,  100. 
Synchytrieas,  103. 
Synchytriuin,  103. 
Syncrypta.  15. 
Synedra,  21. 
S.ynergidaa,  248. 
Syngeneticae,  1,  14,  15, 17, 

4S. 

Syngouium,  3^06. 
Synura,  15. 

Syringa,455,54G,547,550. 
Systegium,  196. 
Systematic  division  of  the 
A'gse,  14. 

of  Filices,  210. 

of  Fungi,  95. 

of  Monocotyledons,  277. 

of  Thallophytes,  4. 

of      Vascular      Crypto- 
gams, 204. 

Tabellaria,  19. 


618 


INDEX. 


Tabellarieee,  21. 
Tabernsemontana,  544. 
Taccarum,  306. 
Tacona,  284. 
Tagetes,  564,  572. 
Takamahaka,  438. 
Talinum,  373. 
Talipot,  298. 
Tallow-tree,  434. 
Tamaricaceee,  411. 
Tamarind,  466,  468. 
Tatnarindus,  467. 
Tamarisk,  411. 
Tamarix,  411,  412. 
Tamus,  323. 
Tanacetum,  572,  574. 
Tanghinia,  544. 
Tannin,  490. 
Tansy,  572. 

Tapetum,  203,  239,  240. 
Taj.hrina,  116,  117,  118. 
Taphrinacese,  116. 
Tapioca,  434. 
Tar,  266. 

Taraxacum,  571,  566,  574. 
Targiouia,  191. 
Tassel  Pond-weed,  279. 
Taxaceae,  259,  272. 
Tuxes,  261. 

Taxodiace8B,257,  267,272. 
Taxodium,  267. 
Taxoidero.  258,  259. 
Taxus,  237,  238,  255,  257, 

259,  261,  262,  272. 
Tea,  415. 

Tea-plant,  False,  521. 
Tea-rose,  460. 
Teak-tree,  535. 
Tear-Fuugus,  166. 
Teasel,  494,  558,  560. 
Tecoma,  529. 
Tectona,  535. 
Teesdalia,  398,  401. 
Telegraph-plant,  466. 
Teleutospores,  146. 
Tellima,  452. 
Terebinthinse,  435. 
Terfezia,  124. 
Terminalia,  437. 
Ternstroemiaceae,  414. 
Testa,  247,  248. 
Testudinaria,  323. 
Tetmemorus,  44. 
Tetracyclicee,  505,  514. 
Tetradynamia,  398. 
Tetragonia,  375. 
Tetragonolobus,  471. 
Tetraphis,  195.  196,  197. 
Tetrapoma,  400. 
Tetrapteris,  442. 
Tetraspora,  51. 


Tetrasporacese,  47, 48,  51. 
Tetraspores,  10,  76. 
Teucrium,  567. 
Thalassia.  283. 
Thalia,  327. 
Thalictrum,  379,  385. 
Thallophyta,  1,  4. 
Thallus,  1,  4. 
Thamnidiacese,  99. 
Thamnidium,  100. 
Thea,  414,  415. 
Thecaphora,  110,  114. 
Them,  374. 
Thelebolaceee,  109. 
Thelebolus,  109  120. 
Thelephora,  162,  176. 
Thelephoraceae,  162. 
Thelygonum,  372. 
Thelypodieae,  404. 
Theobroma,  422,  423. 
Theobromine,  423. 
Theophrasta,  513. 
Thesium,  500. 
Thistle,  569. 
Thladiantha,  481. 
Thlaspi,  400,  401,  402. 
Thomasia,  422. 
Thorn-apple,  520. 
Thottea,  499. 
Thrift,  514. 
Thrinax,  300. 
Thrush,  180. 
Thuidium,  197. 
Thuja,  241,  268. 
Thujopsis,  269. 
Thunbergia,  530. 
Thyme,  539,  541. 
Tbymelaaa,  449 
Thymel«aceag,  449. 
Thymelffiinae,  448. 
Thymus,  537,  539,  541. 
Tiaridium,  533. 
Tibouchina,  484. 
Ticorea,  437. 
Tigridia,  321. 
Tilia,  424,  425. 
Tiliacea?,  423. 
Tillandsia,  320. 
Tilletia,  111,  112,  113. 
Tilletiaceae,  111),  113. 
Tiiopteridace*,  72. 
Tilopteris,  72. 
Timothy-grass,  294,  296. 
Tinnantia,  308. 
Tmesipteris,  228. 
Toad-flax,  525. 
Toad-rush,  284. 
Toadstools,  159,  166. 
Tobacco,  520,  529. 

Virginian.  522. 
Toddaliese,  437. 


Todea,  203. 
Tofieldia,  310. 
Tofieldiese,  310. 
Tolu,  Balsam  of,  473. 
Toluifera,  473. 
Tolypella,  67. 
Tol'ypellopsis,  67. 
Tolyposporium,  110. 
Tolypothrix,  26. 
Tomato,  521. 
Tomentella,  161. 
Tomentellaceee,  161. 
Tonquin-bean.  466,  472. 
Tooth-wort,  526. 
Tordylium,  496. 
Torenia,  525. 
Torilis,  497. 
Torreya,  262,  272. 
Touchwood,  164. 
Tournefortia,  533. 
Trabeculaa,  241. 
Tracheides.  251. 
Trachylobium,  468. 
Tradescantia,  308. 
Trama,  167,  174. 
Trametes.  164,  165. 
Tragacanth,  Gum,  473 
Tragopogon,  113, 564,  5' 

574. 

Trapa,  485,  486. 
Travellers'  Palm,  325. 
Tremandraoeee,  412. 
Tremella,  156,  157,  159. 
Tremellaceaa,  146,  156. 
Trentepohlia,  8,  51. 
Tribulus,  438. 
Trichia,  8. 

Trichocoma,  176. 

Tricbodesmium,  22. 

Trichogyne,  58.  81. 

Tricholoma,  168,  171. 

Trichomanes,  206,  215. 

Trichophilus,  8,  54. 

Triohosantbes,  48 1. 

Trichosphseria,  l'2(},  130. 

Trichospheeriaceae,  129. 

Trichostomum,  196. 

Tricoccee,  430. 

Tricyrtis,  310. 

Trientalis,  512,  513. 

Trifolieas,  471. 

Trifolium,  469,  471,  473 

Triglochin,  278  279. 

Trigoniacese,  442. 

Trillium,  314. 

Triodia,  294. 

Triphasia,  438. 

Triphragmium,  147,  151 

Triplaris,  361. 

Triteleia,  312. 

Triticum,  288,  295,  29.6. 


INDEX. 


610 


Tritonia,  321. 

Urena,  428. 

Triumfetta,  424.  425. 

Ureneae,  428. 

Trollius,  379,  381. 

Urginea,  312,  314. 

Tropeeolaceae,  419. 

Urocystis,  113. 

Tropseolum,  420. 

Uroglena,  15. 

True  Ferns,  204,  205. 

Uromyces,  148,  151. 

True  Laurels,  391. 

Urophlyctis,  103. 

True  Mosses,  192. 

Urospora.  58. 

Truffles,  124. 

Urtica,  134,  151,  351,  353. 

Trumpet-tree,  356. 

Urticacese,  352. 

Trumpet-wood,  529. 

Urticiflorae,  351. 

Tryblidiaceaa,  133. 

Usnea,  143. 

Tryblidiales,  133. 

Ustilaginaceas,  110,  113. 

Tryblidium,  133. 

Ustilaginea3,  109. 

Tsuga,  265,  266. 

Ustilago,  111,  113. 

Tuber,  124. 

Ustulina,  131. 

Tuberaceee,  124. 

Utricularia,  527,  528. 

Tubercles,  8,  466. 

Utriculariacese,  518,  527. 

Tubercularia,  127. 

Utriculus,  287. 

Tuberose,  318. 

Uvularia,  310. 

Tubiflorje,  505,  514,  532. 

Tuburcinia,  110,  111,  113. 

Vaccines,  41. 

Tulip,  312. 

Yacciniacea3,451,508,  509. 

Tulipa,  312,  314. 

Yaccinium,  134,  160,  161, 

Tulipese,  312. 

509,  510. 

Tupa,  563. 

Vaginula,  189. 

Turkish-millet,  296. 

Vahea,  544. 

Turmeric,  326. 

Vaillantia,  552. 

Turneracese,  476. 

Valeriana,  557,  558. 

Turnip,  405. 

Valerianaceee,  549,  556. 

Turpentine,  266,  439. 

Valerianella,  557,  558. 

Turritinas,  404. 

Vallisneria,  282,  283. 

Tussilago,    151,  569,  571, 

Valloons,  348. 

574. 

Vallota,  318. 

Tydeea,  528. 

Valonia,  59.  62. 

Tylostoma,  174. 

Valoniacese,  47,  62. 

Ty]ostomacea3,  174. 

Valsa,  130. 

Typha,  30-'.  303. 

Valsacese,  130. 

Tvphaceae,  302. 

Vanda,  332. 

Typhula,  161. 

Vandellia,  525. 

Vandese,  332. 

Ulex,  472. 

Vanilla,  331,  333. 

Ullucus,  371,  372. 

Vascular  Cryptogams,    2, 

Ulmaceee,  351. 

198,  240. 

Ulmeas,  351. 

Isosporous,  200. 

Ulmus,  351. 

Heterosporous,  200. 

Ulothricaceae,  47,  53. 

Vateria,  415. 

Ulothrix,  12,  14,  53,  54. 

Vaucheria,  10,  33,  61. 

Ulva,  10,  53. 

Vaucheriacese,  47,  60. 

Ulvaceaa,  47,  53. 

Vegetable-ivory,  301,  302. 

UmbelliferfE,  491. 

Vegetable-silk,  545. 

Umbelliflorse,  490. 

Velamen,  332. 

Umbilicaria,  143. 

Vella,  400. 

Umbilicus,  451. 

Veiling,  404. 

Uncaria,  553. 

Vellosia,  318. 

Uncinia,  287. 

Vellosieaa,  318. 

Uncinula,  122. 

Velltheimia,  312. 

Upas-tree,  356. 

Velum  partiale,  167,  168. 

Urare,  546. 

uuiversale,  167.    » 

Uredinacese,  145,  146. 

Venter,  184. 

Uredo,  148. 

Ventral-canal-cell,  185. 

Venturia,  130. 
Veratreaa,  310. 
Veratriu,  311. 
Veratrum,  310,  311. 
Verbascuui,  523,  525,  527. 
Verbena,  535. 
Verbenaceaa,  532,  535,  537. 
Vernonia,  571. 
Veronica,   335,  523,   525r 

526,    527,    530,   536, 

55y. 

Verpa,  136. 
Verrucaria,  140,  142. 
Vesicaria,  400. 
Vetcb,  470. 
Vibriones,  27. 
Viburnum,  455,  553,  555r 

556. 

Vicia,  469,  470,  473. 
Vicieee,  469,  470. 
Victoria,  386,  387. 
Vigna,  471. 
Vinca,  544. 

Vincetoxicum,  155,  546. 
Vine,  121,  444. 
Vinegar -bacterium,  31,  32, 

35. 

Viola,  410,  411. 
Violaceaa,  410. 
Violets,  114,  410. 
Violet-stone,  54. 
Viper's-bugloss,  533. 
Virginian-creeper,  447. 
Viscaria,  364,  367. 
Viscoideee,  501. 
Viscum,  501,  5.02,  504. 
Vismia,  414. 
Vitex,  535. 
Vitis,  445,  446,  447. 
Vochysiacese,  442. 
Volkmannia,  225. 
Volva,  167. 
Volvaria,  171. 
Volvocaceaa,  14,  47,  48, 
Volvox,  48,  50. 
Vomic  nut,  546. 
"Vorblatt,"  275. 

Wahlenbergia,  562. 
Wallflower,  402,  405. 
Wall-lichen,  143. 
Wall-rue,  213. 
Walnut,  165,  349,  350, 
Water-cress,  402,  405. 
Water-dropwort.  4(J8. 
Water-ferns,  2u5,  215, 
Water-fungi,  96. 
Water-hyssop,  525. 
Water-lilies,  385. 
Water-melon,  481. 
Water-milfoil,  486, 


620 


INDEX. 


Water-net,  52. 
Water-purslane,  483. 
Water -so  Idler,  282. 
Water-wort,  413. 
Water-violet,  512. 
Wax-flower,  546. 
Weberia,  197. 
Weigelia,  554. 
Weingaertneria,  294. 
Weisia,  196. 
Weisiacese,  196. 
Wellingtonia,  267. 
Wehvitschia,  270,271. 
"  Wendungszellen,"  67. " 
West-Indian      arrowroot, 

327. 

Weymouth  Pine,  266,  267. 
Wheat,  113,  291,  292,  295, 

296. 

Wheat-grain,  292. 
Wheat  seedling  of,  292. 
White-beam,  465. 
White  Bryony,  481. 
White-cabbage,  405. 
White-mustard,  405. 
White  Pine,  266. 
White-pepper,  363. 
White-rot,  164,  165. 
White  Water-lily,  387. 
Whitlavia,  515. 
Whortleberry,  509. 
Wig-tree.  439. 
Wild  Basil,  540. 
Wild  Cabbage,  404. 
Willow,  124,  133,  338. 
Willow-herb,  4ti4. 
Winter-aconite,  382. 
Winter-cherry,  521. 


Winter-cress,  402. 
Winter-green,  507. 
Winter-spores,  146. 
Wistaria,  470,  473. 
Witches'-brooms,  85,  117, 

155. 

Woad,  403,  405. 
Wolffia,  307. 
Wood,  251. 
Wood-rush,  284. 
Wood-sorrel,  416. 
Woodruff,  552,  553. 
Woodsia,  214. 
Wormwood,  572,  574. 
Woundwort,  538. 

Xanthellacese,  15. 
Xanthidium,  44. 
Xanthium,  569,  573. 
Xanthorhiza,  379,  383. 
Xanthorrheea,  312. 
Xeranthemum,  566,  570. 
Xerotes,  312. 
Xylaria,  131. 
Xylariacese,  131. 
Xylem,  251. 
Xylopia,  388. 
Xylophylla,  431,  432. 
Xylosteum,  554. 
Xyridaceee,  30$. 

Yam,  323. 

Yeast-formation,  94. 
Yeast-fungi,  31,  36. 
Yellow  bird's-nest,  507. 
Yellow-rattle,  525,  526. 
Yellow  Water-lily,  387. 
Yellow-wort,  543. 


Yew,  259,  261,  C66. 
Ylang-ylang,  388. 
Yorkshire-log,  294,  296. 
Yucca,  312,  313,  316. 

Zamia,  253. 
Zaimardinia,  12,  72. 
Zannichellia,  278,  279. 
Zantedeschia,  305,  306. 
Zanthoxylese,  436. 
Zanthoxylum,  436. 
Zea,  290,  293. 
Zelkova,  351. 
Ziugiber,  326. 
Zingiberacese,    277,    323, 

325. 

Zinnia,  572. 
Zizania,  293. 
Zizyphus,  448. 
Zoochlorella,  9. 
Zoogametes,  12. 
Zoogloea,  27. 
Zoogonicffi,  68,  70. 
Zoosporangia,  10. 
Zoospores,  10,  87. 
Zooxantella,  9. 
Zostera,279,280,306,316. 
Zostereae,  278. 
Zygadenus,  310. 
Zygochytriacese,  103. 
Zygomorphy,  277. 
Zygomycetes,  95,  96. 
Zygophyllaceee,  438. 
Zygophyllum,  438. 
Zygospore,  12. 
Zygote,  12. 
Zygnema,  44,  45. 
Zygnemaceae,  44. 


Butler  &  Tanner,  The  Selwood  Printing  Works,  Frome,  and  London. 


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